3 Sade ty alt ae nae Lyte Begs Melts ete neha Terral i ae HUG | VOL. 105 JANUARY 2003 NO. 1 aed & (ISSN 0013-8797) EG i ™" PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED CONTENTS “Ib ADAMSKI, DAVID and CHRIS T. MAIER—A new Holcocera Clemens (Depron Gelechioidea: Coleophoridae: Holcocerini) associated with Pinaceae in North America .... ADLER, PETER H., CHARLES L. BROCKHOUSE, and DOUGLAS C. CURRIE—A new species of black fly (Diptera: Simuliidae) from Nova Scotia ..............22222.0eeeee eee ee ees 9 BAUMGARDNER, DAVID E.—New synonyms and stage description for three species of Bentohyphidac(Ephemeroptera) | mem cnc eciece ee eer tee reciente ieee ice eiies 203 BYERS, GEORGE W. and DOUGLAS A. ROSSMAN—A new Dactylolabis Osten Sacken (Dipterashipulidac)xtronvN orth Catolinarerde-se ac oe cis fates etesere snieeaoae each eeisl eee ans 70 CARROLL, J. F—Survival of larvae and nymphs of /vodes scapularis Say (Acari: Ixodidae) in foumbhabitatsiin Maryland. = a.8\5- 4 sonic uistisene since nicleinines cloteie encase aie incite mesa en aes 120 COSTACURTA, NISE DO CARMO, CLAUDIO JOSE BARROS DE CARVALHO and MARCIA SOUTO COURI—A new species of Plumispina Albuquerque (Diptera: Muscidae) from SouthegneBS razr ay eestor... . isos ee iaertetencts ieee lee tamer sere eab ris ails ceil aeltaathane eet 127 DARSIE, RICHARD FE, JR. and JONATHAN F. DAY—Studies of the genus Cu/ex Linnaeus in Florida. I. Redescription of the pupae of Culex nigripalpus Theobald and Cx. tarsalis Coquillett, vectors of St. Louis encephalitis, and a key to pupae of Cu/ex species in the east- ermilenited!Statesi(Diptera:(Culicidac) inner eso coscet tse beat cece cls teacide eters ate 100 HENRY, THOMAS J.—Proboscidotylus nigrosquamis (Maldonado) (Heteroptera: Miridae: Orthotylinae): New combination and first record for the United States ...................... 59 HOEBEKE, E. RICHARD and MAUREEN E. CARTER—Halyomorpha halys (Stal) (Heter- optera: Pentatomidae): A polyphagous plant pest from Asia newly detected in North America 225 HOLZENTHAL, RALPH W. and GISELE L. DE ALMEIDA—New species of Polycentropodidae (Trichoptera) from southeastern and southern Brazil .................00ecceeeesceeeseseeseeeee 22 JENSEN, PETER D., KEVIN M. O’NEILL, and MATTHEW LAVIN—Pollen provision records for three solitary bee species of Megachile Latreille and Heriades Spinola (Hymenoptera: Merachilidae) anysonthwesterm MMOMtama) miter a rrais clcletets eeeietctele emis e\=)e ota Wermisieder ste (oie iors le sp elajaleiatsl) Fig. 1. Cernotina antonina, male genitalia. A, Lateral. B, Tergum X and intermediate appendages, dorsal C, Sternum IX and inferior appendages, ventral. D, Phallus, lateral. 24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON branch of the intermediate appendage which bears a pair of short spinelike setae. Male.—Length of forewing 4.0 mm. Col- or in alcohol, pale yellowish brown. Geni- talia as in Fig. 1. Sternum IX short, pro- duced anterolaterally; anterior margin, in ventral view, narrowly excavated; tergum IX membranous, not evident. Inferior append- age elongate rectangular, truncate apically; apicoventral and mesal surfaces pigmented, setose; apicomesal lobe wide, truncate, pig- mented apically, with 4 short thick apico- mesal setae; basodorsal lobe shelflike, with long apicomesal setae and shorter thicker se- tae mesally. Tergum X membranous basally and mesally, divided mid-dorsally; lateral lobes lightly sclerotized, rounded apically, setose dorsally; base of X internally with pocket of fine setae. Preanal appendage ap- parently absent. Intermediate appendage bi- partite; dorsal branch rodlike, bearing short fine setae and pair of short thick spinelike apical and subapical setae; ventral branch shorter, digitate, setose. Phallus long, slen- der, tubular, membraneous internally, with internal sclerite and 3 short spines. Female.—Unknown. Type material.—Holotype, ¢. BRAZIL: Parana: Antonina, Reserva de Sapitandu- va, 25°28’S, 48°50'W, el. 60 m, 27.iii.1987, PROFAUPAR-lampada (UFPR). Paratypes. Same data as holotype, 26.iv.1987, 1 ¢ (UFPR):; Minas Gerais: Rio Santo Antonio, downstream from Morro do_ Pilar, I9%08-134"S; 43°21-256'W, el. 530° m, 17.x.2000, Paprocki & Ferreira, 1 6 (UMSP): Paranda: Guarapuava, Estancia Agua Santa Clara, 25°40'S, 52°01'W, el. 740 m, 1.xi1.1986, PROFAUPAR-lampada, 1 3d (UFPR). Etymology.—The species is named for the type locality. Cernotina lazzarii Holzenthal and Almeida, new species (Fig. 2) Cernotina lazzarii appears to be closest to C. abbreviata Flint 1971 and C. perpen- dicularis Flint 1971, the three species form- ing a distinct group within the genus. All share a divided tergum X with its apex bearing a sclerotized process either apically (C. abbreviata, C. perpendicularis) or api- coventrally (C. lazzarii). The dorsal branch of the preanal appendage of each species is short and bears a pair of spinelike setae, but in the new species this branch is directed posteriorly rather than posteroventrally. Male.—Length of forewing 3.5—3.7 mm. Body sclerites pale yellowish brown; anten- na mostly white, more apical flagellomeres cream colored with light brown setae basal- ly; head and thorax with white setae dorso- mesally and light brown setae laterally; legs light brown with darker brown setae; wings light brown with scattered yellowish brown setae and line of white to cream colored se- tae along anal margin from base to arculus. Genitalia as in Fig. 2. Sternum IX short, pro- duced anterolaterally; anterior margin broad- ly excavated ventrally; tergum IX membra- nous, not evident. Inferior appendage elon- gate rectangular; apex truncate, pigmented, setose apicoventrally; mesal surface with short, broad, setose lobe; apicomesal lobe narrow, acute, pigmented apically, with pair of mesal setae; basodorsal lobe shelflike, with long apicomesal setae and short thick mesal setae. Tergum X lightly sclerotized basally and mesally; in lateral view appear- ing bulbous and broadly rounded; tergum X divided middorsally, lateral lobes apically acute, slightly mesally directed, subapicoy- entrally with small sclerotized process. Pre- anal appendage short, rounded, setose. Inter- mediate appendage bipartite; dorsal branch short, rodlike, bearing pair of short apical and subapical spinelike setae; ventral branch longer, triangular, with apical and ventral se- tae. Phallus long, slender, tubular, membra- nous internally, with internal sclerite and 4 short spines. Female.—Length of forewing 3.5—4.0 mm. Color and structure similar to male. Genitalia typical for genus. Type material.—Holotype, ¢. BRAZIL: Parana: Municipio de Corbélia, Rio Novo headwaters, 24°53.886'S, 53°14.895’'W, el. VOLUME 105, NUMBER 1 to 7) Cernotina lazzarii Fig. 2. Cernotina lazzarti, male genitalia. A, Lateral. B, Tergum X, preanal and intermediate appendages, dorsal. C, Sternum IX and inferior appendages, ventral. D, Phallus, lateral. 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 700 m. 4—7.iv.1998, Holzenthal & Huisman (MZUSP). Paratypes. Same data as holo- type, 3 6,5 2 (UMSP), 1 6,5 2 (NMNH), 2 3,5 2 (MZUSP); Parand: Fénix, Reser- va Estadual ITCE 23°54’S, 51°58'W, el. 350 m, 20.xi.1987, PROFAUPAR-lampada, 1 do (UFPR). Etymology.—This species is named with gratitude and affection for Dr. Flavio Laz- zari, agronomist and plant pathologist, Cur- itiba, Brazil, on whose farm the new species was collected. Polycentropus urubici Holzenthal and Almeida, new species (Fig. 3) This species appears to be a member of the jorgenseni complex of the gertschi spe- cies group as defined by Hamilton (1986), but the dorsal band in the apical membranes of the phallus in P. wrubici is not as distinct as illustrated by Hamilton for other mem- bers of the complex. Polycentropus urubici is not close to any of the described species in the complex, but resembles a few of the undescribed species illustrated by Hamilton (1986). Male.—Length of forewing 5.5—6.0 mm. Body and wings entirely fuscous, legs dark brown. Genitalia as in Fig. 3. Sternum IX short; anterior margin rounded in lateral view, shallowly emarginate in ventral view; tergum IX membranous. Inferior appendage short, triangular, heavily setose, broad ba- sally, narrowing to acute, slightly upturned apex, with mesoventral toothlike projection; mesal surface setose (in paratypes from Santa Catarina, inferior appendage narrow- er basally and apex more strongly up- turned). Tergum X entirely membranous, broad basally, narrow apically. Intermediate appendage long, rodlike, narrowed apically, extending ventrad to apex of inferior ap- pendage (in paratypes from Santa Catarina, intermediate appendage narrow throughout length). Preanal appendage elongate oval, setose; mesoventral process short, digitate, setose, slightly upturned. Phallobase short; apicoventral processs long, thick, pointed, apex directed ventrad; phallic sclerite and dorsal band lightly sclerotized, indistinct. Subphallic sclerite broad, Y-shaped in cau- dal view. Female.—Unknown. Type material—Holotype, ¢. BRAZIL: Parana: Telémaco Borba, Reserva Samuel Klabin, 24°177S) 50°37 W;, els 750 ims 7.vill. 1986, PROFAUPAR-lampada (UFPR). Paratypes. Same data as holotype, 23.x.1987, 1 3d (UFPR), 17.1.1988, 1 3 (UFPR). Santa Catarina: Morro da Igreja, Urubici, Cach- oeira Véu da Noiva, 28°04.595’S, 49°31.090'W, el. 1,300 m, 5.iii.1998, Holzen- thal, Froehlich, Paprocki, 3 ¢ (UMSP), 2 3 (MZUSP): Urubici, Cachoeira Avencal, 28°02.839'S, 49°36.997'W, el. 1,260 m, 6.111.1998, Holzenthal, Froehlich, Paprocki, 1 3d (UMSP), 1 d (NMNH). Etymology.—The species is named for the town in Santa Catarina where paratype specimens were collected. Polyplectropus profaupar Holzenthal and Almeida, new species (Fig. 4) This species is very similar to Polyplec- tropus annulicornis Ulmer, also known from Santa Catarina, Brazil, but differs in the following ways. The dorsolateral pro- cess of the preanal appendage, which is thick and spinose in P. profaupar is long, slender and without spines in P. annulicor- nis; When viewed ventally, the ventromesal process of the inferior appendage is longer and more digitiform in P. annulicornis than in the new species and the apex of the in- ferior appendage is more rounded in P. an- nulicornis. Male.—Length of forewing 5.5—6.5 mm. Body sclerites pale yellowish brown; palps pale yellow with scattered brown setae; an- tenna pale yellow, scape with long white se- tae, antennal flagellomeres with light brown setae basally; dorsum of head with white se- tae mesally and light brown setae laterally; pronotum with white and brown setae and mesonotum with white setae: legs yellow- ish with scattered brown setae; wings light VOLUME 105, NUMBER I 27 ) 3C Polycentropus urubici Fig. 3. Polycentropus urubici, male genitalia. A, Lateral. B, Segment LX, preanal and intermediate append- ages, dorsal. C, Sternum IX and inferior appendages, ventral. D, Phallus, lateral. E, Tergum X, dorsal. F Inferior appendage, caudal. G, Inferior appendage, paratype from Santa Catarina, lateral. H, Intermediate appendage, paratype from Santa Catarina, lateral. 3B 28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4B Fig. 4. Polyplectropus profaupar, male genitalia. A, Lateral. B, Segment X, preanal and intermediate ap- pendages, dorsal. C, Sternum LX and inferior appendages, ventral. D, Phallus, lateral. E, Inferior appendages and ventral portion of preanal appendages, caudal. VOLUME 105, NUMBER 1 brown, covered with whitish setae and scat- tered patches of light brown setae, giving a mottled appearance, these patches darker at base of costa and at pterostigma. Genitalia as in Fig. 4. Sternum IX short, triangular; anterior margin straight in lateral view, shal- lowly emarginate in ventral view; tergum IX membranous. Inferior appendage with lateral lobe setose, elongate oval; ventromesal pro- cess triangular, bearing acute projections dorsally and ventrally. Tergum X entirely membranous, short. Intermediate appendage short, digitate, apically setose. Preanal ap- pendage tripartite; dorsolateral process heavily sclerotized, long, thick, recurved, with lateral spines; mesolateral process oval, setose; ventrolateral process spatulate, se- tose, with digitate mesal process bearing spinelike ventral projection. Phallus with narrow basal portion and membranous apical portion, which bears pair of sclerotized pointed processes ventrally and patch of fine spines subapically; apex of phallus acute; in- ternal phallic sclerites or spines not evident. Female.—Length of forewing 6.0—8.0 mm. Color and structure similar to male. Genitalia typical for genus. Type material—Holotype, ¢. BRAZIL: Santa Catarina: Morro da Igreja, Urubici, Cachoeira Véu da Noiva, 28°04.595'S, 49°31.090'W, el. 1,300 m, 5.ii1i.1998, Hol- zenthal, Froehlich, Paprocki (MZUSP). Paratypes. Same data as holotype, 2 d, 10 2 (UMSP), 5 2 (MZUSP);). Parand: Sao José dos Pinhais, Serra do Mar, BR 277, km 54, 24°17’S, 50°37'W, el. 750 m, 21.1.1988, PROFAUPAR-lampada, | d (UFPR). San- ta Catarina: Urubici, Cachoeira Avencal, 28°02.839’S, 49°36.997'W, el. 1,260 m, 6.111.1998, Holzenthal, Froehlich, Paprocki, 1 dg, 1 2 (NMNH). Etymology.—This species is named after the entomological inventory of Parana state, Brazil, PROFAUPAR or “Projeto de Lev- antamento da Fauna Entomologica no Esta- do do Parana” (Marinoni and Dutra 1993). ACKNOWLEDGMENTS We are very grateful to the Brazilian fed- eral agency CAPES (Coordenagao de Aper- 29 feigoamento de Pessoal de Nivel Superior) for its financial support to the senior author during a visiting professorship at the Univ- ersidade Federal do Parana in 1997-1998 and to the junior author during her graduate program in entomology at UFPR (BEX 0386/01-4). We are also grateful to Dr. Rog- er J. Blahnik, University of Minnesota, and Dr. Oliver S. Flint, Jr, Smithsonian Insti- their invaluable sorting material, verifying identifications, tution, for assistance in and critically reviewing the manuscript. The senior author is also grateful to Dr. Claudio G. Froehlich and Mr. Henrique Pa- procki, Universidade de Sao Paulo, for as- sistance with field work. Subsequent sys- tematics research was supported by NSF grant DEB-9971885 to Holzenthal and Blahnik. This paper is contribution number 1342 trom the Department of Zoologia, Universidade Federal do Parana (Curitiba). LITERATURE CITED Almeida, G. L. de and L. Marinoni. 2000. Abundancia e sazonalidade das espécies de Leptoceridae (In- secta, Trichoptera) capturadas com armadilha lu- minosa no Estado do Parana, Brasil. Revista Bras- ileira de Zoologia 17(2): 347-359. Flint, O. S., Jr. 1971. Studies of Neotropical caddis- flies, XII: Rhyacophilidae, Glossosomatidae, Phil- opotamidae, and Psychomyiidae from the Amazon Basin (Trichoptera). Amazoniana 3(1): 1-67. Flint, O. S., Jr, R. W. Holzenthal, and S. C. Harris. 1999. Catalog of the Neotropical caddisflies (In- secta: Trichoptera). Special Publication, Ohio Bi- ological Survey. Columbus, Ohio. 239 pp. Hamilton, S. W. 1986. Systematics and biogeography of the New World Polycentropus sensu stricto (Trichoptera: Polycentropodidae), Ph.D. Disserta- tion, Clemson University, Clemson, South Caro- lina, 257 pp. Marinoni, L. and G. L. de Almeida. 2000. Abundancia e sazonalidade das espécies de Hydropsychidae (Insecta, Trichoptera) capturadas com armadilha luminosa no Estado do Parana, Brasil. Revista Brasileira de Zoologia 17(1): 283-299. Marinoni, R. C. and R. R. C. Dutra. 1993. Levanta- mento da fauna entomologica no Estado do Pa- rana, I. Introdugao. Situagdes climaticas e floris- ticas de oito pontos de coleta. Dados faunisticos de agosto de 1986 a julho de 1987. Revista Bras- ileira de Zoologia 8(1/2/3/4): 31-73. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 30-49 A REVISION AND PHYLOGENETIC STUDY OF LIPOCHAETA COQUILLETT (DIPTERA: EPHYDRIDAE) WAYNE N. MATHIS AND MICHELLE D. TRAUTWEIN (WNM) Department of Systematic Biology, Entomology Section, PO. Box 37012, NMNH, MRC- 169, Smithsonian Institution, Washington, DC 20013-7012, U.S.A. (e-mail: mathis.wayne @nmnh.si.edu); (MT) 2845 Shoal Crest, Auistin, TX 78705, U.S.A. (e-mail: traut4 @ yahoo.com) Abstract.—The genus Lipochaeta Coquillett is revised and a phylogenetic analysis of the genera of the tribe Lipochaetini is provided. Lipochaeta is known thus far only from the New World, where there are now two species. The second species, L. ranica, n. sp., (type locality: California), is described herein. The species of Lipochaeta occur in saline or alkaline habitats, especially along maritime coasts. The monophyly of Lipochaeta is well corroborated and its putative sister group is the Old World genus Homalometophus Becker. Key Words: eny Among 114 genera of shore flies (Dip- tera: Ephydridae), less than 17% (19 gen- era) are monotypic (Mathis and Zatwarnicki 1995), and that percentage is decreasing as we better sample the extant fauna and dis- cover additional species. Field work during the last two decades, for example, has re- vealed hundreds of new species but very few new genera. There is also a decrease in monotypic genera as we clarify and rechar- acterize higher level taxa to be more inclu- sive, monophyletic clades, usually compris- ing more than one species. The genus Li- pochaeta Coquillett, the subject of this re- is an example of this evolving pattern in the classification of shore flies. Coquillett (1896) described Lipochaeta slossonae in the late 19th Century, and until now it was the only included species in the genus. Coquillett recognized that this new genus and species were unusual and appro- priately suggested that they be placed in a separate and new subfamily, but he did not vision, revision, Diptera, Ephydridae, Lipochaetini, L. ranica, New World, phylog- provide a subfamilial name. Becker (1896: 275) quoted Coquillett’s entire paper but substituted ““Lipochaetinae” for Lipochaeta in the introductory portion, thus making the subfamilial name available. Attribution of Lipochetinae as a family-group name to Becker was followed in all recent catalogs and is continued here. Sabrosky (1999: 179), however, suggested that, “It seems reasonable to correct the obvious lapse and credit Coquillett with the subfamily.”” Co- quillett clearly was the first person to rec- ognize these taxa, from species to subfam- ily, but the subfamilial name is correctly at- tributed to Becker and the generic and spe- cies names to Coquillett. The bizarre external appearance of Li- pochaeta, being highly adapted to psam- mophilous habitats, initially confused some authors about its familial affiliation. A year after its description, Williston (1897: 8) pre- ferred placement of Lipochaeta **. . . among the Ochthiphilinae in the vicinity of Rhic- VOLUME 105, NUMBER 1 noessa |Tethinidae]’” and added that, ““No Ephydrid that I know of lacks bristles, while both of these latter families [Oscini- dae and Agromyzidae] have numerous forms without them. The face is too short, the antennae too different in structure to be- long with the Ephydridae. Moreover the pollinose body and white wings, while not absent among Ephydridae, are not at all common.” Townsend (1898: 168) de- scribed a second species in Lipochaeta (L. texensis from Padre Island, Texas) and commented on the taxonomic placement of the genus. Townsend wrote that while Li- pochaeta is **... clearly allied to the Ephy- dridae” it “*... is truly one of singular as- pect and anomalous position.”” Aldrich’s (1905) catalog of Nearctic Diptera followed Coquillett and Townsend in listing Lipo- chaeta as a genus in the Ephydridae and also in placing L. texensis as a junior syn- onym of L. slossonae. The synonymy of L. texensis with L. slossonae was apparently based on information in a letter that Willis- ton had written to Aldrich. Although Jones (1906) did not accept Lipochaeta as a shore fly in his worldwide catalog of Ephydridae, he offered no alternative placement. Nearly all subsequent authors have followed Co- quillett, Townsend, and Becker in recogniz- ing Lipochaeta as an ephydrid, usually in the tribe Lipochaetini, subfamily Parydri- nae (Sturtevant and Wheeler 1954; Wirth 1965, 1968; Mathis 1977; Cogan 1980). A notable exception was E. T. Cresson, Jr., the doyen of 20th Century shore-fly workers, who was silent on the subject. Although recognition of Lipochaeta as an ephydrid is now virtually universal, its placement in available shore-fly classifica- tions, especially in catalogs, has varied. Cresson’s. successors (Sturtevant and Wheeler 1954; Wirth 1965, 1968; Mathis 1977: Cogan 1980) accorded tribal status to the genus in the subfamily Parydrinae, ap- parently considering Lipochaeta to be re- lated to the tribes Parydrini and Hyadinini. Giordani Soika (1981), however, was of the opinion that the tribe Lipochaetini 1s related 31 to Isgamera Giordani Soika and Asmeringa Becker, two Old-World genera that occur on the seashores of the Mediterranean and East Africa. Giordani Soika further suggested that the occurrence of Lipochaetini in the New World was due to continental drift and that the group has greater antiquity than was previously thought. Mathis (1984a) concurred with Giordani Soika in removing Lipochaeta from Parydrinae and in placing it close to /sgamera and Asmeringa in the subfamily Gymnomyzinae. The above-cited studies were based on phenetic or overall differences and similar- ities, and although the placement of Lipo- chaeta did change trom Parydrinae to Gymnomyzinae, its position close to /sga- mera and Asmeringa is unsupported. Al- though the latter two genera appear similar externally, these features represent, for the most part, independent and convergent ad- aptations to psammophilous environments that are associated with seashores, not syn- apomorphies that indicate phylogenetic re- lationships. Another advance in the phylogenetic po- sition and composition of Lipochaetini was the recognition that the tribe Atissini, as then characterized, was polyphyletic and in- cluded genera that are more closely related to Hecamede Haliday (tribe Hecamedini, Mathis 1993) and to Lipochaeta (tribe Li- pochaetini, Zatwarnicki 1992, Mathis 1995a). Zatwarnicki (1992) provided evi- dence indicating that the tribe Atissini is re- lated to taxa in the subfamily Hydrelliinae, Whereas Hecamedini and Lipochaetini are in the subfamily Gymnomyzinae. Zatwar- nicki (1992) cited further evidence that He- camedini and Lipochaetini are sister groups. The principal source of characters (synapomorphies) for these studies derives from structures of the male terminalia. The latter studies, especially Zatwarnicki (1992), indicate that the tribe Lipochaetini is closely related to the tribe Hecamedini and includes the following four genera (date and author(s) who first placed the ge- nus in Lipochaetini are noted in parenthe- es) NM sis): Glenanthe (Mathis and Zatwarnicki 1990b), Paraglenanthe Wirth (Zatwarnicki 1992), Lipochaeta (Becker 1896), and Homalometopus Becker (Mathis 1984b). Recent revisionary and phylogenetic stud- ies of genera of the tribe Lipochaetini in- clude Homalometopus (Mathis 1984b, Mu- nari 1988) and Glenanthe (Mathis 1992). METHODS AND MATERIALS The descriptive terminology, with the ex- ceptions noted in Mathis (1986) and Mathis and Zatwarnicki (1990a), follows that pub- lished in the Manual of Nearctic Diptera (McAlpine 1981). Because specimens are small, usually less than 3.5 mm in length, study and illustration of the male terminalia required use of a compound microscope. We have followed the terminology for most structures of the male terminalia that other workers in Ephydridae have used (see ref- erences in Mathis 1986; Mathis and Za- twarnicki 1990a, b). The terminology for structures of the male terminalia is provided directly on Figs. 17-18. The species de- scriptions are composite and not based sole- ly on the holotypes. One head and two ven- ational ratios that are used in the descrip- tions are defined below (all ratios are based on three specimens: the largest, smallest, and one other). Gena-to-eye ratio is the ge- nal height measured at the maximum height divided by the eye height. Costal vein ratio is the straight line distance between the api- ces of R,,, and R,,; divided by the distance between the apices of R, and R,,,. M vein ratio is the straight line distance along vein M between crossveins (dm-cu and r-m) di- vided by the distance apicad of dm-cu. The phylogenetic analysis was performed with the assistance of Hennig86©, a com- puterized algorithm that produces clado- grams by parsimony. Character data were polarized primarily using outgroup proce- dures. Although autapomorphies were not included in the cladistic analysis (they were made inactive), which would skew the con- sistency and retention indices, we listed them on the cladogram and included them PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON as part of generic treatments and phyloge- netic considerations to document the mono- phyly of the lineages, particularly at the ge- neric level. Although many specimens for this study are in the National Museum of Natural His- tory, Smithsonian Institution, Washington, DC (USNM), we also borrowed and studied numerous specimens that are deposited in the following museums: ANSP Academy of Natural Sciences of Philadelphia, Pennsylvania, USA. American Museum of Natural His- tory, New York, USA. The Natural History Museum (for- merly the British Museum (Natural History)), London, England, Unit- ed Kingdom. Canadian National Collection, Ot- tawa, Canada. AMNH BMNH CNC SYSTEMATICS Tribe Lipochaetini Becker Lipochaetini Becker 1896: 275 [as Lipo- chaetinae]. Type genus: Lipochaeta Co- quillett 1896.—Zatwarnicki 1992: 89, 118—119 [listing of included genera, phy- logenetic placement].—Mathis and Za- twarnicki 1995: 160—163 [world cata- log].—Mathis 1995a: 2—4 [description, key to genera]. Diagnosis (synapomorphies indicated by an asterisk (*)).—Head: Frontal vitta (or ocellar triangle) setulose*; ocellar seta ei- ther greatly reduced or absent (sometimes with a pair of intrafrontal setae slightly larger than other setulae, this pair inserted in front of anterior ocellus)*; pseudoposto- cellar seta reduced or lacking*; fronto-or- bital setae 3 (reduced secondarily in Lipo- chaeta), anterior 2 setae proclinate, poste- rior seta reclinate. Pedicel lacking promi- nent, spinelike seta; arista with cuticular hair dorsally and ventrally, appearing ma- cropubescent or brushlike, without dorsal rays*. Eye bearing numerous interfacetal microsetulae (apparently arising from each interfacet). Genal seta reduced or lacking. VOLUME 105, NUMBER | Thorax: Dorsocentral setae weakly de- veloped, only posteriormost pair conspicu- ous; acrostichal setulae in 2—4 rows, fre- quently with a prescutellar pair better de- veloped: postsutural supra-alar seta lacking; frequently postpronotal and presutural supra-alar seta reduced or lacking; posterior notopleural seta inserted above level of an- terior seta, sometimes only slightly so (as in Homalometopus and Lipochaeta). Wing with vein R,,, long, extended nearly to lev- el of apex of vein R,,;. Legs lacking con- spicuous setae; femora and tibiae usually gray to pale brown; tarsi yellow. Abdomen: 5th tergite of male longer than 4th. Male terminalia as follows: epandrium attenuate, either emarginate posteriorly or incomplete dorsally; surstylus well devel- oped, usually elongate, frequently as long or longer than epandrium; aedeagus elon- gate, slender, tubular, apex with recurved flap oriented posterodorsally, apical flap in groove at rest, base of aedeagus bifurcate, sometimes with arms elongate*; ejaculatory apodeme present, compressed laterally *; ae- deagal apodeme L-shaped, sometimes with extended ventromedial process: gonites (pre- and postgonites) lacking, possibly fused with hypandrium*; hypandrium well sclerotized, usually V- or U-shaped; Sth sternite deeply V- or U-shaped into which the surstyli and aedeagus lie at rest. Natural history.—This tribe is unusually tolerant of alkaline or saline aquatic envi- ronments, and species of most genera occur on seashores or are associated with inland aquatic habitats that are saline or alkaline. Discussion.—Although the tribe Lipo- chaetini and each of the included genera are readily characterized, often with substantial evidence that they are monophyletic, the re- lationships among the genera are not well understood. In the key that follows, for ex- ample, we have included numerous char- acters, nearly all autapomorphies, that fa- cilitate identification of the genera but con- tribute virtually nothing to resolution of their phylogenetic relationships. A further complicating factor is the debatable status 33 or polarity of the few characters that may indicate relationships. For example, a gap- ing oral cavity and a wide clypeus are char- acter states that are common to Lipochaeta and Homalometopus, whereas a narrow oral opening and clypeus occur in Glenanthe and Paraglenanthe. The problem is that both the narrow and wide conditions occur in the outgroup, Hecamedini, as well as commonly elsewhere in the Ephydridae, thus confusing issues about which character state is apomorphic within the Lipochaetin1. The tribe Lipochaetini, which is one of six tribes now placed in the subfamily Gymnomyzinae (Mathis and Zatwarnicki 1995), appears to be most closely related to the tribe Hecamedini (Zatwarnicki 1992). Lipochaetini’s sister-group relationship with Hecamedini is corroborated by two syna- pomorphies that we have identified (includ- ing Zatwarnicki’s [1992] characters 59-60): 1) pre- and postgonites apparently fused or greatly reduced; 2) posterior notopleural seta inserted much farther dorsad from no- topleural suture than anterior seta. Lipochaetini are distinguished from He- camedini and other tribes of the subfamily Gymnomyzinae and the tribe’s monophyly is confirmed by the following characters (synapomorphies are noted by an asterisk (*)): 1) body densely invested with micro- tomentum, generally dull colored; *2) ocel- lar seta lacking or weakly developed; *3) pseudopostocellar seta lacking; *4) arista microsetulose, lacking dorsal rays; *5) fron- tal vitta bearing many setulae; 6) posterior margin of gena broadly rounded onto oc- ciput: 7) scutellum with 2 pairs of marginal setae; 8) foreleg normal, forefemur slender, foretibia not having a large spur apically; 9) abdominal tergites 2—4 subequal in width, microtomentose, but more or less smooth; *10) aedeagus with apical flap or appendix that is folded back dorsally; and *11) gonite reduced. With the phylogenetic background of the tribe Lipochaetini within the subfamily Gymnomyzinae established and the mono- phyly of Lipochaetini documented, we now 34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hecamede LOP MG L9=20 Bige I: steps, consistency index 0.82; retention index 0.62). proceed with the cladistic analysis and re- sultant relationships among the included genera, but with a few explanatory remarks first. In the presentation on genus-level re- lationships that follows, the characters used in the analysis are noted first. Each char- acter is immediately followed by a discus- sion to explain its states and to provide per- spective and any qualifying comments about that character. After presentation of the information on character evidence, a hy- pothesis of the cladistic relationships is pre- sented and briefly discussed. The clado- gram (Fig. 1) is the primary mode to con- vey relationships, and the discussion is to supplement the cladogram and is intended only to complement the latter. In the dis- cussion of character data, a “‘O” indicates Table 1. cladistic analysis of Lipochaetini (numbers for char- Matrix of characters and taxa used in the acters correspond with those used in the text). Characters ODODOO0000 1111111 1112 Taxa 123456789 0123456 7890 Hecamede OOO0O000000 0000000 0000 Glenanthe 110100001 O000000 1011 Lipochaeta 000011110 1110211 2102 Homalometopus 000000110 1001101 0011 Paraglenanthe 011100000 1000001 1011 Glenanthe Paraglenanthe 12 14 Se 8 19)" 20 Lipochaeta Homalometopus Cladogram depicting hypothetical cladistic relationships among genera of Lipochaetini (length 17 the state of the outgroup: a “1” or “2” in- dicates the derived states. All multistate characters (7, 14, 17, and 20) were treated as nonadditive (—), and characters 1, 3, 5, 6, 9, 11, 12, 15, and 18, which are auta- pomorphies for various genera or tribes, were made inactive (|) for the analysis so that they do not alter the calculation of the consistency index. The numbers used for characters in the presentation are the same as those on the cladogram, and the se- quence is the same as noted in the character matrix (Table 1). The genus Hecamede, which is the nominate genus for the tribe Hecamedini, was the outgroup in our phy- logenetic analysis. CHARACTERS USED IN THE PHYLOGENETIC ANALYSIS (RUNNING COUNT IN PARENTHESIS.) Head 1(1). Shape of eye: (O) round or oval; (1) pyriform (an autapomorphy for Glenanthe). Shape of clypeus: (0) wide, band- like; (1) narrow (a synapomorphy for Glenanthe and Paraglenanthe). 3(3). Shape or ventral facial margin: (0) flat; (1) emarginate (an autapomor- phy for Paraglenanthe). VOLUME 105, NUMBER | 4(4). 6(6). UG): 8(9). 99). 1(10). 211). 3(12). 4(13). Oral opening: (0) wide, gaping; (1) narrow (a synapomorphy for Glen- anthe and Paraglenanthe). Height of face: (0) normal, usually higher than wide; (1) short (an au- tapomorphy for Lipochaeta). Size of antenna: (0) normal; (1) re- duced, especially the arista (an au- tapomorphy for Lipochaeta). Distance between antennal bases: (O) narrow, distance between anten- nal bases less than antennal width; (1) wide, antennal bases separated by width greater than antennal width (an autapomorphy for Homalome- topus); (2) very wide, antennal bases separated by 3 antennal width (an autapomorphy for Lipochaeta). Shape of mesofrons: (0) narrow, V- shaped, with wide parafrons and fronto-orbits (Glenanthe and Para- glenanthe); (1) wide, broadly U- shaped, parafrons and fronto-orbits comparatively morphy tor Homalometopus and Li- pochaeta). Height of gena: (O) high, 0.33 to more than 0.5 eye height; (1) short, less than % eye height (an au- tapomorphy for Glenanthe). smaller (a synapo- Thorax Katepisternal seta: (0) present, con- spicuous, well developed; (1) great- ly reduced (a synapomorphy_ for Homalometopus, Lipochaeta, and Paraglenanthe). Postpronotal seta: (O) present; (1) re- duced (an autapomorphy for Lipo- chaeta). Presutural supra-alar seta: (O) pre- sent; (1) absent (an autapomorphy for Lipochaeta). Position of posterior notopleural seta: (QO) inserted above level of an- terior seta; (1) inserted at about same level as anterior seta or only slightly elevated (a synapomorphy 5(14). 6(15). 7(16). 1(17). 2(18). 3(19): 4(20). for Homalometopus and Lipochae- ta). Setae along posterior margin of ane- pisternum: (0) 2 setae and some smaller setulae; (1) 1 seta (an auta- pomorphy for Homalometomus); (2) no large setae, only setulae (an au- toapomorphy for Lipochaeta). Wing near apex of subcostal vein and vein R,: (0) normal, membra- nous; (1) sclerotized and thickened, yellow (an autapomorphy for Lipo- chaeta). Wing membrane coloration: (O) hy- aline, transparent; (1) white, trans- lucent (a synapomorphy for Glen- anthe, Paraglenanthe, and Lipo- chaeta). Abdomen Epandrium: (0) entire dorsally; (1) attenuate with a posterodorsal notch (an autapomorphy for Glenanthe and Paraglenanthe); (2) incomplete dorsally with a gap between two lat- eral portions (an autapomorphy for Lipochaeta). Cerci: (OQ) separate, unfused with medial margin of epandrium;: (1) fused laterally with median margin of epandrium (an autapomorphy for Lipochaeta). Length of surstylus: (0) about same length as epandrial length: (1) much longer than epandrial length, usually by more than twice (a synapomor- phy for Homalometopus, Glenanthe, and Paraglenanthe). Base of aedeagus: (0) truncate or nearly so; (1) bifurcate (a synapo- morphy for Homalometopus and Paraglenanthe); (2) arms of basal bifurcation more elongate and curled beneath (an autapomorphy for Li- pochaeta). ANALYSIS AND RESULTS Using the implicit enumeration (ie*) op- tion of Hennig86, which is an exhaustive 36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Index = Retention Index. Analysis of characters based on the cladogram (Fig. 1). Con. Index = Consistency Index; Ret. Characters 1 2 3 4 5 6 7 8 9 10 Steps 1 1 l 1 1 | | 1 I 2 Con. Index 100 LOO 100 100 100 100 100 100 100 50 Ret. Index 100 100 100 100 100 100 100 100 100 0 Characters 11 12 13 14 15 16 17 18 19 20 Steps ] 1 l 2 ] 2 2; 1 2 2 Con. Index 100 100 100 100 100 50 100 100 50 100 Ret. Index 100 100, 100 100 100 0 100 100 0 100 search, a single most parsimonious tree was generated from the analysis of the 20 char- acters. The cladogram has a length of 17 steps and consistency and retention indices of 0.82 and 0.62 respectively. The matrix was then subjected iteratively to successive cc) to determine a character’s contribution or weight and to find cladograms supported by the most con- sistent characters (Carpenter 1988, Dietrich and McKamey 1995). The analysis of the characters for this cladogram is given in Ta- ble 2 and the weights of the various char- acters is given in Table 3. Given these char- acter weights, the analysis of the resultant cladogram resulted in consistency and re- tention indices of 1.00 and 1.00 respective- ly. As indicated on the cladogram (Fig. 1), the tribe Lipochaetini is divided into two basal sublineages. The first basal sublineage comprises half of the genera (number of species in parenthesis): Lipochaeta (2) and Homalometopus (7), with 9 species. The weighing (xs w, ie*, Table 3. = ]) of characters after successive weighting. Weights (varying between 1—10) and status (additive = +, monophyly of this sublineage is corrobo- rated by characters 7, 8, and 14. From a biogeographic standpoint, this sister-group relationship suggests that the cladogenetic event(s) that resulted in the divergence of Lipochaeta and Homalometopus occurred some time ago. Homalometopus is only found in the Old World where it now has relatively high diversity, 1.e., seven species representing several speciation events, es- pecially in the area of the Mediterranean Sea. Lipochaeta occurs only in the New World where speciation has been slower, re- sulting in only two described species and the possibility of a third in Chile. The second basal sublineage includes the other two genera of Lipochaetini, Glenan- the (15) and Paraglenanthe (3), with 18 species. The monophyly of this sublineage is corroborated by characters 2, 4, and 17. Glenanthe occurs in both the New and Old Worlds, but there is apparently greater spe- cies diversity in the New World, assuming the present number of described species is nonadditive = —, active = [, inactive Characters i 2 3 4 5 6 7 8 9 10 Weight, status 10+] 10+[ 10+] 10+[ 10+] 10+] 10—-[ 10+[ 10+] O+[ Characters Il 2 13 i4 1S 16 17 18 19 20 Weight, status 10+] 10+] 10+[ 10-[ 10+] 0+[ 10-[ 10+] O+| LO={ VOLUME 105, NUMBER | a good indicator. Paraglenanthe, on the other hand, has a more limited distribution, occurring only in the New World where it is primarily circumcaribbean. KEY TO GENERA OF LIPOCHAETINI BECKER 1. Eye pyriform, distinctly narrowed ventrally; gena short, less than one-fourth eye height; ka- tepisternal seta present along posterior margin, moderately well developed Glenanthe Haliday [15 species, worldwide; Mathis 1995a] — Eye generally oval or round, not distinctly nar- rowed ventrally; gena high, one-third or more of eye height; katepisternal seta reduced ... . i) i) . Antenna reduced, inserted in well-separated cavity, arista atrophied, budlike; face short, height subequal to length of reduced antenna: clypeus bandlike; oral opening gaping: body setae and setulae pale; costa distinctly thickened at merger of vein R, ..... Lipochaeta Coquillett [2 species, New World: mostly costal marine on sand but also inland where saline conditions exist] — Antenna normally developed, not in deep cav- ity, arista as long as flagellomere 1; face well developed, height much more than length of antenna; clypeus variable; oral opening narrow or gaping; setae and setulae largely black; costa only slightly thickened at merger of vein R, .. 3 3. Mesofrons large, occupying most of frons, platelike, subrectangular, uniformly and evenly setulose; ventral facial margin flat; clypeus wide, bandlike; oral opening large, gaping Bea ie Cler ight CRE eRe eee Homalometopus Becker [7 species, Eastern Hemisphere (Mediterranean); Mathis 1984b, Munari 1988] — Frons lacking differentiated mesofrons, at most with frontal or ocellar triangle or vitta that is weakly differentiated from remainder of frons; ventral facial margin emarginate; clypeus nar- row, exposed through ventral facial emargina- tion; oral opening small Paraglenanthe Wirth [3 species, New World (Caribbean); Wirth 1956| Genus Lipochaeta Coquillett Lipochaeta Coquillett 1896: 220. Type spe- cies: Lipochaeta slossonae Coquillett 1896, original 1896: 274-275 [quote of original de- scription].—Williston 1897: 7 [placement near Rhicnoessa, family Agromyzidae, subfamily “‘“Ochthiphilinae” |; 1908: 306 [generic key].—Townsend 1898: 168 [notes on generic placement].—Aldrich designation.—Becker 1905: 631 [Nearctic catalog].—Jones 1906: 169 [note, diagnosis].—Curran 1934: 346-347 [figs. of head, generic key].—Sturtevant and Wheeler 1954: 215-216 [listed as a genus in Napeae].— Wirth 1965: 750 [Nearctic catalog]; 1968: 22 [Neotropical catalog].—Mathis and Zatwarnicki 1995: 163 [world cata- log]. Diagnosis.—Lipochaeta 1s distinguished from other genera of Lipochaetini by the following combination of characters: Body setae and setulae generally inconspicuous, pale. Antennae widely separate and arista rudimentary. Frons projected and _ large; sub-cranial cavity large, gaping. Body length 2.0—4.0 mm, generally grayish, dor- sum of head and thorax sometimes brown- ish, and almost entirely microtomentose. Description.—Head: Wider than high in anterior view, grayish; entirely microto- mentose, sparsely setulose; mesofrons wide, broadly U-shaped, uniformly setulo- se, differentiated from remainder of frons by darker gray to brownish gray; parafrons a comparatively small triangular anterior area; fronto-orbits comparatively narrow. Head without conspicuous setae. Ocellar setae absent; pseudopostocellar setae ab- sent, fronto-orbital setae absent (reduced secondarily); ocelli arranged to form equi- lateral triangle. Antenna reduced, in well- separated (gap between antennal bases about 3X antennal width), deep cavities, oriented laterally; pedicel with fine, pale se- tulae on ventral and dorsal sides, but lack- ing dorsoapical seta, arista rudimentary and budlike. Eye generally irregularly round with distinct, angular margin posterodorsal- ly near vertex; interfacetal present, numerous, conspicuous. Face short, height subequal to length of reduced anten- na, narrowly triangular in profile, tapered to acute angle posteroventrally; clypeus band- like, wide. Gena bearing fine, pale setulae. Subcranial cavity large and gaping. Thorax: Entirely grayish to brownish, microtomentose, pleural area generally pal- microsetulae 38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON er than dorsal coloration; mesonotum bear- ing numerous, fine, pale setulae, setulae most conspicuous on posterior portion of scutellum, particularly in females; post- pronotal seta reduced: posterior notopleural seta at only slightly elevated position rela- tive to anterior seta; anepisternum bearing numerous setulae: katepisternal seta vari- able. Wing with costal vein extended to vein M: costa distinctly thickened at merger of vein R,; dorsal setulae along costal vein ended just beyond vein R,.;; wing white, translucent, wing venation and halter yel- low. Midfemur bearing comblike row of 6— 7 white, longer setae anteriorly at apical 4— Ys: tarsi yellowish; midtarsus with basitar- somere longer than remaining four; each tarsomere with dark, ventroapical setae; midtarsus bearing twice as many black se- tae as others, these paired at apices of tar- someres, basitarsomere bearing 4—5 pairs, apical tarsomere variable; pulvillus present. Abdomen: Male: tergites 2—4 about equal in length; length of Sth tergite slightly more than combined length of 3rd and 4th tergites; 5th tergite triangular in dorsal view: sternites 2—4 linear, much narrower than long, weakly sclerotized; 5th sternite deeply U-shaped and better sclerotized along inner margin of U. Male Terminalia: epandrium greatly reduced, in lateral view about as long as cerci, fused dorsolaterally with cerci, incomplete dorsally, with a gap between lateral portions: surstylus digiti- form, moderately to conspicuously elon- gate, mostly parallel-sided, usually shallow- ly curved in lateral view, generally evenly setulose, setulae longer apically: gonites ei- ther indistinguishably fused with hypan- drium or lacking: ejaculatory apodeme ev- ident as a simple, slightly angulate, long, narrow process: aedeagus very long, nar- row, tubular, length twice or slightly longer than surstylus, apex with membranous flap folded back on itself, length of flap about %—'4 length of aedeagus, basal portion of aedeagus angled ventrally and forked with aedeagal apodeme between arms of basal fork; hypandrium well sclerotized, more or less Y-shaped in ventral view, with forked portion basal, lateral phalanges from basal arms of fork. Female Terminalia: Segments 6-7 telescoped out to form tube; 8th seg- ment apparently lacking: hypoproct Y- shaped. Remarks.—The poorly developed epan- drium and an aedeagus with a terminal membranous flap are similar to Glenanthe and Homalometopus Becker. The female terminalia is also similar to that of Homa- lometopus. Until now, Lipochaeta has been mono- typic. with L. slossonae as the only includ- ed species. The second species, L. ranica, is very similar and obviously closely relat- ed. Differences between these two species are seemingly slight but are consistent and significant (see key and descriptions of spe- cies below). In addition to describing a sec- ond species, which is found primarily on the west coast of California and Mexico, we also report the possibility of a third species from Chile. The potential third species is represented by two female specimens from Atacama. One of the females has an elon- gate scutellum and bears a fringe of long, white setulae along the margin. An elongate scutellum also occurs in L. slossonae and L. ranica, and to a degree, the scutellar fringe of setulae is also apparent within the variation of these two species. Thus we are unsure about the status of the Chilean spec- imens, i.e., is the variation in the scutellar features intra- or interspecific. Additional specimens, especially males, are needed to assess better these possibilities. KEY TO SPECIES OF LIPOCHAETA 1. Tarsi often entirely yellow: surstylus elongate (distinctly longer than height of epandrium), tapered apically to narrowly rounded apex (east coast of North America from Massachusetts south to Florida and Texas, islands of the Ca- ribbean, Belize [Stann Creek]. Mexico [Chia- pas], and Panama [Playa Santa Clara] .... SO hes SEE blo ee L. slossonae Coquillett — Apical tarsomere always slightly to distinctly darkened; surstylus shorter (subequal to height of epandrium), parallel sided to bluntly round- ed apex (west coast of North America from VOLUME 105. NUMBER | 39 Chip Figs. 2-7. Scanning electron micrographs of Lipochaeta slossonae. 2, Head, anterodorsal view. 3, Same, lateral view. 4, Frons, anterodorsal view. 5, Face, anterior view. 6, Right antenna, anterior view. 7, Left compound eye and interfacetal setae, anterior view. Scale bars equals 100 pm. San Francisco south to the Mexican states of Florida, Maryland, New Jersey, Texas, etal California Sur and Nayarit, and to the Ga- comments on habitat and behavior].— lapagos Islands) ....... L. ranica, new species J : az Wirth 1956: 18 [list, Bahamas]; 1965: Lipochaeta slossonae Coquillett 751 [Nearctic catalog (partim)]; 1968: 22 (Figs. 2-19) [Neotropical catalog (partim)].—Foote Lipochaeta slossonae Coquillett 1896: 1995: 422 [sandy beach habitat ].—Math- 220.—Becker 1896: 275 [description].— is and Zatwarnicki 1995: 163 [world cat- Aldrich 1905: 631 [Nearctic catalog (par- alog (partim)].—Mathis 1997: 28—29 [re- tim)]—Jones 1906: 169 [note, diagno- view, Belize]. sis]|—Johnson 1913: 86 [list, Florida] — Lipochaeta texensis Townsend 1898: Sturtevant and Wheeler 1954: 216 [list, 168.—Aldrich 1905: 631 [synonymy ].— 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 8-15. view. 10, Scutellum, dorsal view. 11, Pleural area, lateral view. 12, Anepisternum, dorsoblique view. 13, Left Scanning electron micrographs of Lipochaeta slossonae. 8, Thorax, lateral view. 9, Same, dorsal midfemur, anterior view. 14, Left midtarsus, ventral view. 15, Abdomen, dorsal view. Scale bar equals 100 pm. Sturtevant and Wheeler 1954: 216 [syn- onymy]. Diagnosis.—Moderately small to moderately large shore flies, body length 2.0—4.0 mm. Description.—Head (Figs. 2—7): In pro- file as long as high. Mesofrons slightly dif- ferentiated by darker brown or gray micro- tomentum. Setae and setulae generally in- conspicuous; genal setulae shorter than ane- pisternal setulae. Thorax (Figs. 8-16): Mesonotum vary- ing from slightly darker brown to gray. Anepisternal setae moderately long; kate- pisternal seta reduced. Wing as in Fig. 16. Tarsi usually yellow, apical tarsomere slightly darkened in some specimens. Abdomen (Figs. 17-18): Surstylus elon- gate (distinctly longer than height of epan- drium), tapered apically to narrowly round- ed apex. Type material.—Described from two od. The lectotype ¢ Coquillett, here designated to preserve sta- of Lipochaeta slossonae bility and make more universal the use of this name, is labeled ‘**‘CH[arlotte]. H[ar]B[o]R, FL[orid]A [folded]/Mrs- Slosson Collector/Type No 4300 U.S.N.M. [red; “4300” handwritten |/Lipochaeta slos- sonae Coq. [handwritten; black sub-bor- The double (glued to a paper triangle along with a sec- der.” lectotype is mounted ond specimen, also a male, that lacks its head), is in good condition, and is deposited in the USNM (4300). Coquillett (1896) wrote that the two syntypes were specifi- cally from Punta Gorda, which is a city on the eastern shore of Charlotte Harbor bay. The second male is a paralectotype. The syntype(s) of Lipochaeta texensis is from USA. Texas: Cameron County, Padre Island; ST (sex ?), BMNH. We were unable to locate this specimen, which apparently VOLUME 105, NUMBER 1 was in the BMNH. Its whereabouts is un- known. Other specimens examined.—ANGUIL- LA. Long Pond Bay (18°13’N, 63°01'W), 29 Mar 1992, W. E. Steiner, J. M. Swear- ingen (37 6, 41 2; USNM); Prickly Pear, (18°16'10"N, 63°10’30"W), 30 Mar 1992, W. E. Steiner, J. M. Swearingen (7 d, 11 2; USNM); Sandy Ground (18°12'20"N, 63°05'30"W), 24-30 Mar 1992, W. E. Steiner, J. M. Swearingen (2 2; USNM). BAHAMAS. Great Inaqua Island: Mat- thew Town, 31 Jan 1953, E. B. Hayden, G. B. Rabb (1 2: AMNH). Turks and Caicos Islands: Grand Turk Island, 19 Feb 1953, E. B. Hayden (3 6, 2 2; AMNH); South Caicos Island, 11 Feb 1953, E. B. Hayden (1 36,3 2; AMNH). BELIZE. Stann Creek: Carrie Bow Cay, 31 May 1985, W. N. Mathis (2 6, 4 &; USNM). CUBA. Sancti Spiritus: La Boca (4 km S; 21°45.9'N, 80°01.5’W), 12 Dec 1994, W. N. Mathis (8 36, 3 2; USNM); Playa Ancon (21°44.1’N, 79°59.9'W), 12 Dec 1994, W. N. Mathis (2 6, 3 2; USNM). CURACAO. Coral Specht (3 km E Wil- lemstad), 8-15 Feb 1987, W. E. Steiner, J. M. Swearingen (1 2; USNM). DOMINICAN REPUBLIC. Azua: Puerto Viejo (18°20.9'N, 70°50.4’'W), 14 May 1995, W. N. Mathis (2 d, 2 2; USNM). Barahona: Barahona (18°12'N, 71°5.3'W), 20 May 1998, D. and W. N. Mathis (4 6, 1 2; USNM). La Romana: Isla Saona, Mano Juan (18°08.1'N, 68°44.5'W), 13 May 1995, W. N- Mathis (il 6, 2 9; USNM). Monte Cristi: Monte Cristi (beach; 19°51.5'N, 71°39:5'W), 18 May 1995, W. N. Mathis (6 3d; USNM). GRAND CAYMAN. Double Head (19°23.4'N, 81°22.3’W), 27 Apr 1994, D. and W. N. Mathis (8 d, 4 2; USNM). GUYANA. Mahaica (6°42.8'N, 57°55.6'W), 14 Apr-20 Aug 1994, 1997, W. N. Mathis (1 d, 1 2; USNM). JAMAICA. Clarendon: Barnswell Beach (17°45'N, 77°08.5'W), 13 May 1996, D. and W. N. Mathis, H. Williams (1 6; 41 USNM); Farquhars Beach (17°50.9'N, 77°22.8'W), 9 May 1996, D. and W. N. Mathis, H. Williams (5 6, 5 2; USNM); Jackson Bay (17°44.7'N, 77°12.6’W), 13 May 1996, D. and W. N. Mathis, H. Wil- liams (6 d, 3 2; USNM). St. Elizabeth: Great Pedro Pond (17°52.5'N, 77°45.2'W), 19 Apr 2000, W. N. Mathis (4 6, 4 9; USNM); Salt Pond, Parottee Beach (17°58.1'N, 77°50.2'W), 19 Apr 2000, W. N. Mathis (2 2; USNM). MEXICO. Chiapas: Puerto Arista (2 km E), 18 May 1985, A. Freidberg, W. N. Mathis (10 6, 31 2; USNM). PANAMA. Cocle: Playa Santa Clara, 2 Jul 1967, W. W. Wirth (1 36, 1 2; USNM). PUERTO RICO. Bahia Salinas (beach; 17°57.5'N, 67°12'W), 20 Sep 1995, D. and W. N. Mathis (4 6, 7 2; USNM). Punta Jacinto (near Guanica; WoT ENG 66°52.6'W), 20 Sep 1995, D. and W. N. Mathis (1 6; USNM). UNITED STATES. Florida. Charlotte: Charlotte Harbor (2 6; USNM); Punta Gor- da, 27 Jan 1932, A. L. Melander (1 6; ANSP). Duval: Little Talbot Island State Park, 28 May 1953, M. R. Wheeler (2 6, 2 2; USNM). Lee: Bonita Beach, 17 Apr 1970, W. W. Wirth (10 6, 19 2; USNM); Sanibel Island, Lighthouse Park, 17 Apr 1989, D. and W. N. Mathis (4 ¢; USNM). Manatee: Anna Maria, 19 May 1953, M. R. Wheeler (2 6, 1 2; USNM). Monroe: Ev- erglades National Park (on white sand beach), 3 Apr 1958, H. V. Weems, Jr. (1 6; USNM); Key West, 5 Feb 1938, A. L. Me- lander (6 6, 4 2; ANSP); Matecumbe, 2 Feb 1938, A. L. Melander (2 3; ANSP). Pinellas: St. Petersburg, 29 Jan-30 Mar 1932, E. T. Cresson, Jr., A. L. Melander (4 3d, 13 2; ANSP). Georgia. Liberty: St. Catherines Island, 21 Apr—30 Oct 1977, 1978, A. Hook, R. W. Matthews, J. Krispyn (1 gd, 1 2; USNM). Chatham: Chatham (at light); ict 1953, Hi. FE Schoof. (1 6; USNM). Maryland. Calvert: Chesapeake Beach, 20 Aug—20 Sep 1914, J. M. Aldrich, N. (Banks; (G. He ‘Greéne (28 6, 15 2; ANSP, USNM). Massachusetts. Barnstable: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 16 cercus epandrium es aedeagal apodeme — aedeagus Yaz Figs. 16-18. Wing and structures of the male terminalia of Lipochaeta slossonae (ejaculatory apodeme not illustrated; Florida. Lee: Bonita Beach). 16, Wing, dorsal view. 17, Cerci (left side only), epandrium (left side only), surstylus, aedeagus, aedeagal apodeme, and hypandrium, lateral view. 18, Same (hypandrium lacking), posterior view. Scale bar equals 0.2 mm. VOLUME 105, NUMBER | Sagamore Beach, 14 Jul 1932 (1 9; USNM). Mississippi. Harrison: Ship Island (north side; 30°12.6'N, 88°58’W), 27 Jun 1962, D. L. Deonier (1 2; USNM). Jack- son: Horn Island (taken in inter-tidal zone on sand surface; 30°15'N, 88°42’W), 13 Jun 1962, D. L. Deonier (1 2; USNM). New Jersey. Cape May: Cape May, 6 Jun 1971 (1 3; ANSP); Wildwood, 18 Jul-18 Sep 1908, 1920, E. T. Cresson, Jr. (25 6, 29 @; ANSP). New York. Weschester: Oak Island, 31 Jul 1913 (3 36, 1 2; ANSP). North Car- olina. Onslow: Ashe Island (emergence trap), 7 Oct 1976, T. D. Edwards (2 6; USNM). Oklahoma. Alfalfa: Cherokee Salt Plains, 18 Jun 1930, R. D. Bird (3 6; USNM); Great Salt Plains (salt marsh), 22 May 1969, W. W. Wirth (6 ¢, 3 2; USNM). South Carolina. Beaufort: Hilton Head Is- land (UV light trap), 29 May 1980, P. H. Adler (2. 6; USNM). Texas. Brazoria: Freeport, 9 Mar 1951 (1 ¢6; USNM). Cal- houn: Indianola, 28 Apr—S Jun 1951, 1953 (4 36, 1 2: USNM). Cameron: Port Isabel, 23 Mar-11 Dec 1910, 1951 (1 6, 6 @; ANSP, USNM). Galveston: Galveston, May—26 Dec 1900, 1917, 1951, J. M. Al- drich, E H. Snow, M. R. Wheeler (21 ¢, 19 2; ANSP, USNM). Kleberg: Riviera, 23 Mar 1951 (8 6, 4 2: USNM). Nueces: Cor- pus Christi, 18 Mar 1908, Jones and Pratt (1 do, 3 2; USNM). San Patricio: Sinton, 23 Mar 1951 (2 2; USNM). Distribution (Fig. 19).—Nearctic: USA (FL, GA, MA, MS, NC, NJ, NY, OK, SC, TX). Neotropical: Bahamas, Belize, Cura- cao, Guyana, Mexico (CHI), Panama, West Indies (Anguilla, Cuba, Dominican Repub- lic, Grand Cayman, Jamaica, Puerto Rico). Natural history.—This species occurs on the bare surface of wet, sandy, maritime beaches, especially in embayments and ar- eas where the beach is somewhat protected from the direct and abrading actions of waves. Although saline habitats are primar- ily coastal maritime, this species also oc- curs inland, such as in Oklahoma, where beaches associated with saline environ- 43 ments, usually as saline playas, are also found. Remarks.—Although we were unable to locate and study the syntypes of the junior synonym, L. fexensis, we are reasonably confident that the specimens are conspecific with the senior synonym and that the syn- onymy Aldrich (1905) proposed is correct. All specimens we have examined from Tex- as, including specimens from coastal sites near Padre Island, the type locality of L. texensis, are conspecific. If the syntype is found, the question of conspecificity should be confirmed and a lectotype designated. Lipochaeta ranica Mathis and Trautwein, new species (Figs. 20-31) Lipochaeta slossonae of authors, not Co- quillett [misidentification].—Townsend 1898: 168 [abundance on seashore in southern California].—Aldrich 1905: 631 [Nearctic catalog (partim), note on abun- dance in southern California].—Cole 1912a: 838 [note]; 1912b: 161-162 [list, habitus fig.]; 1923: 478 [list, Loreto, Isla Partida]; 1969: 392, 404 [generic key, discussion].—Wirth and Stone 1956: 465, 470 [generic key, list, California].— Wirth 1965: 751 [Nearctic catalog (par- tim)]; 1968: 22 [Neotropical catalog (par- tim)].—Cheng and Lewin 1974: 167-168 [behavior, feeding habits].—Mathis 1995b: 4 [Galapagos Islands].—Zatwar- nicki 1992: 112 [figs. of male termina- lia].—Mathis and Zatwarnicki 1995: 163 [world catalog (partim)]. Diagnosis.—Moderately small to medi- um-sized shore flies, body length 2.10—3.80 mm. Description—Head (Figs. 20-23): In profile appearing projected, longer than high. Mesofrons differentiated as dark brown or dark gray. Setae and setulae gen- erally more conspicuous than L. slossonae; length of genal setulae equivalent to length of anepisternal setulae, giving a more se- tulose appearance. 44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 19 Distribution map for Lipochaeta slossenae Thorax (Figs. 24—28): Mesonotum vary- ing from dark brown to dark gray. Anepis- ternal setae long, setae appearing brushlike; Katepisternal seta reduced but other setulae usually evident. Legs, particularly tibiae, bearing long setae, appearing brushlike; apical tarsomere partially to completely darkened. Abdomen (Figs. 29-30): Surstylus short- er (subequal to height of epandrium), par- allel sided to bluntly rounded apex. Type material—The holotype ¢ is la- beled “‘CoronaDelMar 29/6/42 [29 Jun 1942] CAL|[IFORNIA] AlMelander/AIMe- lander Collection 1961 [right % with green stippling|//HOLOTYPE <¢ Lipochaeta rani- ca W. N. Mathis & Trautwein USNM [red: species name, gender symbol, and ““& Trau- twein” handwritten].”° The holotype is dou- ble (minuten in a card), is in excellent condition, and is de- posited in the USNM. Eighty-one paratypes (37 6,44 2; USNM) bear the same locality data as the holotype. Other specimens examined.—ECUA- DOR. Galdpagos Islands: Isabela: Puerto Villamil (littoral, sweeping sand beach), 7 Mar 1989, B. J. Sinclair (5 6, 1 2; CNC). Santa Cruz: Charles Darwin Research Sta- tion (littoral, intertidal rocks, beach at high tide line), 17-27 Jan 1989, B. J. Sinclair (10 3, 11 2; CNC, USNM). mounted rectangular VOLUME 105, NUMBER 1 RGR 5 er ROT ETI Figs. 20-28. ‘ Scanning electron micrographs of Lipochaeta ranica. 20, Head, anterodorsal view 21, Same, lateral view. 22, Same, anterior view. 23, Interfacetal setae, lateroblique view. 24, Left pleural area, lateral view 25, Left anepisternum, lateral view view. 28, Left midfemur, anterior view MEXICO. Baja California Norte: San Felipe, 19 Feb 1954, P. H. Arnaud, Jr. (15 3, 7 2; USNM); San Felipe (3 mi N), 25 Mar 1964, M. E. Irwin (2 6, 2 2; USNM); Ensenada, 24 Jun 1950, A. L. Melander (20 3, 16 2; USNM); San Quintin, 7 May—25 Sept 1925, 1950, H. H. Keifer, R. L. Lang- ston (3 6; USNM); Bellandra Bay: Isla Carmen, Gulf of California, 28 Mar 1953, P.H. Arnaud, Jr. (6 ¢, 6 2; USNM); Loreto (Baja), 29 Mar—19 May 1921, 1953, P. H. Arnaud, Ir, EP Van Duzee (2 ¢, 12 9; USNM). Nayarit: Agua Verde Bay, 23 Mar 1953, P. H. Arnaud, Jr. (6 6, 1 2; USNM); San Blas, 2 Apr 1964, E. I. Schlinger (1 USNM). Sonora: Puerto Penasco, 28 Apr 1948 (1 2; USNM). UNITED STATES. California. Los An- geles: Huntington Beach, 7 Oct 1929, L. D. Anderson (1 ¢; USNM); Laguna Beach, 27 26, Notopleuron and anepisternum, dorsoblique view. 27, Thorax, dorsal Scale bar equals 100 2m Aug 1943, E R. Cole, A. L. Melander (2 USNM); Long Beach, 4 Aug—21 Oct 1911, 1935, R. W. Doane, A. L. Melander (17 6, 17 2; USNM); San Pedro, 8 Jul 1950; TE D: A. Cockerell @ ANSP. USNM); Seal Beach, 26 Jul 1942, A. L. Melander (1 6: USNM). Santa Barbara: Carpinteria, 12—17 Jun 1955, A. L. Melan- der (6 6, 1 2; USNM); Santa Barbara, 6 Jul 1917, J. M. Aldrich (1 d, 2 2; USNM). San Benito: Castroville, 19 May 1950, A. H. Sturtevant (11 ¢, 8 2: USNM). San Di- ego: La Jolla, 22 Jun—16 Aug 1935, 1950, M. T. James, A. L. Melander (2 ¢; ANSP. USNM); San Diego, 5 Apr—3 Aug 1915, 1917, 1932, J. M. Aldrich, M. Van Duzee (11 6, 12 2; ANSP, USNM). San Francis- co: San Francisco Bay (salt marshes), 11 Oct 1906, R. W. Doane (3 6, 1 2; ANSP). San Luis Obispo: Morro Bay, 30 Aug 1945, 46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 29-30. Lipochaeta ranica (ejaculatory apodeme not illustrat- ed; Mexico. Baja California Norte: San Fellipe). 29, Cerci (left side only), epandrium (left side only), sur- stylus, aedeagus, aedeagal apodeme, and hypandrium, lateral view. 30, Same, posterior view. Scale bar equals 0.2 mm. Structures of the male terminalia of A. L. Melander (1 36, 1 2; USNM). San Mateo: San Mateo, 19 May 1950, A. H. Sturtevant (7 6, 2 2; USNM). Distribution (Fig. 31).—Nearctic: USA (CA). Neotropical: Ecuador (Galapagos Is- lands), Mexico (BCN). Etymology.—The species epithet, rani- ca, is of Latin derivation and refers to the froglike face of this species. Townsend (1898: 169), in referring to the head of Li- pochaeta, described it as **frog-mouthed” or “‘frog-faced,”” and we concur (Fig. 22). Remarks.—All specimens from Califor- nia and Baja California Norte (Mexico) that we have examined were previously mis- identified as L. slossonae and represent this new species (see species synonymy). Cheng and Lewin (1974) reported on the peculiar ““fluidisation” of the substrate as part of the feeding behavior of this species (as L. slossonae). From observations in both the field and laboratory, they discov- ered that this species fluidizes the wet sand on the beach through a shaking motion that apparently loosens the interstitial microflo- ra, which is then imbibed as a kind of soup. Variation is evident between some pop- ulations of this species. Externally, the specimens from the Galapagos Islands are very similar to L. slossonae, especially in the length and number of setae and setulae and to a lesser degree in coloration. Struc- tures of the male terminalia indicate that these specimens are L. ranica, however. ACKNOWLEDGMENTS We gratefully acknowledge the assis- tance and cooperation of many organiza- tions and individuals who contributed to the field work and production of this paper. To David A. Grimaldi (AMNH), Jon K. Gelhaus and Donald KF Azuma (ANSP), Nigel Wyatt (BMNH), J. Richard Vocker- oth (CNC), and their institutions, who loaned specimens, we express our sincere thanks. The illustrations were carefully inked by Young T. Sohn. Hollis B. Williams provid- ed technical support. Susann Braden and Scott Whittaker assisted with the prepara- tion of the scanning electron micrographs. For reviewing a draft of this paper, we thank Amnon Freidberg, Stephen D. Gai- mari, and Tadeusz Zatwarnicki. We are also grateful to David Challinor, former Assis- tant Secretary for Research, and to Anna K. Bernensmeyer and David L. Pawson, for- mer Associate Directors for Science, Na- tional Museum of Natural History, Smith- sonian Institution, for financial support to conduct research at The Natural History Museum (BMNH), London, England, through grants from the Research Oppor- tunity Fund. In 1995, 1996, and 1998, field work on the West Indies was funded in large measure by grants from the Biodiver- sity Program (Biotic Surveys and Invento- ries, BSI), National Museum of Natural History, Smithsonian Institution (Lynne R. VOLUME 105, NUMBER 1 Fig. 31. Distribution map for Lipochaeta ranica. Parenti, former chair, George R. Zug, chair). Field work on the West Indies was greatly expedited through the able and pleasant assistance of N. Dianne Mathis, Hollis B. Williams, Kelvin Guerrero, Dan- iel E. Pérez-Gelabert, and Oliver S. Flint, Jr. Field work in Guyana was supported by the Smithsonian Institution’s Biological Di- versity of the Guianas Program (publication number 65; Vicki A. Funk, Director; Carol L. Kelloff, Coordinator). LITERATURE CITED Aldrich, J. M. 1905. A catalogue of North American Diptera (or two-winged flies). Smithsonian Insti- tution, Smithsonian Miscellaneous Collections 46(1444): 1-680. Becker, T. dae. Berliner Entomologische Zeitschrift 41(2): 91-276. Carpenter, J. M 1896. Dipterologische Studien IV. Ephydri- 1988. Choosing equally parsimonious cladograms. Cladistics 4(3): 291-296. Cheng, L. feeding 250(5462): 167-168. Cogan, B. H. 1980. 71. Family Ephydridae, pp. 655— 669. In Crosskey, R. W., ed. Catalogue of the Dip- tera of the Afrotropical Region, 1,437 pp. British among multiple 1974. Fluidisation as a Nature and R. A. Lewin. mechanism in beach flies. Museum (Natural History), London. Cole, E R. 1912a. Notes on the Diptera of Laguna 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Beach. Pomona College Journal of Entomology 4(3): 837-840. . 1912b. Some Diptera of Laguna Beach. First Annual Report of Laguna Marine Laboratory 1: 150-162. . 1923. Expedition of the California Academy of Sciences to the Gulf of California in 1921. Dip- tera from the islands and adjacent shores of the Gulf of California. Il. General report. Proceedings of the California Academy of Sciences, Series 4 12(25): 457-481. Cole, FE R. (with the collaboration of E. I. Schlinger). 1969. The Flies of Western North America. Uni- versity of California Press, Berkeley and Los An- geles, xi + 693 pp. Coquillett, D. W. 1896. A new subfamily of Ephydri- dae. Entomological News 7(7): 220-221. Curran, C. H. 1934. The Families and Genera of North American Diptera. The Ballou Press, New York, 512 pp. Dietrich, C. H. and S. H. McKamey. 1995. Two new neotropical treehopper genera and investigation of the phylogeny of the subfamily Membracinae (Homoptera: Membracidae). Proceedings of the Entomological Society of Washington 97(1): 1— 16. Foote, B. A. 1995. Biology of shore flies. Annual Re- view of Entomology 40: 417—442. Giordani Soika, A. G. 1981. Sulla presenza nelle la- gune delle coste atlantiche dell’ America Centro- Settentrionale della Lipochaeta slossonae Cogq., dittero efidride di tipo paleosteppico-mediterra- neo. Bollettino del Museo civico di Storia naturale di Venezia 31: 69-77. Johnson, C. W. 1913. Insects of Florida I. Diptera. Bul- letin of the American Museum of Natural History 32: 37-90. Jones, B. J. 1906. Catalogue of the Ephydridae, with bibliography and description of new species. Uni- versity of California Publications in Entomology 1(2): 153-198. Mathis, W. N. 1977. Key to the Neotropical genera of Parydrinae with a revision of the genus Eleleides Cresson (Diptera: Ephydridae). Proceedings of the Biological Society of Washington 90(3): 553-565. 1984a. A revision of the genus Asmeringa Becker (Diptera: Ephydridae). Israel Journal of Entomology 17(1983): 67-79. . 1984b. A revision of the shore fly genus Hom- alometopus Becker (Diptera: Ephydridae). Pro- ceedings of the Biological Society of Washington 97(2): 251-262. . 1986. Studies of Psilopinae (Diptera: Ephy- dridae), I: A revision of the shore fly genus Pla- copsidella Kertész. Smithsonian Contributions to Zoology 430: iv + 30 pp. . 1992. The first shore fly of the genus Glen- anthe Haliday from the Australasian Region (Dip- tera: Ephydridae). Proceedings of the Entomolog- ical Society of Washington 94(1): 78-82. 1993. Studies of Gymnomyzinae (Diptera: Ephydridae), 1V: A revision of the shore-fly genus Hecamede Haliday. Smithsonian Contributions to Zoology 541: 46 + iii pp. . 1995a. Studies of Gymnomyzinae (Diptera: Ephydridae), [V: A revision of the shore-fly genus Glenanthe Haliday from the Western Hemisphere. Smithsonian Contributions to Zoology 567: ii + 26 pp. . 1995b. Shore flies (Diptera: Ephydridae) of the Galapagos Islands. Annals of the Entomolog- ical Society of America 88(5): 627—640. . 1997. Shore flies of the Belizean Cays (Dip- tera: Ephydridae). Smithsonian Contributions to Zoology 592: vi + 77 pp. Mathis, W. N. and T. Zatwarnicki. 1990a. A revision of the western Palearctic species of Athyroglossa (Diptera: Ephydridae). Transactions of the Amer- ican Entomological Society 116(1): 103-133. . 1990b. Taxonomic notes on Ephydridae (Dip- tera). Proceedings of the Biological Society of Washington 103(4): 891-906. . 1995. A world catalog of the shore flies (Dip- tera: Ephydridae). Memoirs on Entomology, In- ternational 4: vi + 423 pp. McAlpine, J. F 1981. Morphology and terminology- Adults. Jn McAlpine, J. FE, et al., eds. Manual of Nearctic Diptera, 1: 9-63. Ottawa. [Vol. 1 is Monograph 27 of Research Branch Agriculture Canada. | Munari, L. 1988. Il genere Homalometopus Becker: Considerazioni e ipotesi sulla sua biogeografia e filogenesi, con descrizione di una nuova specie della Sardegna (Diptera, Ephydridae). Societa Ve- neziana di Scienze Naturali - Lavori 13: 5—16. Sabrosky, C. W. 1999. Family-group names in Diptera. Myia 10: 1-360. Sturtevant, A. H. and M. R. Wheeler. 1954. Synopses of Nearctic Ephydridae (Diptera). Transactions of the American Entomological Society 79: 151— 2575 Townsend, C. H. T. 1898. Some characteristic maritime Diptera from the south end of Padre Island, and the adjacent Texas coast. I. Entomological news 9(7): 167-169. Williston, S. W. 1897. Diptera Brasiliana. Part IV. Kan- sas University Quarterly, series A 6(1): 1-12. 1908. Manual of North American Diptera. New Haven: James T. Hathaway, 3rd edition, 405 PP- Wirth, W. W. 1956. The Ephydridae (Diptera) of the Bahama _ Islands. 1817, 20 pp. . 1965. Family Ephydridae, pp. 734-759. In Stone, A., et al., eds. A Catalog of Diptera of America North of Mexico. United States Depart- American Museum Novitates VOLUME 105, NUMBER | ment of Agriculture, Agricultural Research Ser- vice, Agriculture Handbook, 276: iv + 1,696 pp. . 1968. 77, Family Ephydridae, pp. 1-43. In Papavero, N., ed. A Catalogue of the Diptera of the Americas South of the United States. S40 Pau- lo: Departamento de Zoologia, Secretaria da Agri- cultura. 49 Wirth, W. W. and A. Stone. 1956. Chapter 14. Aquatic Diptera, pp. 372-482. Jn Usinger, R. L., ed. Aquatic Insects of California. University of Cali- fornia Press, Berkeley. Zatwarnicki, T. 1992. A new classification of Ephy- dridae based on phylogenetic reconstruction (Dip- tera: Cyclorrhapha). Genus 3(2): 65-119. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 50-58 BRUMOIDES SEPTENTRIONIS DAVISI (LENG) (COLEOPTERA: COCCINELLIDAE): DISTRIBUTION, HOST-PLANT ASSOCIATIONS, AND HABITATS OF A SELDOM-COLLECTED LADY BEETLE A. G. WHEELER, JR. Department of Entomology, Clemson University, Clemson, SC 29634, U.S.A. (e-mail: awhlr@clemson.edu) Abstract.—The chilocorine coccinellid Brumoides septentrionis davisi (Leng) is an in- frequently collected lady beetle whose broad distribution in eastern North America has been mapped; few specific localities, however, have been published. Despite a known association with pines, the Pinus species on which it is found have not been recorded. Historical locality data are provided based on specimens in five museum collections, and new records from recent field work are presented. This coccinellid’s plant and habitat associations are discussed. Its populations apparently have declined in the Northeast in recent years. Certain scale insects probably serve as prey. Key Words: The most frequently encountered of all predacious beetles (Clausen 1940), cocci- nellids are associated with biological con- trol more than any other group of predators (Obrycki and Kring 1998). Coccidopha- gous lady beetles have proved more effec- tive as classical biological control agents than those that are aphidophages (Dixon 2000). The establishment, either from bio- control introductions or accidentally from commerce, of several Old World coccinel- lids that are generalist aphid predators has led to more stringent regulations for their and has prompted interest in native lady beetles whose densities have de- clined importation as those of adventive coccinellids have increased (e.g., Schaefer et al. 1987, Wheeler and Hoebeke 1995, Colunga- Garcia and Gage 1998, Cottrell and Year- gan 1998, Obrycki et al. 1998, Evans 2000). I provide distributional records of the Nearctic davisi Brumoides septentrionis Brumoides septentrionis davisi, coccinellids, insect distribution, pine barrens (Leng) and notes on its plant associations and habitats to call attention to this infre- quently collected lady beetle. I also suggest possible prey associations and a recent de- cline in its numbers. The availability of this background information might prove useful if studies on the bionomics of this cocci- nellid are initiated. Brumoides septentrionis davisi (Leng) The chilocorine Brumoides septentrionis davisi (sensu Gordon 1985) is the eastern North American subspecies of B. septen- trionis (Weise), whose other subspecies are found farther west in Canada (McNamara 1991) and the United States (Gordon 1985). Leng (1908) described the eastern subspe- cies as Exochomus (Brumus) septentrionis var. davisi from Alabama, District of Co- lumbia, “Lake Superior,” Massachusetts, Michigan, New Jersey, Pennsylvania, and Virginia; except for Lakehurst, N.J., specif- ic localities were not mentioned. The sub- VOLUME 105, NUMBER 1 genera of Exochomus that Leng (1908) rec- ognized—Arawana, Brumus, and Exocho- mus—were later considered valid genera (see Gordon 1985). Chapin (1965) proposed the new genus Brumoides, with the Old World Coccinella suturalis Fabricius designated as type, and included both B. davisi (Leng) and B. sep- tentrionis as valid species. I am retaining the generic name Brumoides for septen- trionis davisi, although Kovar (1995) re- stricted this genus to the Old World; he did not, however, propose a new generic name to accommodate the New World species. Brumoides septentrionis davisi is broadly oval, convex, 3.5 to 4.5 mm long, with the upper surface glabrous. The head and pro- notum are black and the elytra yellowish brown to red and coarsely punctured. Each elytron typically has two large black discal spots with the sutural area broadly black; the black areas often are confluent. Gordon (1985) illustrated the adult habitus and male and female genitalia. DISTRIBUTION Historical (Table 1).—State and provin- cial records that have been added since Leng’s (1908) original description are New York (Leonard 1928), Minnesota and Wis- consin (“‘Lake Superior” was cited by Leng 1908) (Wingo 1952), South Carolina (Kirk 1970), and Ontario and Quebec (McNamara 1991). In his key to species of Brumoides, Gordon (1985) gave the general range of B. septentrionis davisi as “eastern United States, New York and New Jersey west to Wisconsin,” but under its distribution he stated that the subspecies ranges from “southeastern Canada to Virginia, west to Minnesota.” Gordon (1985) did not include Alabama, listed by Leng (1908), in the known distribution. The distribution given by Belicek (1976) for B. septentrionis in- cluded Illinois and Indiana, and his map ap- pears also to include Tennessee; he did not recognize subspecies, but his records for these three states might apply to B. septen- trionis davisi. Neither Belicek (1976) nor 51 Gordon (1985) listed locality data for the specimens they examined. Downie and Ar- nett’s (1996) inclusion of Connecticut, Del- aware, Maryland, Massachusetts, New Hampshire, Ohio, Pennsylvania, Rhode Is- land, Vermont, and Virginia in the known distribution apparently is based on their in- terpretation of the stippled area on Gordon’s (1985) map. New records.—Brumoides septentrionis davisi was collected mainly during studies on mirids (Wheeler 1991; A.G.W., unpub- lished data) and other hemipterans found in northeastern pitch pine-scrub oak barrens (Wheeler 1996, 1999a, b; Wheeler and Wil- son 1996). Voucher specimens have been deposited in the Clemson University Ar- thropod Collection; Pennsylvania Depart- ment of Agriculture Collection, Harrisburg (PADA); and the National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C. (USNM). CONNECTICUT: Hartford Co., Shaker Pines, Enfield, 28 Sept. 1991; Windham Co., Windham Airport, | Sept. 1991. MAS- SACHUSETTS: Barnstable Co., North Fal- mouth, 29 May 1988; Franklin Co., Mon- tague Sand Plain, 15 June 1991. NEW HAMPSHIRE: Merrimack Co., Concord barrens, 14 Sept. 1991. NEW YORK: Suf- folk Co., Yaphank, 29 May 1982; Ulster Co., Sam’s Point Dwarf Pine Ridge Pre- serve NE of Cragsmoor, | & 14 June and 29 Sept. 1991. NORTH CAROLINA: Rockingham Co., Rt. 220, 5.6 km S of Vir- ginia state line, 3.3. km NNW of Stoneville, 9 Apr. 1979; Guilford Co., Rt. 68, 6.4 km S of Rt. 421, 5 km SE of Sandy Ridge, 10 April 1983. PENNSYLVANIA: Chester Co., Goat Hill Serpentine Barrens, SW of Nottingham, 8 July 1990; Lancaster Co., New Texas Serpentine Barrens, 7 July 1988; Luzerne Co., Milnesville, 17 Sept. 2001; Schuylkill Co., jet. Rt. 81 & Rt. 61, S of Frackville, 20 June & 6 Oct. 1991. SOUTH CAROLINA: Oconee Co., Rt. 76, 10.5 km NW of Westminster, 11 Nov. 2001. VIRGINIA: Botetourt Co., Eagle Rock, 8 May 1999; Rt. 81 Rest Area, 2.4 km N of an i) jet. Rt. 640, SE of Fincastle, 26 May & 29 July 1978, 23 May 1981, 9 June 1982, 4 Mar. & 8 Apr. 1983: Henry Co., Rt. 220, 44 km SW of Ridgeway, 14+ Mar. 1982. WEST VIRGINIA: Mercer Co., Rt. 77, 0.6 km N of Bluestone River SE of Camp Creek, 8 May 1999. PLANT ASSOCIATIONS AND HABITATS Brumoides septentrionis davisi is a con- iter inhabitant found almost exclusively on pines. Leng (1908) noted its occurrence on pines [likely pitch pine, Pinus rigida Mill.] at Lakehurst, N.J. Few specimens trom the localities listed in Table | bear host-plant data, but those from Berlin, Framingham, and Sherborn, Mass., are labeled as found on pitch pine. At least historically, pitch pine was present at nearly all the north- eastern localities trom which this coccinel- lid is known (e.g... Bromley 1935, Cryan L985, Motzkin et al. 1999). The specimen trom near Gordon, Wisc., was taken on jack pine (P. banksiana Lamb.). The adults that [ collected from the New England states, New York, and Pennsylva- nia were from pitch pine except for a spec- imen trom eastern red-cedar (Juniperus vir- giniana L.) in Lancaster County, Pennsyl- vania, Specimens trom North Carolina, South Carolina, and Virginia were from Virginia pine (P. virginiana Mill.): the West Virginia record is based on an adult trom table mountain pine (P. pungens Lamb.). In the Northeast. B. septentrionis davisi was found in pitch pine-scrub oak barrens and similar communities. It was collected in the dwarf pine plains (ridges) near Lake Maratanza in Ulster County, New York: the Montague Sand Plain in Franklin County, Massachusetts: a pitch pine community in Schuylkill County, Pennsylvania: and rem- nant pine barrens such as Concord, N.H.: Shaker Pines near Enfield, Conn.: and Yap- hank (Long Island), N-Y. It also was found on pitch pine in serpentine barrens in Ches- ter and Lancaster counties. Pennsylvania. In Luzerne County, Pennsylvania, it was found on pitch pine in a ruderal site, a coal PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON spoilbank. Collections from table mountain pine and Virginia pine in North Carolina, South Carolina, Virginia, and West Virginia also did not involve specialized communi- ties but were from disturbed sites, such as roadsides, in the Piedmont and in the Valley and Ridge ecoregions. POsSIBLE PREY ASSOCIATIONS Predation on scale insects is well known among chilocorine coccinellids (e.g., Bald- uf 1935, Majerus 1994, Hodek and Honék 1996, Dixon 2000). Within Brumoides, the Old World B. suturalis preys mainly on ar- mored scales (Diaspididae), mealybugs (Pseudococcidae), and soft scales (Cocci- dae) (e.g.. Gautam 1990, Williams and Greathead 1990, Carnegie 1997, Ponsonby and Copland 1997), with aphids and other insects serving as alternative prey (e.g. Gautam 1990). In addition, Gordon (1985) stated that members of this genus (species unspecified) feed on cochineal or dactylo- pud scales (Dactylopiidae) and mealybugs. For B. septentrionis, Belicek (1976) report- ed that the balsam woolly adelgid, Adelges piceae Ratzeburg, serves as prey, apparent- ly referring to predation by one of the west- ern subspecies of this lady beetle. No prey records for the easternmost subspecies, B. septentrionis davisi, are available, although Leng (1908) noted its occurrence at Lake- hurst, N.J., on aphid-intested pines. No definite prey records were obtained during my field work. Aphids of several genera (mainly Cinara) were observed at all sites where the coccinellid was found and might serve at least as alternative prey. All 10 adults of B. septentrionis davisi that were observed in Luzerne County, Penn- sylvania, in September 2001 were on a sin- gle pitch pine growing on a coal spoilbank with S—10 other pitch pines: the coccinellid was beaten only trom branches infested with a pine needle scale, Chionaspis het- erophyllae Cooley. In Franklin County, Massachusetts, and Schuylkill County. Pennsylvania, this lady beetle also was found on pitch pines that harbored a mar- VOLUME 105, NUMBER | garodid scale, Matsucoccus gallicolus Mor- rison. The frequent co-occurrence (ca. 5 ad- ditional sites) of the coccinellid with a more or less specialized predator of Matsucoccus scales (e.g., Lussier 1965), the anthocorid Elatophilus inimicus (Drake and Harris), suggests that M. gallicolus also was present at other collection sites for the coccinellid. DISCUSSION Brumoides septentrionis davisi is about as large as many of our familiar and well- studied coccinelline lady beetles. Moreover, its coloration—black and yellowish brown to red—is only slightly more subdued than that of black-spotted coccinellids. Its restriction to our readily recognized reddish, pines, a consistent association with special- ized communities such as pitch pine-scrub oak apparent (northeastern states) on pines in Christmas barrens, and an absence tree plantations and landscape plantings might partly explain the infrequency of its collection and lack of attention given to its bionomics. Brumoides septentrionis davist was col- lected from New Hampshire to South Car- olina at elevations from near sea level on Long Island, New York (17 m), to about 685 m in Ulster County, New York. Adults were found in all months from March to November (range: 4 Mar—11 Nov.), sug- gesting that this coccinellid is multivoltine like most Iperti 1999). In the Northeast, this lady beetle has been collected in several well-known pine barrens, including the New Jersey Pine Bar- rens and those on Cape Cod, Massachu- setts, and Long Island, New York. Despite intensive collecting on pitch pines, it was not observed in New York’s Albany Pine (e:2;, other Chilocorini Bush or in pine barrens such as Fryeburg, Shapleigh, and Waterboro in Maine and Os- sipee in New Hampshire. In addition to the field work in those northeastern pine barrens in which B. septentrionis davisi has yet to be found, the southern extent of its distribution need for more 53 requires greater resolution. Specifically, the Alabama record, listed by Leng (1908) in describing this coccinellid but not men- tioned by Gordon (1985), requires verifi- cation. It was not found in the survey of the Alabama Coccinellidae by Grimes (1965). Attempts to collect it in Alabama (and Georgia) might focus on pines in upland ar- eas, including the Appalachian Plateau. The taxonomic status of B. septentrionis davisi might also be reevaluated. Gordon (1985) indicated that he was not satisfied with his arrangement of species and sub- species of Brumoides. Following Leng’s (1908) description of B. septentrionis var. davisi, Casey (1908) reduced this variety to a synonym of Brumus septentrionis Weise. According to Leng (1911), Casey, in recon- sidering the status of davisi, thought it rep- “should be Most subse- quent authors also listed davisit as a valid resented a new species that called Brumus davisi Leng.” species, including Leonard (1928) (as Ex- ochomus davisi), Stehr (1930) (as BE. [Bru- mus] davisi), Wingo (1952) (as Brumus davisi), Chapin (1965) (as Brumoides dav- isi), and Kirk (1970) (as Brumoides davisi). Following Gordon’s (1985) monographic treatment of North American Coccinellidae, however, subsequent authors (e.g., McNa- mara 1991, Downie and Arnett 1996) have continued to list davisi as a subspecies of B. septentrionis. Another gap in our knowledge of North American Brumoides is the lack of larval descriptions. The larval key to genera (and selected species) of Nearctic coccinellids (Rees et al. 1994) excluded this genus be- cause larvae were unavailable for study. The determination of this chilocorine’s trophic habits also is needed. My sugges- tion that the diaspidid Chionaspis hetero- phyllae serves as acceptable prey is based only on the beating of 10 adults of the lady beetle from scale-infested branches of a pitch pine. The suggestion that B. o septen- trionis davisi might prey on the margarodid scale Matsucoccus gallicolus is offered be- cause of the co-occurrence of these species 54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Locality data for Brumoides septentrionis davisi from five museum collections and previously published localities (minimum of county level). Locality Year of Collection Museum or Reference United States District of Columbia Washington (Rock Creek) 1901 USNM Maryland Glen Echo 1992 USNM Massachusetts Berkley 1936 MCZ Berlin 1937 MCZ Dover 1900 MCZ Fall River NA? MCZ Framingham 1944 MCZ Marion 1902-1904 MCZ, USNM Sherborn 1934 MCZ Springfield NA MCZ Tyngsboro 1893, 1908 MCZ Michigan Whitefish Point NA USNM Minnesota Duluth 1918 Stehr 1930 Hubbard County 1929 Stehr 1930 New Hampshire Durham 1899 UNH New Jersey Browns Mill 1911 USNM Clementon 1906 USNM Jamesburg NA Smith 1910 Lakehurst 1903-1943 Leng 1908, Smith 1910; CUIC, USNM Manchester NA USNM Milltown NA Smith 1910 New York Keeseville 1952 CNC Lake Ronkonkoma 1922 CUIC Melville 1924 CUIC Mt. Whiteface 1922 USNM Riverhead 195051953 CUIC Rockaway Beach NA USNM Southold 1941, 1942 CUIC Wyandanch 1911 Davis 1911, Leonard 1928; USNM Yaphank NA Davis 1911, Leonard 1928 Pennsylvania Hazleton NA MCZ Rhode Island Lonsdale 1920 MCZ Watch Hill 1909 MCZ South Carolina Long Creek NA Kirk 1970 VOLUME 105, NUMBER 1 Table 1. Continued. a 7) Locality Year of Collection Virginia Mt. Vernon 1911, 1960 Museum’ or Reference CNC, USNM Wisconsin Gordon (4 mi. east) 1952 USNM Canada Ontario Constance Bay 1892 CNC Sudbury 1932, 1985 CNC *CNC: Canadian National Collection, Ottawa, Ont.; CUIC: Cornell University Insect Collection, Ithaca, N.Y.; MCZ: Museum of Comparative Zoology, Harvard University, Cambridge, Mass.; UNH: University of New Hampshire, Durham; USNM: National Museum of Natural History, Smithsonian Institution, Washington, D.C. *NA = Not Available. and the known coccidophagous habits of other chilocorine coccinellids. Trophic re- lationships of a coccinellid species should not be assumed merely from its syntopy with potential prey species, and experimen- tal studies on B. septentrionis davisi are needed to identify both its essential and ac- cepted or alternative prey (sensu Hodek and Honék 1996). Leng (1908) commented on the abun- dance of the coccinellid at Lakehurst in the New Jersey Pine Barrens. About 75-80% of the specimens in the USNM are from this locale (R.D. Gordon, pers. comm.). I have not found this coccinellid in the New Jersey Pine Barrens during my studies of mirids (Wheeler 1991; A.G.W., unpub- lished data) and other insects (Wheeler 1996; 1999a, b; Wheeler and Wilson 1996). In the absence of baseline data on popula- tion trends of this lady beetle, any evidence pointing to a recent decline in its densities is tenuous. My impression, though, 1s that it has become increasingly difficult to find in northeastern pine barrens. Since the early 1990s, my attempts to recollect this species have been unsuccessful at several north- eastern sites where 10 or more adults had been observed one or more times in 199]. Recent collecting was most intensive at Sam’s Point Dwarf Pine Ridge in New York (13 Aug. 2000, 5 Sept. 2001), but also in- cluded efforts to recollect B. septentrionis davisi in the pitch pine community near Frackville, Pa. (4, 6 Sept. 2001) and the Montague Sand Plain in Massachusetts (15 Aug. 1993, 6 Sept. 2001). In contrast, adults of another conifer-associated lady beetle, the coccinelline Mulsantina picta (Randall), were found at these and other sites during the 1990s and in 2000-2001. The numbers of coccinellid species, es- pecially those in crop fields, can fluctuate widely at a particular site from year to year (e.g., Foott 1974, Elliott and Kieckhefer 1990, Kieckhefer and Elliott 1990). If pop- ulations of B. septentrionis davisi actually have declined recently in pine barrens, hy- potheses to account for its reduced densities might include competition from and intra- guild predation by Old World coccinellids that have become established in North America (e.g., Evans 1991, 2000; Elliott et al. 1996; Obrycki et al. 1998). The most abundant and widespread of these adventive species are Coccinella septempunctata L. and Harmonia axyridis (Pallas). The former coccinelline spread rapidly during the 1980s (e.g., Hoebeke and Wheeler 1980, Schaefer et al. 1987, Staines et al. 1990, Gordon and Vandenberg 1991), whereas the latter species began to assume dominance in some natural and managed systems in the 1990s (e.g., Tedders and Schaefer 1994, 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Brown and Miller 1998, Colunga-Garcia and Gage 1998, Hesler et al. 2001). Al- though the establishment of H. axyridis in the eastern United States might be the result of an accidental introduction (Day et al. 1994), this predator was intentionally re- leased numerous times between 1916 and 1985, mainly to control arboreal homopter- an pests, including margarodid scales (e.g., Gordon 1985, McClure 1987). Adults and larvae of H. axyridis were found in the ma- jority of the stands of pitch pine and Vir- ginia pine that I surveyed in the late 1990s and in 2000-2001. Adults of two other ad- ventive coccinellids—Hippodamia varie- gata (Goeze) and Propylea quatuordecim- punctata (L.)—were observed less often on pines (A.G.W., unpublished data). In apple orchards of eastern West Virgin- ia, the Old World H. axyridis has become the dominant coccinelline, displacing an- other Old World species, C. septempunctata (Brown and Miller 1998). It is, however, the potential adverse effects of nonindigenous coccinellids on lady beetles (and other in- sects) native to North America that is a cause of concern (e.g., Ehler 1990, Evans 1991, Horn 1991, Elliott et al. 1993, Wheeler and Hoebeke 1995). A detrimental effect on native coccinellids such as Adalia bipunctata (L.), Coccinella novemnotata Herbst, C. transversoguttata richardsoni Brown, Cycloneda munda (Say), and Hip- podamia convergens Guérin-Méneville al- ready has been suggested (Wheeler and Hoebeke 1995, Elliott et al. 1996, Wheeler and Stoops 1996, Brown and Miller 1998, Colunga-Garcia and Gage 1998). Coccinel- lids that have more specialized habitat re- quirements than those associated with agroecosystems might be especially vulner- able to competitive displacement by adven- tive coccinellids, their numbers perhaps de- clining more rapidly than those of agricul- turally important lady beetles (Elliott et al. 1996). Any long-term monitoring of our na- tive coccinellid fauna, therefore, might in- clude not only species associated with agroecosystems but also those such as B. septentrionis davisi that are found in spe- cialized natural communities. ACKNOWLEDGMENTS For providing label data from specimens of the coccinellid | am grateful to Edward Becker (CNC), Richard Hoebeke (CUIC), David Furth (MCZ), Donald Chandler (UNH), and Robert Gordon and Natalia Vandenberg (USNM). Natalia Vandenberg additionally called my attention to and pro- vided a copy of the paper by Kovar. My appreciation also is extended to Randall Schuh (American Museum of Natural His- tory, New York) for checking for possible specimens in the AMNH, Cecil Smith (Uni- versity of Georgia, Athens) for checking for material in the UGA collection, Paul Huth (Mohonk Preserve, New Paltz, N.Y.) for fa- cilitating my access to Sam’s Point Dwarf Pine Ridge Preserve, James Stimmel (PADA) for identifying the scale insects, and Peter Adler (Clemson University) for comments that improved the manuscript. LITERATURE CITED Balduf, W. V. 1935. The Bionomics of Entomophagous Coleoptera. John S. Swift, St. Louis, 220 pp. Belicek, J. 1976. Coccinellidae of western Canada and Alaska with analyses of the transmontane zoogeo- graphic relationships between the fauna of British Columbia and Alberta (Insecta: Coleoptera: Coc- cinellidae). Quaestiones Entomologicae 12: 283— 409. Bromley, S. W. 1935. The original forest types of southern New England. Ecological Monographs 5: 61-89. Brown, M. W. and S. S. Miller. 1998. Coccinellidae (Coleoptera) in apple orchards of eastern West Virginia and the impact of invasion by Harmonia axyridis. Entomological News 109: 143-151. Carnegie, A. J. M. 1997. Sugarcane and bamboo, pp. 333-341. In Ben-Dov, Y. and C. J. Hodgson, eds. Soft Scale Insects: Their Biology, Natural Ene- mies and Control, Vol. 7B. Elsevier, Amsterdam. Casey, T. L. 1908. Notes on the Coccinellidae. Cana- dian Entomologist 40: 393-421. Chapin, E. A. 1965. The genera of the Chilocorini (Co- leoptera, Coccinellidae). Bulletin of the Museum of Comparative Zoology 133: 227-271. Clausen, C. P. 1940. Entomophagous Insects. Mc- Graw-Hill, London, 688 pp. Colunga-Garcia, M. and S. H. Gage. 1998. Arrival, VOLUME 105, NUMBER 1 establishment, and habitat use of the multicolored Asian lady beetle (Coleoptera: Coccinellidae) in a Michigan landscape. Environmental Entomology 27: 1574-1580. Cottrell, T. E. and K. V. Yeargan. 1998. Intraguild pre- dation between an introduced lady beetle, Har- monia axyridis (Coleoptera: Coccinellidae), and a native lady beetle, Coleomegilla maculata (Cole- optera: Coccinellidae). Journal of the Kansas En- tomological Society 71: 159-163. Cryan, J. EF 1985. Retreat in the barrens. Defenders 60(1): 18-29. Day, W. H., D. R. Prokrym, D. R. Ellis, and R. J. Chianese. 1994. The known distribution of the predator Propylea quatuordecimpunctata (Cole- optera: Coccinellidae) in the United States, and thoughts on the origin of this species and five oth- er exotic lady beetles in eastern North America. Entomological News 105: 244-256. Dixon, A. F G. 2000. Insect Predator—Prey Dynamics: Ladybird Beetles and Biological Control. Cam- bridge University Press, Cambridge, 257 pp. Downie, N. M. and R. H. Arnett. Jr. 1996. The Beetles of Northeastern North America, Vol. II: Polypha- ga: Series Bostrichiformia through Curculionoi- dea. Sandhill Crane Press, Gainesville, Florida. pp. 891-1721. Ehler, L. E. 1990. Environmental impact of introduced biological-control agents: Implications for agri- cultural biotechnology, pp. 85—96. /n Marois, J. J. and G. Bruening, eds. Risk Assessment in Agri- cultural Biotechnology: Proceedings of the Inter- national Conference. University of California Di- vision of Agriculture and Natural Resources Pub- lication 1928. Elliott, N. C. and R. W. Kieckhefer. 1990, Dynamics of aphidophagous coccinellid assemblages in small grain fields in eastern South Dakota. Envi- ronmental Entomology 19: 1320-1329. Elliott, N. C., R. W. Kieckhefer, and W. C. Kauffman. 1993. Effects of an invading coccinellid, Cocci- nella septempunctata L., on the structure of native coccinellid species assemblages in agricultural crops. Bulletin of the Ecological Society of Amer- ica 74(2 suppl.): 226. Elliott, N., R. Kieckhefer, and W. Kauffman. 1996. Ef- fects of an invading coccinellid on native cocci- nellids in an agricultural landscape. Oecologia 105: 537-544. Evans, E. W. 1991. Intra versus interspecific interac- tions of ladybeetles (Coleoptera: Coccinellidae) attacking aphids. Oecologia 87: 401—408. . 2000. Morphology of invasion: Body size pat- terns associated with establishment of Coccinella septempunctata (Coleoptera: Coccinellidae) in western North America. European Journal of En- tomology 97: 469-474. Foott, W. H. 1974. Observations on Coccinellidae in nN ~ corn fields in Essex County, Ontario. Proceedings of the Entomological Society of Ontario 104: 16— Pile Gautam, R. D. 1990. Mass-multiplication techniques of coccinellid predator, lady bird beetle (Brumoi- des suturalis). Indian Journal of Agricultural Sci- ences 60: 747-750. Gordon, R. D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. Journal of the New York Entomological Society 93: 1-912. Gordon, R. D. and N. Vandenberg. 1991. Field guide to recently introduced species of Coccinellidae (Coleoptera) in North America, with a revised key to North American genera of Coccinellini. Pro- ceedings of the Entomological Society of Wash- ington 93: 845-864. Grimes, W. H. 1965. The Coccinellidae (Coleoptera) of Alabama, Ph.D. Dissertation, Auburn Univer- sity, Auburn, Alabama, 144 pp. Hesler, L. S.. R. W. Kieckhefer, and D. A. Beck. 2001. First record of Harmonia axyridis (Coleoptera: Coccinellidae) in South Dakota and notes on its activity there and in Minnesota. Entomological News 112: 264-270. Hodek, I. and A. Honek. 1996. Ecology of Coccinel- lidae. Kluwer Academic Publishers, Dordrecht, 464 pp. Hoebeke, E. R. and A. G. Wheeler, Jr. 1980. New dis- tribution records of Coccinella septempunctata L. in the eastern United States (Coleoptera: Cocci- nellidae). Coleopterists Bulletin 34: 209-212. Horn, D. J. 1991. Potential impact of Coccinella sep- tempunctata on endangered Lycaenidae (Lepidop- tera) in northwestern Ohio, USA, pp. 159-162. Jn Polgar, L., R. J. Chambers, A. F G. Dixon, and I. Hodek, eds. Behaviour and Impact of Aphidopha- ga. SPB Academic Publishing, The Hague. Iperti, G. 1999. Biodiversity of predaceous Coccinel- lidae in relation to bioindication and economic im- portance. Agriculture, Ecosystems and Environ- ment 74: 323-342. Kieckhefer, R. W. and N. C. Elliott. 1990. A 13-year survey of the aphidophagous Coccinellidae in maize fields in eastern South Dakota. Canadian Entomologist 122: 579-581. Kirk, V. M. 1970. A list of the beetles of South Car- olina. Part 2—Mountain, Piedmont, and Southern Coastal Plain. South Carolina Agricultural Exper- iment Station Technical Bulletin 1038, 117 pp. Kovar, I. 1995. Revision of the genera Brumus Muls. and Exochomus Redtb. (Coleoptera, Coccinelli- dae) of the Palaearctic Region. Part I. Acta En- tomologica Musei Nationalis Pragae 44: 5—124. Leng, C. W. 1908. Notes on Coccinellidae.—III. Jour- nal of the New York Entomological Society 16: 33-44. Leng, [C. W.]. 1911. [Proceedings of the Society, May 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 17, 1910]. Journal of the New York Entomologi- cal Society 19: 203. Leonard, M. D., ed. 1928. A list of the insects of New York with a list of the spiders and certain other allied groups. Cornell University Agricultural Ex- periment Station Memoir 101, 1121 pp. Lussier, S. J. 1965. A study of Elatophilus inimica D. & H. (Hemiptera: Anthocoridae) and its role in the natural control of the red-pine scale Matsu- coccus resinosae B. & G. (Homoptera: Margarod- idae), M.S. Thesis, University of Massachusetts, Amherst, 42 pp. McClure, M. S. 1987. Potential of the Asian predator, Harmonia axyridis Pallas (Coleoptera: Coccinel- lidae), to control Matsucoccus resinosae Bean and Godwin (Homoptera: Margarodidae) in the United States. Environmental Entomology 16: 224-230. McNamara, J. 1991. Family Coccinellidae, ladybird beetles, pp. 229-237. In Bousquet, Y., ed. Check- list of Beetles of Canada and Alaska. Research Branch Agriculture Canada Publication 1861/E. Majerus, M. E. N. 1994. Ladybirds. HarperCollins, London, 367 pp. Motzkin, G., W. A. Patterson HI, and D. R. Foster. 1999. A historical perspective on pitch pine-scrub oak communities in the Connecticut Valley of Massachusetts. Ecosystems 2: 255-273. Obrycki, J. J. and T. T. Kring. 1998. Predaceous Coc- cinellidae in biological control. Annual Review of Entomology 43: 295-321. Obrycki, J. J., K. L. Giles, and A. M. Ormord. 1998. Interactions between an introduced and indige- nous coccinellid species at different prey densi- ties. Oecologia 117: 279-285. Ponsonby, D. J. and M. J. W. Copland. 1997. Preda- tors. Coccinellidae and other Coleoptera, pp. 29— 60. In Ben-Dovy, Y. and C. J. Hodgson, eds., Soft Scale Insects: Their Biology, Natural Enemies and Control, Vol. 7B. Elsevier, Amsterdam. Rees, B. E., D. M. Anderson, D. Bouk, and R. D. Gordon. 1994. Larval key to genera and selected species of North American Coccinellidae (Cole- optera). Proceedings of the Entomological Society of Washington 96: 387-412. Schaefer, P. W., R. J. Dysart, and H. B. Specht. 1987. North American distribution of Coccinella sep- tempunctata (Coleoptera: Coccinellidae) and its mass appearance in coastal Delaware. Environ- mental Entomology 16: 368-373. Staines, C. L., Jr, M. J. Rothschild, and R. B. Trum- bule. 1990. A survey of the Coccinellidae (Cole- optera) associated with nursery stock in Mary- land. Proceedings of the Entomological Society of Washington 92: 310-313. Stehr, W. C. 1930. The Coccinellidae (ladybird beetles) of Minnesota (Coleoptera). University of Minne- sota Agricultural Experiment Station Technical Bulletin 75, 54 pp. Tedders, W. L. and P. W. Schaefer. 1994. Release and establishment of Harmonia axyridis (Coleoptera: Coccinellidae) in the southeastern United States. Entomological News 105: 228-243. Wheeler, A. G., Jr. 1991. Plant bugs of Quercus ilici- folia: Myriads of mirids (Heteroptera) in pitch pine-scrub oak barrens. Journal of the New York Entomological Society 99: 405-440. 1996. Eremocoris depressus Barber: Hosts, seasonality, and first New England records of a pine seed specialist (Heteroptera: Lygaeidae). Pro- ceedings of the Entomological Society of Wash- ington 98: 767-773. 1999a. Scaphytopius angustatus (Osborn) (Homoptera: Cicadellidae), a leafhopper charac- teristic of pitch pine-scrub oak barrens. Proceed- ings of the Entomological Society of Washington 101: 219-220. . 1999b. Phoenicocoris claricornis and Pino- phylus carneolus (Hemiptera: Miridae): Distribu- tion and seasonality of two specialists on micro- sporangiate strobili of pines. Journal of the New York Entomological Society 107: 238-246. Wheeler, A. G., Jr. and E. R. Hoebeke. 1995. Cocci- nella novemnotata in northeastern North America: Historical occurrence and current status. Proceed- ings of the Entomological Society of Washington 97: 701-716. Wheeler, A. G., Jr. and C. A. Stoops. 1996. Status and spread of the Palearctic lady beetles Hippodamia variegata and Propylea quatuordecimpunctata (Coleoptera: Coccinellidae) in Pennsylvania, 1993-1995. Entomological News 107: 291-298. Wheeler, A. G., Jr. and S. W. Wilson. 1996. Planthop- pers of pitch pine and scrub oak in pine barrens communities (Homoptera: Fulgoroidea). Proceed- ings of the Entomological Society of Washington 98: 100-108. Williams, J. R. and D. J. Greathead. 1990. Sugar cane, pp. 563-578. Jn Rosen, D., ed. Armored Scale In- sects: Their Biology, Natural Enemies and Con- trol, Vol. 4B. Elsevier, Amsterdam. Wingo, C. W. 1952. The Coccinellidae (Coleoptera) of the Upper Mississippi Basin. Iowa State College Journal of Science 27: 15-53. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 59-65 PROBOSCIDOTYLUS NIGROSQUAMIS (MALDONADO) (HETEROPTERA: MIRIDAE: ORTHOTYLINAE): NEW COMBINATION AND FIRST RECORD FOR THE UNITED STATES THOMAS J. HENRY Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, Washington, DC 20506-0168, U.S.A. (e-mail: thenry @sel.barc.usda.gov) Abstract.—Parthenicus nigrosquamis Maldonado, described and previously known only from Puerto Rico, is reported for the first time in the United States from Key West, Florida. Based on head morphology, vestiture, and male genital structures, this species is removed from Parthenicus Reuter and transferred to the monotypic genus Proboscidotylus Henry (n. comb.), previously known only from Mexico. The type species of Probosci- dotylus, P. carvalhoi Henry, and P. nigrosquamis are redescribed and their relationships are discussed. Scanning electron micrographs of selected structures, dorsal and lateral photographs, and illustrations of male genitalia for both species are provided to help facilitate recognition. Key Words: Insecta, Heteroptera, Miridae, Orthotylinae, new combination, descriptions, distribution dorsal and lateral photographs, and illustra- tions of male genitalia. Also given are an While studying material housed in the Florida State Collection of Arthropods in Gainesville, I discovered four specimens of Parthenicus nigrosquamis Maldonado from Key West, Florida, that represent a new re- cord for the United States. Maldonado (1969), in describing P. nigrosquamis from Puerto Rico based on seven specimens, in- dicated that this species might not be con- generic with other species of Parthenicus, but refrained from erecting a new genus. My study of external characters and male genitalia indicates that P. nigrosquamis is congeneric with Proboscidotylus carvalhoi Henry described from Veracruz, Mexico (Henry 1995). In this paper, I transfer Parthenicus ni- grosquamis to the genus Proboscidotylus, redescribe P. carvalhoi and P. nigrosquam- is, and provide for both species scanning electron micrographs of selected structures, identification key, a discussion of relation- ships, and new records of P. nigrosquamis from Puerto Rico and the United States. Acronyms for collections cited in this pa- per are as follows: FSCA (Florida State Collection of Arthropods, Florida Depart- ment of Agriculture, Gainesville); UNAM (Instituto de Biologia, Universidad Nacion- al Autonoma de Mexico, Mexico, DF); USNM ([{United States] National Museum of Natural History, Smithsonian Institution, Washington, DC). Proboscidotylus Henry Proboscidotylus Henry 1995: 340. Type species: Proboscidotylus carvalhoi Hen- ry 1995. Original designation. Diagnosis.—Members of this genus may be separated from all other Orthotylinae by 60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1 4 Figs. 1-3. 2; arrow indicating swollen frons. 3, setigerous tubercle on genital capsule. the strongly swollen frons extending well in front of the eyes in both sexes (Figs. 4, 6, 12), particularly in females; prominent tylus, sometimes extended into an elongate tubercle (Figs. 3—5); relatively stout anten- nal segment I; uniformly cylindrical anten- nal segment II, sometimes slightly more thickened basally in males; erect simple se- tae on the dorsum, intermixed with black scalelike setae on the pronotum and hem- elytra, with a small, more dense, patch of black scalelike setae on each paracuneus (Figs. 1, 10); and a cluster of stout, apically swollen, bristlelike and, possibly, glandular setae on the ventral surface of the genital capsule (Figs. 8-9, 15-17). Male genitalia: Vesica simple, secondary gonopore typical with indistinct horseshoe-shaped opening, bearing one slender or clavate spiculum (Figs. 19, 22); left paramere L-shaped (Figs. 18, 21); right paramere simple, elon- gate to elongate oval (Figs. 20, 23). Discussion.—The relationship of Pro- boscidotylus to other orthotylines is not ful- ly known. Henry (1995) indicated that it belonged in the nominate tribe Orthotylini, somewhere near the Zanchius group as de- fined by Schuh (1974), and noted that only Proboscidotylus carvalhoi. 1, Dorsal aspect, 2 Lateral aspect, ¢; arrow a indicating extended tylus; arrow b indicating ; arrow indicating cuneal patch. 2, Lateral aspect, a few other orthotyline genera bear black, scalelike, dorsal setae, including Brooksetta Kelton, //nacora Reuter, Knight, Macrotyloides Van Duzee, and Parthenicus Reuter. Synapomorphies dis- tinguishing Proboscidotylus from these and Ilnacorella other orthotylines possessing black, scale- like, dorsal setae are the strongly swollen frons, the uniformly cylindrical second, and the cluster setae on the ventral surface of the male genital capsule. o KEY TO THE SPECIES OF PROBOSCIDOTYLUS 1. Length of male 2.68—2.84 mm, female 2.64— 3.04 mm; tylus of male extended into an elon- gate, apically setigerous tubercle (Figs. 3—5); antennal segment Il of male and female much longer than the basal width of the pronotum, at least by the dorsal width of an eye; Mexico (Veracruz) carvalhoi Henry Length of male 2.35—2.55 mm, female 2.40— 2.65 mm; tylus of male not extended into an yy) elongate tubercle (Figs. 11-12); antennal seg- ment II of male and female only slightly longer than the basal width of the pronotum, much less than by the dorsal width of an eye: Puerto Rico and United States (Florida) a ees Rees) nigrosquamis (Maldonado) VOLUME 105, NUMBER | Figs. 4-9. aspect of 6l —’ Scanning electron micrographs of Probosidotylus carvalhoi (after Henry 1995). 4, Head, lateral § (150%); arrow indicating extended tylus. 5, Extended tubercle on tylus of d (260). 6, Head, lateral aspect of 2 (151); arrow indicating swollen frons. 7, Scalelike setae on hemelytra (1,610). 8, Genital capsule, caudal aspect (426%); arrow indicating setal cluster. 9, Genitala capsule, lateral aspect (447); arrow indicating setal cluster. Proboscidotylus carvalhoi Henry (Figs. 1-9, 18-20) Proboscidotylus carvalhoi Henry 1995: 344 (n. sp.). Diagnosis.—Separated from P. nigros- quamis by the larger size, more strongly swollen frons, elongate tubercle on the tylus found only in the male, and the proportion- ately longer second antennal segment. Description (modified after Henry 1995).—Male (n = 4): Length 2.68—2.84 mm, width 1.20—1.26 mm across hemelytra). Overall coloration pale (widest area brownish yellow. Dorsum clothed with scattered erect, simple setae, intermixed with slender, black, scalelike setae (Fig. 7) on hemelytron and similar scattered scale- like setae on posterior half of pronotum (more dense transverse patches as in P. ni- grosquamis possibly rubbed away); each paracuneus with a small, dense patch of 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1 ae Figs. lO-11. aspect. black scalelike setae. Head: Width 0.64— 0.66 mm, 0.24-—0.28 mm; strongly swollen (Pig. 4); tylus extended vertex frons into an elongate, apically setigerous tuber- cle (Figs. 4—5), length 0.10 mm. Rostrum: Length 0.74-—0.78 mm, extending nearly to bases of metacoxae. Antenna: Segment I, length 0.28—0.30 mm; Il, 1.20—1.24 mm; II, O.56-0.64 mm; TV, 0.34-0.40 mm. Pronotum: Length 0.28-0.30 mm, basal 0.84-0.86 mm. Genitalia: capsule (Figs. 8-9); left paramere (Fig. 18); 19); right width Genital slender vesical spiculum (Fig. paramere (Fig. 20). Female (n = 8): Length 2.64—3.04 mm, width 1.24—1.28 mm. Head: Width 0.60 0.62 mm, vertex 0.28—0.30 mm. Rostrum: Length 0.80—-0.86 mm. Antenna: Segment I, length 0.30—0.32 mm; II, 1.12—1.20 mm; Il, 0.74-0.78 mm; IV, 0.40-—0.46 mm. Pronotum; O.28—0.32 basal width 0.84—0.88 mm. Similar to male in overall coloration and mm, 11 Proboscidotylus nigrosquamis (d ). 10, Dorsal aspect; arrow indicating cuneal patch. 11, Lateral pubescence, but differs by the broader form (Figs. 1-2), more strongly swollen frons (Fig. 6), and tylus lacking an elongate tu- bercle. Host.—Unknown. Distribution.—Known only from Vera- cruz, Mexico. Specimens examined.—MEXICO: holo- type ¢ and 4 6, 8 & paratypes, Veracruz, UNAM Los Tuxlas Estac. Biol., N. Cate- maco, 16-19 Sept. 1989, E. Barrera, T. J. Henry, & I. M. Kerzhner, taken at light (1 3, 3 2) UNAM:;) holotype, 3'6,>5> 2 USNM). Proboscidotylus nigrosquamis (Maldonado), new combination (Figs. 10-17, 21-23) Parthenicus nigrosquamis Maldonado 1969: 66 (n. sp.); Schuh 1995: 179 (cat.). Diagnosis.—Separated from P. carvalhoi by the smaller size, less strongly swollen VOLUME 105, NUMBER | Figs. 12-17. (151). ve crt ss eh Pas (to Bits Nn cir SP Scanning electron micrographs of Probosidotylus nigrosquamis (< ). 13, Patch of scalelike setae on cuneus (1,710). 63 a 31 ama. ¢ 12, Head, lateral aspect 14, Claw (1,300). 15, Genital capsule, lateral aspect (895); arrow indicating setal cluster. 16, Genital capsule, caudal aspect (530%); arrow indicating setal cluster. 17, Highly magnified bristlelike setae on genital capsule (1,270%). frons, absence of a tubercle on the tylus of the male, and the proportionately shorter antennal II that is only slightly longer than the width of the pronotum by a length much less than the dorsal width of an eye. Description.—Male (n = 5): Overall col- oration uniformly green to yellowish green. Dorsum clothed with scattered erect, simple setae, intermixed with slender, black, scale- like setae (Figs. 10, 13) on hemelytron and posterior half of pronotum; black scalelike setae on pronotum thicker and clustered into 5 or 6 more dense, transverse patches; each paracuneus with a dense patch of black scalelike setae. Length 2.35—2.55 mm, width 1.01-1.09 mm (widest area across hemelytra). Head: Width 0.57—0.59 mm, vertex 0.20—0.21 mm; frons swollen, tylus prominent (Fig. 12), but not extended into a long tubercle and without apical clus- ter of long setae. Rostrum: Length 0.74— 0.78 mm, extending nearly to bases of me- tacoxae (extends past metacoxae to base of abdomen in curled or more teneral speci- 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 18 Figs. 18-23. vd | Male genitalia of Proboscidotylus spp. 18-20, P. carvalhoi (after Henry 1995): 18, Left par- amere. 19, Vesical spiculum. 20, Right paramere. 21—23, P. nigrosquamis: 21, Left paramere. 22, Vesical spic- ulum. 23, Right paramere. mens). Antenna: Segment I, length 0.20— 0.21 mm; II, 0.94—0.96 mm; III, 0.35—0.38 mm; IV, 0.22—0.29 mm. Pronotum: Length 0.27—0.29 mm, basal width 0.91—0.82 mm. Legs: Slender, concolorous with body; tib- ial spines pale, indistinct; claws slender, parempodia fleshy, typically convergent apically (Fig. 14). Genitalia: Genital cap- sule (Figs. 15-17); left paramere (Fig. 21); apically clavate vesical spiculum scalloped along dorsal edge (Fig. 22); right paramere (Fig. 23): Female (n = 5): Length 2.40—2.65 mm, width 1.12—1.14 mm. Head: Width 0.52— 0.57 mm, vertex 0.26—0.26 mm; frons more strongly swollen than in male. Rostrum: Length 0.78—0.79 mm, extending nearly to metacoxae (extending past metacoxae near- ly to base of Ovipositor in curled or more teneral specimens). Antenna: Segment I, length 0.21—0.22 mm; II, 0.85—0.86 mm; II, 0.36-0.40 mm; IV, 0.22-0.26 mm. Pronotum: Length 0.23—0.25 mm, _ basal width 0.79—0.81 mm. Similar to male in color and pubescence, but differs by the more strongly swollen frons and slightly broader hemelytra. Host.—Unknown. Distribution.—Previously known only from Puerto Rico. Now newly recorded from the United States (Florida). Discussion.—Although this species lacks the extended tylus that is characteristic of P. carvalhoi males, it shares all other char- acters that define the genus, including the swollen frons in both sexes, the stout an- tennal segment I, the uniformly cylindrical antennal segment II, the black scalelike se- tae on the dorsum (including the cuneal patch), and the male genital capsule with a field of stout setae on the ventral surface. Specimens examined.—PUERTO RICO: Holotype d, allotype 2 (and 3 badly dam- aged paratypes), Mayaguez, Julio 1961, J. Maldonado C. (USNM); 2 6, 1 &, EI Verde, June 1967, J. Maldonado C. (USNM); 2 6, Caribbean N. F, El Verde Field Sta., 8-14 May 1985, at UV light, 300 mel. CULE 198s Exp! ES Ro iHoebeke: J. K. Liebherr, & S. W. Nichols (USNM; additional material in Cornell University collection); 3 d, 5 2, El Verde Field Sta- tion, 12 June 1990, A. G. Wheeler, Jr., taken at UV light (USNM). UNITED STATES: 3 S, 1 2, Florida, Monroe Co., Flemming Key [man-made key off north edge of Key West], 23 January 1979, 19 & 21-25 De- cember 1979, 27 January 1980, John A. Acree & H. V. Weems, Jr., insect flight trap (FSCA, 1 6 USNM). ACKNOWLEDGMENTS I thank Julieta Brambila and Susan Hal- bert (FSCA) for lending specimens repre- senting the first U.S. record of P. nigros- quamis, and A. G. Wheeler, Jr. (Clemson University, Clemson, SC) for the series of this species he collected in Puerto Rico. I VOLUME 105, NUMBER | also extend thanks to Michele Touchet (Systematic Entomology Laboratory [SEL], ARS, USDA, % USNM) for the adult dig- ital photographs, and to the late Richard C. Froeschner (USNM), David R. Smith (SEL), and E. E. Grissell (SEL) for review- ing the manuscript and offering suggestions for its improvement. LITERATURE CITED Henry, T. J. 1995. Proboscidotylus carvalhoi, a new genus and species of sexually dimorphic plant bug from Mexico (Heteroptera: Miridae: Orthotyli- 65 nae). Proceedings of the Entomological Society of Washington 97: 340-345. Maldonado, J. Capriles. 1969. The Miridae of Puerto Rico (Insecta, Hemiptera). University of Puerto Rico Agricultural Experiment Station Technical Paper 45: 1-133. Schuh, R. T. 1974. The Orthotylinae and Phylinae (He- miptera: Miridae) of South Africa with a phylo- genetic analysis of the ant-mimetic tribes of the two subfamilies for the world. Entomologica Americana 47: 1-322. . 1995. Plant bugs of the world (Insecta: Het- eroptera: Miridae). Systematic catalog, distribu- tions, host list. and bibliography. The New York Entomological Society, New York, 1,329 pp. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 66-69 PEORIA INSULARIS, A NEW SPECIES OF PEORIINI (LEPIDOPTERA: PYRALIDAE: PHYCITINAE) FROM MISSISSIPPI AND LOUISIANA JAY C. SHAFFER Department of Biology 3E1, George Mason University, Fairfax, Wa 22030-4444, U.S.A. (e-mail: jshaffe | @gmu.edu) Abstract.—Peoria insularis is described from four male specimens as a new species of pyralid moth from Mississippi and Louisiana. It is distinguished as the only North American species of Peoria with both a transverse posterior forewing band and prominent white markings on the forewing veins. Photographs of the adult moth and genitalia are included. Range extensions are given for two Peoria species. Peoria floridella Shafter, previously known only from the east coast of Florida is recorded from coastal dunes of North Carolina and Alabama, and P. punctata Shaffer, described from a single locality in Texas is now recorded from additional localities there and from Louisiana. Key Words: A perusal of an excellent series of spec- imens of Peortini from the Mississippi En- tomological Museum, Mississippi State University, turned up four specimens of a new species of Peoria as well as some note- worthy range extensions for P. floridella Shaffer 1968 and P. punctata Shaffer 1976. MATERIALS AND METHODS Four genitalia slides and one right fore- wing slide (the right hind wing was missing from the specimen) were prepared from the four specimens. All were mounted in Eu- paral. Moths and slides were examined us- ing a Wild MS5 stereo microscope; slides were also examined at high power with a Nikon L-Ke microscope. Genitalia photo- graphs were taken with an Olympus Vanox AHBT3 microscope. Color names follow the ISCC-NBS Color-Name Charts (Kelly 1965) except for small structures where only general designations could be given. Moths and color samples were viewed to- gether using the Wild microscope and a Peoria floridella, Peoria punctata fluorescent ring light. The holotype is de- posited in the National Museum of Natural History, Smithsonian Institution, and the three paratypes are in the collection of the Mississippi Entomological Museum. Peoria insularis Shaffer, new species (Figs. 1-5) Diagnosis.—Among North American species of Peoria the presence of a trans- verse posterior forewing band is possessed only by P. insularis, P. longipalpella (Ra- gonot), and P. punctata. P. longipalpella is devoid of the white markings on the fore- wing veins exhibited by P. insularis and in varying degree by several other Peoria spe- cies. These white vein markings are rather indistinct in P. punctata, a species which is unique within Peoria in that its uncus arms are of very unequal length, the dorsal arm only about one third as long as the ventral one (see Shaffer, 1976, Fig. 10). The dark central island on the forewing cell (Fig. 1) VOLUME 105, NUMBER | 67 Figs. 1-2. (1), 1 mm (2). is unique to P. insularis, though it may be difficult to discern in worn specimens. The genitalia provide no unique apomor- phies or combination of characters distin- guishing this species. Indeed, most Peoria genitalia are rather uniform with the long, slender, digitate medial process of the gna- thos being shared with eight other species and the single well developed cornutus with nine other species. Description.—Male: Head: Frons coni- cal, brown dorsally, lighter laterally. Labial palpus (Fig. 2) porrect, 4% as long as eye diameter; basal segment white, second brown on dorsal half, light brown at base and on ventral half, third brown. Maxillary palpus cylindrical, not quite extending to tip of frons. Antenna shaft subserrate, scaling light brown. Ocellus normal, black with clear center. Forewing (Fig. 1): Length 8.5-10.0 mm (* = 9.25; n = 4). R, well stalked with R,,;, from just before upper outer angle; M, from the angle; M,,, stalked about half its length, from lower outer angle of cell, well separate from CuA,. Costa dark brown basally along wing margin, elsewhere white; subcosta and ra- dius broadly white, with narrow bands of yellowish pink between costa and subcosta and between subcosta and radius. Cell broadly outlined with white (on anterior Peoria insularis, holotype. 1, Adult showing left wings. 2, Head, right side. Scale bar = 2 mm margin, posterior margin, and closing vein) leaving island of brownish gray in center of cell. Subcosta and radial and medial veins beyond cell narrowly marked by white. Cubital veins beyond cell indistinctly marked with white. Ground rather uniform- ly brownish gray posterior to cell and CuA,. LA+2A indistinctly marked with white. Hind wing: M,., fused, stalked with CuA, about half length of latter, from lower outer angle of cell; CuA, from very near the an- gle. Genitalia (Figs. 3-5): Medial process of uncus a well sclerotized somewhat nar- row band of uniform width, laterally bear- ing numerous, minute, irregular, rounded more protuberances; lateral arms of equal or nearly equal length. Gnathos with long, nar- row, digitate median process; lateral arms broad, quite flat. Juxta scoop-like, devoid of setae. Vinculum broadly rounded, some- what flattened along anterior margin. Valve with costa terminating in small tooth. Ae- deagus somewhat irregular; vesica with a small patch of minute triangular teeth; 2 cornuti (Fig. 5), one cornutus distinct, rounded and bearing 3 or 4 somewhat slen- der teeth, well sclerotized only on side bearing teeth, second cornutus (Fig. 5, ar- row) indistinct, variably developed, lacking teeth. Holotype.—<¢: Mississippi, Franklin Co., 68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs Porter! €reekoeTSNy RAE SECS NWi29 June 1992, T. Schiefer & R. Fontenot; 3 genitalia on slide 2720 J. C. Shaffer. With the permission of Richard Brown, the ho- lotype is deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC. Paratypes.—d, Mississippi, Grenada Co., T22N, R3E, Sec. 31 NW, 7=13 Aug. 1991, R. L. Brown, d genitalia on slide 2732, wing and antenna on slide 2756, J. C. Shaffer; ¢, Louisiana, Bossier Parish, Barksdale A.FB., 32°30'42”N, 93°32'42’W, 26 June 1996, D. M. Pollock, Blacklight in shortleaf pine forest, d genitalia on slide 2731 J. C. Shaffer; ¢, Barksdale A.FB., 32°31'13”"N, 93°35'46"W, 24 August 1996, D. M. Pollock, Blacklight in calcareous prairie, d genitalia on slide 2730 J. C. Shaf- fer. All deposited in the Mississippi Ento- mological Museum, Mississippi State Uni- versity. Etymology.—The specific epithet is de- rived from the Latin insula (island) in ref- erence to the isolated dark patch in the fore- wing cell. Discussion.—Table | (Shaffer 1968, p. 1s. 3-5. Peoria insularis, male genitalia, holotype. 3, Genitalia, aedeagus removed. 4, Aedeagus. 5, Vesica, enlarged and rotated 90° clockwise. Scale bar = 0.5 mm (3, 4), 0.1 mm (5). 12) compares 12 sets of characters for known North American species of Peoria. For P. insularis the symbols 0, x, X, 0, Xx, X, X, p, X, Ss, a, | may be added to columns 1 through 12 respectively for that table, up- dated copies of which are available from the author. Peoria insularis brings the total of described North American species of Pe- oria to 16, 13 of which were covered in Shaffer (1968). Peoria punctata Shaffer (1976), and P. padreella A. Blanchard (1980) were described more recently. In ad- dition, there are 4 known Neotropical Pe- oria species, bringing the grand total of Pe- oria to 20. The genus is limited to the West- ern Hemisphere. RANGE EXTENSIONS FOR PEORIA FLORIDELLA AND PEORIA PUNCTATA In the Mississippi State material there are ten specimens of P. floridella, a species pre- viously known only from the east coast of Florida, ranging from Summer Haven (St. Johns Co.) south to Vero Beach (Indian River Co.). These ten specimens consist of two males from Ft. Macon State Park in North Carolina (Carteret Co.) and eight VOLUME 105, NUMBER | specimens (2 6, 6 2) from Bon Secour Na- tional Wildlife Refuge (Baldwin Co.) Ala- bama. The North Carolina locality is on a barrier island (Bogue Bank), the Alabama locality on a coastal peninsula. J.B. Sullivan (personal communication) reports the spe- cies as common in the coastal dunes of North Carolina, but apparently absent from similar inland habitats. Three male specimens of P. punctata from the same collection are of interest be- cause the species was previously known from only two specimens, the male holo- type from Conroe, Texas (Montgomery Co., collected 13 May 1970), and a second male collected by the author at the Welder Wild- life Refuge near Sinton, Texas (San Patricio Co., 22 May 1977; record not previously published). Of the three Mississippi State specimens, two are also from Texas (13.5 miles east of Seguin, Guadalupe Co.; both 8 May 1993), and the other from Louisiana (6 miles east southeast of Buhler, Calcasieu Par.; 14 June 1993). ACKNOWLEDGMENTS I thank Richard Brown of the Mississippi Entomological Museum for calling this ma- 69 terial to my attention, arranging the loan, and reviewing a draft of the manuscript. Specimens were collected with grants and contracts to Richard Brown and Terence Schiefer from the National Science Foun- dation (BSR-9024810), the Mississippi Natural Heritage Program, and The Nature Conservancy, Louisiana Field Office. | am grateful to J. B. Sullivan for providing ad- ditional information of P. floridella. LITERATURE CITED Blanchard, A. 1980. A new species of the Genus Pe- oria Ragonot (Pyralidae). Journal of the Lepidop- terists’ Society 34(4): 338-339. Kelly, K. L. 1965. ISCC-NBS color-name charts illus- trated with centroid colors. Standard Sample No. 2106. Supplement to National Bureau of Stan- dards Circular 553. 4 pp. (unnumbered), 18 sheets color samples. U.S. Government Printing Office. Washington, D.C. Shaffer, J. C. 1968. A Revision of the Peoriinae and America North of Mexico (Lepidoptera: Pyralidae). Bulletin of the United States National Museum 280: 1-124. . 1976. Two new species of Peoriinae (Lepi- doptera: Pyralidae) from Texas. Proceedings of the Entomological Society of Washington 78(4): 431— 434. Anerastiinae (Auctorum) of PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 70-72 A NEW DACTYLOLABIS OSTEN SACKEN (DIPTERA: TIPULIDAE) FROM NORTH CAROLINA GEORGE W. BYERS AND DOUGLAS A. ROSSMAN (GWB) Snow Entomological Division, Natural History Museum, 1460 Jayhawk Bou- levard, University of Kansas, Lawrence, Kansas 66045-7523; (DAR) Research Associate, Department of Biology, Luther College, Decorah, lowa 52101; Curator Emeritus, Museum of Natural Science, Louisiana State University, Baton Rouge, Louisiana 70803. Abstract.—The male of Dactylolabis (D.) sohiyi, new species, from North Carolina is described and illustrated. It is compared with D. pemetica Alexander, the only other American species of Dactylolabis with a greatly enlarged stigmal area on the wing. Key Words: While collecting Tipulidae in western North Carolina in the spring of 1999, one of us (DAR) found an unusually large Dac- tylolabis Osten Sacken with distinctive wings. A search for this remarkable fly in the spring of 2001 yielded no further spec- imens. We therefore describe it here in the hope that other entomologists in the region will learn more about the species. The de- scription is based on one male, pinned after original preservation in alcohol. Dactylolabis (Dactylolabis) sohiyi Byers and Rossman, new species (Figs. 1-3) Description.—Head: Dorsum gray, pru- inose, with short, pale setae; posterior ver- tex and occiput brown with slight pruinos- ity; longer, black setae above and behind eyes. Rostrum brown, palps darker brown. Antenna dark brown, with 14 flagellomeres. Thorax: Mesonotum dark reddish brown, with weakly defined, broad median stripe of slightly darker brown and diffuse, shorter stripe at each side: posterior scutum and scutellum darker, nearly dull black. Pleural areas and coxae light grayish tan. Halter light tan. Femora yellowish brown Tipulidae, Limoniinae, Dactylolabis, alar stigma near base but mostly brown; tibiae brown; tarsi dark blackish brown. Wings (Fig. 1): Pale yellowish brown; costal margin slightly undulate; stigma un- usually large, light brown, extending from C to R, and R; and from just beyond fork of Rs to apex of wing; veins C, Sc and R dark brown, membrane along Cu and 2A somewhat lighter brown. Anterior arculus absent but vestige of it present as basal spur off M at level of humeral crossvein. Abdomen: Most terga grayish brown with narrow, paler posterior borders; sterna light gray. Segments 8 and 9 light brown. Two bluntly rounded projections from pos- terior edge of tergum 9 (Fig. 2), creating broadly U-shaped median indentation. Out- er dististyles (Fig. 3) tapering from base to tip, with numerous setae; inner dististyles smooth, slightly twisted apically, without setae, directed mesad with apices crossing (in preserved specimen). Measurements: Body length (in fluid) 8.8 mm. Wing length 13.9 mm. Antenna about 2.9 mm. Femora (legs |—3): 7.6, 7.6 and 8.8 mm. Tibiae: 7.8, 6.9 and 7.3 mm. Tarsi: 6.2, 6.4 and 5.4 mm. Holotype.—Male collected from rock VOLUME 105, NUMBER 1 71 Figs. 1-3. abdominal tergum, dorsal aspect. 3, Apex of left basistyle and dististyles, dorsolateral aspect: bs inner dististyle, od—outer dististyle. Scales: a—Fig. 1, b—Figs. wall in Hickory Nut Gorge at Chimney Rock Park, elevation approximately 2,100 feet, Rutherford Co., North Carolina, on 19 April 1999, by D. Rossman. The specimen has had one wing and the terminal abdom- inal segments dissected off but glued (wa- ter-soluble glue) to the cardboard point bearing the rest of the fly. The fore and middle legs are intact; the hind legs are somewhat damaged but also glued to the point. Deposited in the Snow Entomologi- cal Division, Natural History Museum, University of Kansas, Lawrence. When captured, the fly was “‘scuttling up a sheer, dry rock face.” At the time D. sohiyi was collected, D. montana (Osten Sacken) was also present and fairly common, although in more shaded situations at a slightly lower elevation. Etymology.—The specific epithet, sohiyi, comes from the Cherokee and means “place of the hickory nut,” a reference to Hickory Nut Gorge, of which the Chimney Rock area forms the southern wall. The name was chosen to honor the indigenous people of the region, for whom the Gorge Dactylolabis sohiyi, male. 1, Wing, showing enlarged stigmal area (coarser shading). 2, Ninth basistyle, id— 23 was important in one of their myths (Ross- man 1988). Discussion.—Of the 19 Nearctic species of Dactylolabis named and described pre- viously, 13 are western, one is boreal-arctic, and five occur in eastern North America. In only one of these five, D. pemetica Alex- ander, does the male have the stigmal area of the wings greatly enlarged. The known range of D. pemetica extends from Maine southward to North Carolina, where it may overlap the range of D. sohiyi. Accordingly, males of these two species may be usefully compared. Most noticeable are differences in the wings: those of D. sohiyi are about 46% longer than the greatest recorded length for D. pemetica and about 4.8 mm. longer than the body; and the enlarged stig- ma in the former appears smooth (with very fine hairs barely visible at high magnifica- tion) and distinctly darker than the rest of the wing, as contrasted with “‘hairy” and “faintly darkened” in D. pemetica. Sexual dimorphism in the wings is tentatively pre- sumed to occur in D. sohiyi as in D. pe- metica (Alexander 1936: 288, fig. 1). The 72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mesonotum in D. pemetica is almost uni- formly dark brown, while in D. sohiyi it has dark brown stripes on a background of paler reddish brown. Published illustrations of the ninth ab- dominal tergum and dististyles (Osten Sacken 1869, Snodgrass 1904) of the male of North American Dactylolabis are not very useful for comparison with those of D. sohiyi. A species similar to D. sohiyi in these structures is the European D.(D.) transversa (Savchenko 1989: 70, fig. 1), which has more typical wings. All these il- lustrations do, however, indicate a setifer- ous outer dististyle and smooth inner distis- tyle; and Osten Sacken’s figure of D. cub- italis (O.S.) shows a ninth tergum quite dif- ferent from that of D. sohiyi. Alexander (1936) for some reason did not mention male genital structures in describing D. pe- metica. LITERATURE CITED Alexander, C. P. 1936. The crane-flies (Tipulidae) of New England. Fourth supplementary list. Occa- sional Papers of the Boston Society of Natural History 8: 273-292. Osten Sacken, C. R. 1869. Monographs of the Diptera of North America. Part TV. Smithsonian Miscel- laneous Collections 8: 1-345, 4 plates. Rossman, D. A. 1988. Where legends live: A pictorial guide to Cherokee mythic places. Cherokee Pub- lications, Cherokee, North Carolina. Savchenko, E. N. 1989. Limoniid crane flies of the fauna of the U.S.S.R. Academy of Science of the Ukranian S.S.R., Kiev, 380 pp. (In Russian.) Snodgrass, R. E. 1904. The hypopygium of the Tipu- lidae. Transactions of the American Entomologi- cal Society 30: 179-236. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 73-79 INVERTEBRATE BIODIVERSITY (ANTS, BEES, AND OTHERS) ASSOCIATED WITH STEM DOMATIA OF THE INDIAN MYRMECOPHYTE HUMBOLDTIA BRUNONIS WALLICH (MAGNOLIOPHYTA: FABACEAE) FRED R. RICKSON, MELINDA M. RICKSON, KUMAR GHORPADE, BETH B. NORDEN, AND KARL V. KROMBEIN (FRR, MMR) 247 Rainbow Drive, PM.B. 14735, Livingston, TX 77399-2047, U.S.A. (e-mail: ricksonm@hotmail.com); (KG) P.O. Box 8439, 1861 Bethel Street, St. Thomas Town, Bangalore 560 043, India; (BBN, KVK) Department of Systematic Biology—En- tomology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0165, U.S.A. Abstract.—The legume tree genus Humboldtia Vahl contains three mutualistic, myr- mecophytic species. These trees develop inflated, hollow internodes which differentiate a single, self-opening entrance hole for each domicile unit. At leaf flush, hundreds of new, accessible, empty domatia are available to any organism capable of entering and defending the hollow space. In terms of abundance, ants are the primary occupant of the hollow internodes of H. brunonis Wallich of India, followed by a bee, Braunsapis sp., and several minor inhabitants including an arboreal annelid. Humboldtia laurifolia Vahl in Sri Lanka is the closest relative of H. brunonis, and we compare and discuss the internode inhabitants of these two geographically separated, but morphologically very similar tree species. Key Words: India, Sri Lanka Numerous plants have entered into mu- tualistic relationships with ants. The plant evolves either, or both, a solid or liquid food supply, and a domicile in the form of leaf pouches, inflated leaf petioles, or a hol- low stem, while the ants provide a modi- cum of anti-herbivore defense. The simplest arrangement is where a plant possesses ex- trafloral nectaries which are visited by an assortment of ants, wasps, and other inver- tebrates to obtain nectar from these secre- tory tissues (Kopter 1991, Rickson and Rickson 1998). In these “open” systems there is usually no dominant ant species and the visiting species assemblage can change over time. Those plants possessing a pre- dictable domatia, or a food source requiring some innate ability to access, often have a Humboldtia brunonis, Humboldtia laurifolia, ants, Braunsapis, mutualism, particular ant species dominating the plant, although this too can change through time, perhaps through simple competition (Mas- chwitz et al. 1991), or seemingly at a par- ticular age stature of the plant (McKey 1991). Often those ants which are restricted to a particular plant species tend to be very aggressive toward intruders, and the plants, conversely, can have lower innate chemical defenses (Rehr et al. 1973). The actual suc- cess of the protection seems to vary widely from system to system, but where mea- sured, a positive benefit to the plant usually arises from the relationship. Little information is available on the total complement of organisms living within a plant’s domatia, across a series of popula- tions, because the ants dominating the bio- 74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mass of associated organisms are usually collected and reported as the animal part- ners involved (Huxley 1978). In a number of relationships, the ant, of- ten a founding queen, chews a hole into a preformed, hollow domicile. In these situ- ations, there is always an ant present within the chamber, the queen and/or her brood, should another organism seek to occupy the same space. Another group of plants also develops preformed domatia, but the plant provides a self-opening entrance hole to each of the chambers. If there is already an ant colony present on the plant, workers can move eggs or brood into newly available chambers as they become available. Ants are always dominant in these situations. However, in the case of a new leaf flush Where each leaf is associated with an ad- jacent stem domatium having an open en- trance hole, there may be a superabundance of open potential nest sites available to any organism capable of accessing the hole first, without any predisposed ability to find a specific location or method of chewing into the plant tissue. In such cases, other organ- isms may also evolve a predisposition to regularly take up residency in the hollow domatia and either compete with ants for dominance, or coexist. Humboldtia, with four species in south- ern India and one species in Sri Lanka, has three species which develop extrafloral nec- taries on the leaf blades, leaf stipules, and flower buds, and domatia in the form of in- flated, hollow stem internodes (Krombein et al. 1999). Each internodal domatium has a single, consistently placed, self-opening en- trance hole. Potential occupants of the hol- low internodes do not need any special lo- cation or chewing instincts, or special mouth parts, to gain access to the open cav- ity. The node between each hollow internode along a branch is solid. This creates a linear series of independent units, each with a pre- formed access opening. A shoot, which has an average of eight to ten new leaf/inter- nodes associated with each stem flush, pro- duces a similar number of individual, po- tential nest sites with each leaf flush. An annual production of 20—30 new shoots produces a corresponding 200—300 new po- tential nest sites, each year, available to any organism capable of accessing the entrance hole and defending the hollow internode space. The ant-associated Humboldtia spe- cies are gregarious with as many as 100+ individuals growing in close proximity, and so the yearly leaf/domatia production pro- duces a very large number of potential nest- ing sites within a small area. Humboldtia laurifolia Vahl, endemic to the southwestern lowland wet zone of Sri Lanka, has been thoroughly sampled and its internode-inhabiting associates enumerated (Krombein et al. 1999). The inhabitants in- clude: 14 species of ants, four wasps, one bee, two dipterans, three coleopterans, a pseudoscorpion, and an annelid. Included in this list are seven undescribed species and a number of first and second order predators and parasites on the various inhabitants. Fi- nally, the cryptic nature of the Humboldtia associates can be appreciated by noting that the Smithsonian Institution carried out a Fauna of Ceylon project over 12 years and yet the inhabitants of this common tree largely escaped detection and collection. Humboldtia brunonis Wallich, is a small, gregarious, understory tree found along the eastern edge of the Western Ghats in south- ern India from about 13°70'N, 75°00’E to 11°30’N, 76°30’E (Ramesh et al. 1997). This species, along with H. decurrens Bed- dome ex Oliver found in the southwestern part of the Western Ghats, possesses the same extrafloral nectary placement and do- micile traits as described for H. laurifolia (Krombein et al. 1999). MATERIALS AND METHODS Several hundred hollow internodes from random trees of H. brunonis growing in the Makut Reserve Forest, Coorg District, Kar- nataka State, India (12°10'N, 75°50’E) were first collected by KG in 1994, with author KVK making a second collection effort VOLUME 105, NUMBER | within this population of trees in 1997. The Makut population is a little south of the center of the North-South distribution of the species (Ramesh et al. 1997). The inter- nodes were split open, distribution of the included organisms noted, and samples pre- served in 70% alcohol. Only those organ- isms actually collected from within the hol- low internodes, as opposed to specimens found foraging on the tree, were included in this summary. RESULTS We present below, in general order of abundance, the invertebrates collected from Humboldtia brunonis hollow internodes. HYMENOPTERA Formicidae Krombein collected 100 hollow inter- nodes of which 27 contained ants, while an- other 38 were empty, 13 contained a com- mon bee Braunsapis sp. (see below), and the remaining internodes possessed an as- sortment of organisms. Eleven ant species were found nesting within the hollow stem internodes. There was no single dominant ant taxon although Crematogaster spp. were present in approximately 50% of the ant-occupied internodes, and the three spe- cies within that genus contributed approxi- mately equally to that 50%. A surprising finding was the lack of weaver ants, Oe- cophylla smaragdina (F) associating with H. brunonis at Makut. This species is a common visitor to many plant species with extrafloral nectaries, and is found nesting on both AH. laurifolia in Sri Lanka and H. vahliana Wight in south India. Two unde- scribed species of ants were collected. The following species were nesting with- in hollow internodes of Humboldtia bru- nonis. The species are arranged according to Bolton (1995). Dolichoderinae Tapinoma indicum Forel Technomyrmex albipes (FE Smith) Technomyrmex, prob. n. sp. 15 Formicinae Camponotus compressus F. Polyrhachis illaudata Walker Polyrhachis, n. sp. Myrmicinae Crematogaster nilgirica Emery Crematogaster prob. travancorensis Forel Crematogaster sp. | Monomorium sp. Tetramorium pacificum Mayr Anthophoridae Braunsapis sp., an undescribed bee, was found in 13 internodes out of 100 collected by KVK, and was also common in the KG collections. Both sexes and all stages of de- velopment were present. Given that the spe- cies has not been collected before, despite being a common associate of the tree, we assume that it does not venture far from the internodes. There was no evidence of the food used by this bee. This species is the same associated with H. laurifolia in Sri Lanka, however in Sri Lanka, Braunsapis sp. was only found in four internodes out of over 1300 sampled, while two wasps, Krombeinictus nordenae Leclercq, and Crossocerus mukalanae Leclercq were common. Neither of the wasps however, was collected from H. brunonis in India. Bethylidae A solitary, small, pale, bethylid pupa was removed from a single internode. These small wasps are parasitoids of Coleoptera and Lepidoptera, but no other organisms were noted in the internode. COLEOPTERA Cleridae Numerous clerid larvae were collected from three internodes. In one, a single late stage predaceous larva was in an internode containing Braunsapis eggs. Another larva of the same species was found in an inter- node with a queen and worker Cremato- gaster, and approximately 30, 2mm long 76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON clerid larvae were taken from another in- ternode which lacked other organisms. Clerid larvae were also found associated with Braunsapis brood internodes in H. laurifolia from Sri Lanka. Staphylinidae A staphylinid larva was found in an in- ternode without any evidence of prey, while a second larva was present in an internode which had a silken cap across the cavity above the larva. Diptera A dipterous pupa, with black legs, was attached to, and below, a flat closure disk. The cavity had mucus coating the walls. Three puparia of another diptera were re- moved from internodes, and 20 immature diptera were collected from a KG bulk sam- ple. Collembola Collembola were found just within the entrance hole of three unoccupied inter- nodes. Orthoptera Two Blattodae oothecae were found within an internode, and several adult cock- roaches were found on trees. OLIGOCHAETA Megascolecidae Pale, approximately 2.5 cm long annelids were found in a number of internodes. These appeared similar to the undescribed, immature Perionyx sp. collected from H. laurifolia, but were adults in H. brunonis. This arboreal annelid is interesting in that the posterior one-third is curved and acts as a suction devise while the organism moves leech-like over a surface (Krombein et al. 1999), ARANEIDA Salticidae A small salticid, with some webbing, was removed from one internode. Chilopoda A single, small centipede was collected within an otherwise empty internode. DISCUSSION The geologic history of India and Sn Lanka is one of intermittent connection by a land bridge, corresponding with various ice age fluctuations in ocean level. Fossil evidence indicates that numerous plant and animal species traveled this bridge, while others did not seem to make the transition, leading to a mixture of species common to both countries and also numerous endemics on the two land masses. The center of ori- gin for Humboldtia, based on current spe- cies distribution, is within the Indian West- ern Ghats, but there is no fossil record in- dicating how far east, towards Sri Lanka, the genus might have once extended. Based on the fossil record, the southern portion of India was more highly forested in the past, and either H. brunonis, or some extinct an- cestor, could have existed in close proxim- ity to the Indian-Sri Lankan land bridge, making seed dispersal between the two land masses more probable than today. We do not envision a previous presence of other Humboldtia spp. on Sri Lanka with an ex- tinction of all taxa except H. laurifolia. Likewise, given the wide range of environ- ments presently occupied by H. /aurifolia, and its current success within those envi- ronments, we do not think that this species once existed in India and then become ex- tinct. Morphologically, and within a taxonomic framework, H. brunonis and H. laurifolia are the most closely related species within the genus. Additionally, H. brunonis is the sole Indian Humboldtia species existing on the eastern side of the Western Ghats. Giv- en the previous forestation of southern In- dia, we hypothesize a seed dispersal event from the Indian subcontinent giving rise to what today is H. laurifolia. If H. brunonis and H. laurifolia originat- ed by speciation after a seed dispersal VOLUME 105, NUMBER | event, rather than a vicariant separation of existing species, then the present associated invertebrates evolved their association with these Humboldtia spp. in situ, and the oc- cupants found in the two tree species rep- resent independent adaptations to a very similar domatium morphology and nectary location, on two separate land masses. To- day, the two species exist in similar envi- ronments of wet evergreen to wet decidu- ous forests. These similar habitats probably forecast a somewhat invertebrate fauna from which the currently associated organisms evolved their relationship. Of eleven inhabiting ant taxa, there is no dominant species on H. brunonis. Techno- myrmex albipes is strongly dominant on H. laurifolia across habitats, but this ant is similar only an occasional occupant of H. brunonis. Conversely, Crematogaster was found nest- ing on almost every H. brunonis tree, com- prising approximately 50% of the domatia inhabitants, but this genus was collected from only 13 internodes out of over 1,300 domatia sampled from H. laurifolia in Sri Lanka. Tapinoma indicum 1s a minute ant that was found nesting in two internodes on H. brunonis, and four domatia of H. lauri- folia. All colonies were very strong with hundreds of workers and all stages of larvae and pupa in each internode. Tapinoma in- dicum occurs as a minor occupant in several ant plants in the old world and seems to be able to co-exist with a number of ant spe- cies, even in an adjacent domatium on Humboldtia, but all the domatia on even a small branch. The two species of Polyrhachis which nest within H. brunonis internodes contrast with P. bug- niont, associated with H. laurifolia in Sri Lanka, which binds leaves together to build its nests and so nests externally to the hol- low internodes. Polyrhachis gracilior Forel binds leaves of Humboldtia decurrens in southern India, just as P. bugnioni does in Sri Lanka. Thus, H. brunonis is distinct from the other two myrmecophytic Hum- boldtia species in having its associated Po- never dominates Ci lyrhachis species nesting within the hollow internodes rather than in leaf nests. The other four ant species collected from H. brunonis domatia, Camponotus com- pressus FE, Monomorium sp., Tetramorium pacificum Mayr, and Technomyrmex prob. n. sp., were collected from only one to sev- eral internodes each. Of these ant species, T. pacificum and the Monomorium species are identical to those from H. laurifolia, whereas C. compressus and Technomyrmex prob. n. sp. are unique to H. brunonis in our collections. A major difference between H. brunonis and H. laurifolia is the lack of a wasp spe- cies being associated with H. brunonis, and a bee, Braunsapis sp. taking that place. A survey of the area to determine the avail- ability of small wasps capable of using the internodes for nest sites would be very in- teresting, and surveys of other populations of H. brunonis in different environments may yet find wasps using the internodes. The bee Braunsapis sp. which is a minor inhabitant of H. laurifolia becomes a major inhabitant of H. brunonis possibly because of the absence of a competing wasp species. It is interesting that a bee or wasp 1s asso- ciated in a major way with both myrme- cophytic Humboldtia species; this is not just an ant-associated tree. Representatives of the beetle family Cler- idae are associated with Braunsapis, prob- ably as a brood predator, on both H. bru- nonis and H. laurifolia. However, the pseu- doscorpion Haplochernes warburgi, a com- mon ant predator on H. laurifolia, was absent from H. brunonis. Another common aggressive predator associated with numer- ous H. laurifolia, internodes, a new Diptera (Keroplatidae) species, Platyceridion edax Chandler and Matile, is absent from our H. brunonis population. In fact, there seems to be no strong predator present in the Makut population despite a diverse ant fauna as potential prey. Instead, we find the few Cleridae, Staphylinidae, and diptera larvae to be the only possible predators. Annelids, perhaps the same undescribed, 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON arboreal Perionyx species, are found in both Humboldtia species. In both cases the worms are nocturnal or exit the internodes during light daytime rains. We never found the worms associated with another inverte- brate within an internode, but see no obvi- ous method used to exclude competitors from the internodal space. The population of H. brunonis sampled for this study did not possess either the spe- cies diversity or density of domatia occu- pancy found in H. /aurifolia. Of the 100 H. brunonis internodes opened by KVK, 62% were occupied. Domatia occupancy on H. laurifolia was always above 90% for a giv- en tree, and reached 100% on several bulk samples collected from single trees. We have reported on a single branch diversity of H. laurifolia, with 10 adjacent domatia possessing two ant species, Technomyrmex albipes and Cataulacus taprobanae F. Smith, the dipteran Platyceridion edax, and the wasp Krombeinictus nordenae Leclercq (Krombein et al. 1999). We did not find such single-branch diversity on H. brunon- is, rather Crematogaster would usually dominate domatia along a branch, with one of the other invertebrates in single, random internodes. A very preliminary examination of two small populations of H. decurrens, the third myrmecophytic species, in the southern Western Ghats of India, revealed three spe- cles of ants, Polyrhachis gracilior Forel, Technomyrmex prob. brunneus Forel, and an undescribed species of Cataulacus. This very southern species of Humboldtia, which prefers wetter habitats than H. bru- nonis, deserves a complete survey of its as- sociated organisms. It must be pointed out that of the eleven ant taxa collected from H. brunonis, a very common and easily accessible plant, two are new species. These discoveries, along with another undescribed ant collected from a small sample of hollow internodes of H. decurrens from southern India, and up to 11 new insect species across three Orders from H. laurifolia in Sri Lanka, point out how these cryptic plant cavities have never been collected, and how they harbor a unique fauna along with such common spe- cies as the ant Technomyrmex albipes. Finally, it must be mentioned how im- portant the genus Humboldtia is in harbor- ing a unique series of organisms relating to local biodiversity. Humboldtia laurifolia from evergreen wet locations harbors at least 11 undescribed organisms, while a sin- gle population of H. brunonis, in a dryer environment, produced four undescribed species, and H. decurrens, one new species after vary limited collection. The role of Humboldtia and its hollow, self-opening in- ternodes in harboring cryptic, rare species may be unparalleled among invertebrate-as- sociated plants in south Asia. ACKNOWLEDGMENTS We thank Raghavendra Gadagkar, Centre for Ecological Sciences, Indian Institute of Science, Bangalore, India for facilitating the field collecting by KVK. Collectively we thank the following specialists for iden- tification of various Formicidae: Barry Bol- ton, The Natural History Museum, London; Rudolf J. Kohout, Queensland Museum, Australia (Polyrhachis); T. M. Musthak-Ali, Department of Entomology, University of Agricultural Sciences GKVK Campus, Bangalore, India; and Stephen O. Shattuck, Australian National Insect Collection, Can- berra. LITERATURE CITED Bolton, B. 1995. A new General Catalogue of the Ants of the World. Harvard University Press, Cam- bridge, Massachusetts, 504 p. Huxley, C. R. 1978. The ant-plants Myrmecodia and Hydnophytum (Rubiaceae), and the relationships between their morphology, ant occupants, physi- ology and ecology. New Phytologist 80: 231—268. Koptur, S. 1991. Extrafloral nectaries of herbs and trees: modelling the interaction with ants and par- asitoids, pp 213-230. Jn Huxley, C. R. and D. FE Cutler, eds. Ant—Plant Interactions. Oxford Uni- versity Press, Oxford. Krombein, K. V., B. B. Norden, M. M. Rickson and F R. Rickson. 1999 Biodiversity of the domatia oc- cupants (ants, wasps, bees, and others) of the Sri VOLUME 105, NUMBER | Lankan myrmecophyte Humboldtia laurifolia Vahl (Fabaceae). Smithsonian Contributions to Zoology 603: 1—34. Maschwitz, U., B. Fiala, J. Moog and L. G. Saw. 1991. Two new myrmecophytic associations from the Malay Peninsula: ants of the genus Cladomyrma (Formicidae, Camponotinae) as partners of Saraca thaipingensis (Caesalpiniaceae) and Crypteronia griffithii (Crypteroniaceae). Insectes Sociaux 38: 27-35. McKey, D. 1991. Phylogenetic analysis of the evolu- tion of a mutualism: Leonardoxa (Caesalpini- aceae) and its associated ants, pp. 310—334. In veh Huxley, C. R. and D. EF Cutler, eds. Ant—Plant Interactions. Oxford University Press, Oxford. Ramesh, B. R., J-P. Pascal and C. Nouguier. 1997. At- las of Endemics of the Western Ghats (India). In- stitat Francais de Pondichery, Pondicherry, 403 Pp: Rehr, S. S., P. P. Feeny and D. H. Janzen. 1973. Chem- ical defense in Central American non-ant-acacias. Journal of Animal Ecology. 42: 405-416. Rickson, F R. and M. M. Rickson. 1998. The cashew nut, Anacardium occidentale (Anacardiaceae), and its perennial association with ants: Extrafloral nectary location and the potential for ant defense. American Journal of Botany 85: 835-849. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 80-99 A CHECKLIST OF COMMONLY INTERCEPTED THRIPS (THYSANOPTERA) FROM EUROPE, THE MEDITERRANEAN, AND AFRICA AT U.S. PORTS-OF-ENTRY (1983-1999). PART 1. KEY TO GENERA Davip A. NICKLE Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, 10300 Baltimore Avenue, Building 005, BARC-West, Beltsville, MD 20705-2350 Abstract.—Although there are more than 1,000 described species of thrips from Europe, the Mediterranean region, and Africa, since 1983 only 130 species have been intercepted in cargo and shipments of plants coming into the various ports-of-entry in the United States. Of these, only 23 species consistently made up ca. 85% of the identifiable thrips. This paper is a checklist of thrips commonly intercepted on plants coming into the U.S. from Europe, the Mediterranean region, and Africa; keys with figures are included for the 57 represented genera. It is the first of a five-part series aimed to facilitate identifications by port identifiers at U.S. Department of Agriculture, Animal and Plant Health Inspection Service (USDA, APHIS), ports-of-entry. Key Words: | thrips, pests of flowers, Europe, Africa Thrips (Thysanoptera) are a group of mi- nute insects usually less than 1.0 mm in length. adults, with 2 pairs of narrow membranous Most species are fully alate as wings having few or no veins but with well developed fringe-like cilia around the mar- gins. The sexes of thrips are similar in most anatomical structures, but parthenogenesis is Common among some species, and males are often rare in other species (Stannard 1968). Metamorphosis is complex, usually with active larval stages (I and II), a propupa and pupa in Terebrantia and propupa, pupa I and pupa II in Tubulifera, followed by the adult stage (upon which this paper is based). Many species feed on plant tissue of many host plants, including agricultur- ally important fruits and vegetables, and cutflowers imported into this country from abroad. Some species are host specific or feed on a limited variety of closely related plant species; others are polyphagous; some species are predaceous, feeding on mites Figs. 1-14. => Morphological features of Thysanoptera. 1, Thripidae (Thrips sp.), dorsal aspect, showing left half with selected structures. 2, Aeolothripidae (Stomatothrips sp.), dorsal aspect, right half. 3, Phlaeothripidae (Haplothrips sp.), dorsal aspect, showing right wings only. 4, Head and pronotum of a thrips (Frankliniella species), dorsal aspect, showing ocellar setae I, I], and HI. 5, Aeolothrips pronotum, dorsal aspect. 6, Melanthrips pronotum, dorsal aspect. 7, Retithrips syriacus, head and pronotum, dorsal aspect. 8, Aeolothrips sp., antennal segment III. 9, Melanthrips sp., antennal segment III. 10, Retithrips syriacus, terminal antennal segments. 11, Dendrothrips ornatus, terminal antennal segments. 12, Heliothrips haemorrhoidalis, pronotum showing regular sculpturation, dorsal aspect. 13, Rhipiphorothrips miemsae, pronotum showing irregular sculpturation, dorsal aspect. 14, Dendrothrips ornatus, head and pronotum, dorsal aspect. I Pp VOLUME 105, NUMBER 1 81 mesonotum metanotum forewing scale fringe cilia setae of first vein » ES . HA H . ocellar seta I Wy ellar seta I compound eye ocellar seta IT ocellus setae of second vein tergite VILL « NIN? Gl P E . a ? : SOE Pp ? \\ } posteroangular seta anteromarginal seta del Nee ah ea od Coie Pe slice ‘ aby eo oO 0 OY ‘e) taw)) 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and small insects or on other thrips (Mound et al. 1976). Although literature is extensive regarding the biology, systematics, behavior, and pest potential of thrips [e.g., see Kirk (1996), Lewis (1997), and Parker et al. (1991) for reviews with presentations of the literature], there is a need for regional guides to iden- tify species inadvertently transported in in- ternational commerce of agricultural and horticultural shipments. This paper is the first of a series of identification guides for the more commonly intercepted thrips from a particular region arriving in ports of the United States and is meant to facilitate identification of those species most fre- quently found in flowers, fruits, and leaves of a wide range of commodities. It 1s mod- eled in part after a similar recently pub- lished paper on the identification of larval Pyraloidea (Lepidoptera) intercepted in U.S. ports-of-entry (Solis 1999). It con- denses identification keys already in exis- tence on either a worldwide scope (Mound and Kibby 1998) or a more limited regional scope (e.g., parts of Europe only or Africa only) (Dyadechko 1977, Mound et al. 1976, Priesner 1964) to include only the thrips most likely to be encountered in commerce from Europe/Mediterranean Region/Africa. It also makes use of information derived from generic revisions—e.g., Odontothrips (Pitkin 1972), Thrips (Nakahara 1994), Frankliniella (Moulton 1948), Anaphoth- rips, Ceratothrips, and Tenothrips (Bhatti 1967, 1973, 1978, 1990)—or of informa- tion derived from species lists (e.g., Naka- hara 1997, 1999) and catalogs (Jacot-Guil- larmod 1970-1977). Obviously, many gen- era and species have been excluded from this treatment, but conversely, it is unlikely that those species would be received in pro- duce and other commerce that routinely en- ters the United States. This paper is based on the adult female sex only, partly because males are seldom encountered in the small sampling procedures used by port inspec- tors and partly because for many species— even some economically important ones— males have not been described or adequate- ly treated. Because life stages of many spe- cies have not been fully studied, immature stages also are excluded from this paper, al- though keys to immature stages are avail- able for selected species, some of which are included herein (Nakahara and Vierbergen 1998). This paper (Part 1) is the first of a 5-part series directed at identifying thrips from Europe/Mediterranean region/Africa com- monly intercepted by USDA/APHIS port identifiers. It is intended to provide an over- view of the diversity of thrips entering this country and to indicate the frequency that each species was encountered over a 17- year period from 1983—2000. Clearly, some species commonly are encountered, while others are rare. In order to sort out this di- versity, a key to 57 represented genera is included, with figures (both line drawings and scanning electron micro- graphs) to facilitate identifications to genus level. Parts 2 and 3 will treat those species comprising a significant proportion of the intercepted fauna, 1.e., thripine species shar- ing the character of the presence of ctenidia on tergite VIII: part 2 will cover Frankli- niella and related genera (16 species) and part 3, the genus Thrips (23 spp.). Part 4 will treat the 52 remaining species of Thri- pidae (6 spp. in 5 genera of Panchaetoth- ripinae and 46 spp. in 33 genera of Thri- pinae). Part 5 will deal with Aeolothripidae (15 spp. in 3 genera) and Phlaeothripidae (21 spp. in 5 genera). Sources of quarantine interceptions 1n- clude commercial shipments, inspections of passenger baggage, aircraft or ship quarters, stores, galleys, and mail. In fact, most in- terceptions are from aircraft quarters. Not- withstanding, nearly 1,000 samples are an- nually sent the Systematic Entomology Laboratory, USDA, for urgent identification of unknown thrips samples representing species that cannot be reliably identified at ports-of-entry and which require immediate identification for action at the ports. along VOLUME 105, NUMBER | 83 Table 1. Species of thrips intercepted at ports of entry (Numbers represent accumulated interceptions over the period 1983-1999). Ranges of species in Europe, the Mediterranean, and Africa are indicated with an **x.”* Establishment or occurrence of any of these species in the United States also is indicated by an **x’* under U.S. Eur Med Afr US AEOLOTHRIPIDAE Aeolothrips Haliday 1836 brevicornis Bagnall 1915 2 x bucheti Bagnall 1934 4 X collaris Priesner 1919 5 Xx x x x deserticola Priesner 1929 4 x x ericae Bagnall 1920 3 x x fasciatus (Linnaeus 1758) 1 x x x m8 intermedius Bagnall 1934 13 Xx linarius Priesner 1948 l x meridionalis Priesner 1948 1 Xx x x scabiosatibia Moulton 1930 7 Xx tenuicornis Bagnall 1926 1 X undetermined species 31 Franklinothrips Back 1912 vespiformis (Crawtord 1909) i [x] Xx Melanthrips Haliday 1836 fuscus (Sutzer 1776) 11 Xx Xx x gracilicornis Maltbaek 1931 4 x 8 x pallidior Priesner 1919 2 Xx x Xx undetermined species 6 THRIPIDAE Anaphothrips Uzel 1895 articulosus Priesner 1925 I av obscurus (Miiller) 1776 14 Xx x Xx sudanensis Trybom 1911 l x x Apterothrips Bagnall 1908 apteris Daniel 1904 1 Xx secticornis (Trybom 1896) 1 Xx x Aptinothrips Haliday 1836 rufus (Goeze 1776) 1 x Xx x x stylifer Trybom 1894 il Xx Xx Ceratothripoides Bagnall 1918 brunneus Bagnall 1918 7 x Ceratothrips Hood 1919 ericae (Haliday 1836) 7 x Chaetanaphothrips Priesner 1957 undetermined species 4 Chirothrips Haliday 1836 aculeatus Bagnall 1927 2 Xx X manicatus (Haliday 1836) 6 Xx ? X meridionalis Bagnall 1927 1 x x x mexicanus Crawford 1909 | x Xx Dendrothripoides Bagnall 1923 innoxius Karny 1914 1 x X venustus Faure 1941 ] x undetermined species 1 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Eur. Med. Afr. US Dendrothrips Uzel 1895 degeeri Uzel 1895 ornatus (Jablonowski 1894) saltator Uzel 1895 2) undetermined species 1 Mm Ww ~“~ Pal Dichromothrips Priesner 1932 corbetti (Priesner 1936) 1 [7] x undetermined species | Drepanothrips Uzel 1895 reuteri Uzel 1895 4 x x Echinothrips Moulton 1911 americanus Morgan 1913 ] x x Frankliniella Karny 1910 fusca (Hinds 1902) Th x x intonsa (Trybom 1895) 94 x occidentalis (Pergande 1895) 448 Xx Xx x pallida (Uzel 1895) I x schultzei (Trybom 1910) 55 x x tenuicornis (Uzel 1895) 136 x tritici (Fitch 1855) 3 undetermined species M2. Glaucothrips Karny 1921 glaucus (Bagnall 1914) 5 xe Heliothrips Haliday 1836 haemorrhoidalis (Bouché 1838) 5 X x x Xx Hercinothrips Bagnall 1932 bicinctus (Bagnall 1919) 1 dimidiatus Hood 1937 1 Tridothrips Priesner 1940 iridis (Watson 1924) 2 x Xx Leucothrips Reuter 1904 undetermined species 1 Limothrips Haliday 1836 cerealium (Haliday 1836) 18 Xx x x % denticornis (Haliday 1836) 9 x undetermined species 1 Megalurothrips Bagnall 1915 sjOstedti (Trybom 1910) BS) x undetermined species 1 Microcephalothrips Bagnall 1926 abdominalis (Crawtord 1910) 1s, ~ Pv Mycterothrips Trybom 1910 consociatus (Targioni-Tozzetti 1887) latus (Bagnall 1912) undetermined species [— EL) Neohydatothrips John 1929 samayunkur (Kudo 1995) 13 Xx x undetermined species 2 VOLUME 105, NUMBER 1 85 Table 1. Continued. Eur. Med Afr U.S Odontothrips Amyot & Serville 1843 karnyi Priesner 1924 132 x X x undetermined species 2 Oxythrips Uzel 1895 nobilis Bagnall 1927 1 x undetermined species l Palmiothrips zur Strassen 1965 annulicornis zur Strassen 1965 1 x Parthenothrips Uzel 1895 dracaenae (Heeger 1854) 3 X Xx x Prosopothrips Uzel 1895 nigriceps Bagnall 1927 1 x % Retithrips Marchal 1910 syriacus (Mayet 1890) 2 x x x Rhipiphorothrips Morgan 1913 miemsae Jacot-Guillarmod 1937 2 Xx Scirtothrips Shull 1909 aurantii Faure 1929 5 x dorsalis Hood 1919 3 Xx inermis Priesner 1933 1 Xx x undetermined species 3 Scolothrips Hinds 1902 longicornis Priesner 1926 1 x Selenothrips Karny 1911 rubrocinctus (Giard 1901) I X X Synaptothrips Trybom 1910 africanus (Moulton 1936) | X distinctus (Bagnall 1915) 9 x gezinae (Faure 1938) 6 x undetermined species 7 Taeniothrips Amyot & Serville 1845 inconsequens (Uzel 1895) 3 x X Tameothrips Bhati 1978 tamicola (Bagnall 1914) | me Tenothrips Bhatti 1967 discolor (Karny 1907) 9 ? Xx fricit (Uzel 1895) 4 x undetermined species 5 x Thrips Linnaeus 1758 acaciae Trybom 1910 2 x angusticeps Uzel 1895 24 x x x italicus (Karny 1907) 24 [x] atratus (Haliday 1836) 52 Xx Xx x australis (Bagnall 1915) 10 x x Xx x brevicornis Priesner 1920 3 x flavus Shrank 1776 28 Xx fulvipes Bagnall 1923 6 X fuscipennis Haliday 1836 200 Xx x 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table |. Continued. ———— eee trehernei Priesner 1937 1 ? italicus (Bagnall 1926) 30 x major Uzel 1895 178 x Xx x mareoticus (Priesner 1932) l x meridionalis (Priesner 1926) 23 x Xx microchaetus Karny 1920 | minutissimus Linnaeus 1758 7 x Xx nigropilosus Uzel 1895 9 x ? x palmi Karny 1925 11 Xx physapus Linnaeus 1758 | pillichi Priesner 1924 | x simplex (Morison 1930) 26 x Xx Xx x spadix Hood 1932 | x tabaci Lindeman 1889 474 x x x Xx tenellus Trybom 1912 l Xx urticae Fabricius 1781 3 , vulgatissimus (Haliday 1836) 98 x x undetermined species 37 PHLAEOTHRIPIDAE Aleurodothrips Franklin 1909 fasciapennis (Franklin 1908) 1 x x Bolothrips Priesner 1926 cingulatus (Karny 1916) | X Elaphrothrips Buffa 1909 undetermined species p2 Eparsothrips zur Strassen 1968 varicornis (Bagnall 1919) 4 x Gynaikothrips Zimmermann 1900 ficorum (Marchal 1908) l x x Haplothrips Amyot & Serville 1843 aculeatus Fabricius 1803 4 Xx articulosus Bagnall 1926 3 cerealis Priesner 1939 l X clarisetis Priesner 1930 2 Xx distinguendus Uzel 1895 2 x x fulginosus Schille 1912 l x gowdeyi (Franklin 1908) 65 x x nigricornis Bagnall 1910 12 reutert (Karny 1907) 4 setiger Priesner 1921 l subtillissimus (Haliday 1836) 2 undetermined species 23 Hoplandrothrips Hood 1912 ellisi Bagnall 1914 1 flavipes Bagnall 1923 2 Xx x undetermined species l Karnyothrips Watson 1924 flavipes (Jones 1912) 2 melaleucus (Bagnall 1911) 1 a x } undetermined species ee SSSSSSSSSSSSSMMsseF VOLUME 105, NUMBER 1 Table 2. Complete list of species of thrips inter- cepted from commerce from Europe and Africa at U.S. ports of entry, 1994-1999 (species in bold font rep- resent most frequently intercepted species, tabulated in Table 3). Phlaeothripidae Haplothrips articulosus Bagnall 1895? Haplothrips fuliginosus Schille 1912 Haplothrips gowdeyi (Franklin 1908) Haplothrips nigricornis Priesner 1910 Hoplandothrips sp. Aeolothripidae Aeolothrips brevicornis Bagnall 1915 Aeolothrips collaris Priesner 1919 Aeolothrips deserticola Priesner 1929 Aeolothrips intermedius Bagnall 1934 Aeolothrips tenuicornis Bagnall 1926 Franklinothrips sp. Melanthrips fuscus (Sulze 1776) Thripidae Anaphothrips obscurus Miller 1776 Anaphothrips sudanensis Trybom 1911 Chirothrips manicatus (Haliday 1836) Chirothrips meridionalis Bagnall 1927 Dendrothrips ornatus (Jablonowski 1894) Frankliniella intonsa (Trybom 1895) Frankliniella occidentalis (Pergande 1895) Frankliniella schultzei (Yrybom 1910) Frankliniella tenuicornis (Uzel 1895) Limothrips ceralium (Haliday 1836) Limothrips denticornis (Haliday 1836) Neohydatothrips samayunkur (Kudo 1995) Odontothrips karnyi Priesner 1924 Oxythrips sp. Parthenothrips dracaenae (Heeger 1854) Taeniothrips inconsequens (Uzel 1895) Tenothrips discolor (Karny 1907) Thrips atratus (Haliday 1836) Thrips australis (Bagnall 1915) Thrips brevicornis Priesner 1920 Thrips flavus (Schrank 1776) Thrips fuscipennis Haliday 1836 Thrips italicus (Bagnall 1926) Thrips major Uzel 1895 Thrips meridionalis (Priesner 1926) Thrips nigropilosus Uzel 1895 Thrips physapus Linnaeus 1758 Thrips simplex (Morison 1930) Thrips tabaci Lindeman 1889 Thrips trehernei Priesner 1927 Thrips vulgatissimus (Haliday 1836) 87 It is an economically expensive propo- sition to protect U.S. agriculture and horti- culture. This protection requires time in- vested in identifying specimens at each port, sending suspect specimens to taxo- nomic specialists connected with the Sys- tematic Entomology Laboratory for final identifications, and actions taken once iden- tifications are made to resolve issues in- volving the entry status of infested ship- ments. When completed, these papers should provide identifiers and others with a powerful resource for identifying potential European, Mediterranean, and African thrips pests regularly threatening U.S. ag- riculture and horticulture. MATERIALS AND METHODS Two databases of thrips identifications, available to me at the Systematic Entomol- ogy Laboratory, were used to determine the species that were intercepted in shipments from European, Near East, and African countries. The first included species inter- cepted between 1983 and 1993 (Table 1), and the second for species intercepted from 1994 to 1999 (Table 2). Both databases in- cluded information concerning the country of origin, the plant host on which the spec- imen was found, and the identification of the thrips taxon. Records of interceptions originating from The Netherlands are often deceiving, since The Netherlands is host to the international market wherein cut flowers from all parts of the world are auctioned. The Netherlands Plant Protection Service has a staff at the flower market and auction in Alrsmeer. Although cut flowers are in- spected entering The Netherlands and other EU countries, consignments transshipped to the U.S. and other countries are usually not inspected, and many thrips are transported with their hosts in cargos. The first database (containing a total of 2,437 interceptions) (Table 1) and the second database (of 497 specimens) (Table 2) provided the basis for this study. These databases do not include thrips that were intercepted and identified by 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON = Table 3. Most frequently intercepted thrips from Europe and Africa at U.S. ports of entry, 1994-1999, based on a database of 497 identified specimens. Species were ranked from most frequently intercepted to species represented by more than unique specimens. Number of % of Total Cumulative Species Rank Interceptions (n 497) Percent Thrips tabaci | $1 20.0 20.0 Frankliniella occidentalis 2 59 14.6 34.6 Thrips fuscipennis 3 41 10.1 44.7 Thrips major 4 32 7.9. 52.6 Thrips vulgatissimus 5 22 5.4 58.0 Odontothrips karnyi 6 18 4.5 62.2 Frankliniella intonsa 7 15 37 65.9 Haplothrips gowdeyi 8 11 2h 68.6 Frankliniella tenuicornis 9 10 2.4 71.0 Frankliniella schultzei 10 i) Nez WPT) Melanthrips fuscus 1] 6 1.4 74.1 Thrips meridionalis 12 5 1.2 75.3 Thrips flavus 13 5 1.2 76.5 Limothrips cerealium 14 5 1.2 TT: Thrips atratus 15 4 0.9 78.6 Haplothrips nigricornis 16 4 0.9 79.5 Aeolothrips collaris 17 3 0.7 19.3 Thrips simplex 18 3 0.7 80.0 Thrips nigropilosus 19 2 0.5 80.5 Neohydatothrips samayunkur 20 2 0.5 81.0 Limothrips denticornis 21 2 0.5 81.5 Aeolothrips deserticola 22 2 0.5 82.0 Thrips australis 23 2 0.5 82.5 Dendrothrips ornatus 24 2 0.5 83.0 APHIS port identifiers. Ports with air flights from Europe and Africa, such as JFKIA (John E Kennedy International Airport), O’Hare, Atlanta, Houston, Los Angeles, and Seattle, have identifiers with authority to make identifications of some commonly intercepted species without verification from the SEL specialist, and these are not included in this paper. Also, since 1996, routine identifications made by the APHIS/ PPQ thrips specialist (Susan Broda) are also excluded. The problem with inclusion of data from APHIS/PPQ is that these data only include information on species that are not already established in the U.S. Thus, the numerous identifications of Frankliniella occidentalis and Thrips tabaci are not in- cluded. Since port identifiers need to iden- tify all species taken at U.S. ports, I decided to use the more complete SEL data. If APHIS/PPQ identifications had been in- cluded, the numbers in Tables 2 and 3 would have been skewed toward species that do not occur in the U.S., but it would not have changed the composition of the species list. In a review of historical records kept by APHIS/PPQ from 1923 to 1984, I noted two trends. (1) Early records are scant. At this time very few records existed of com- mercial shipments of flowers. Most inter- ceptions were from passenger baggage. These records always were of species found in the top 10 in Table 2, with the exception of Frankliniella occidentalis Pergande. This is a U.S. species that has been transported in commerce to other parts of the world in the late 20th century and has become established in Europe and Africa (as well as other parts of the world). It is now one of the most commonly inter- cepted species (see Tables 2-3). (2) Be- cause of changes in commercial trade, some species that previously were not (or VOLUME 105, NUMBER | only rarely) intercepted are now more fre- quently encountered. For example, Thrips palmi Karny, a tropical Asian species, be- came established in Africa and the Western Hemisphere, and more recently has be- come established in the U.S. (in Florida and Hawaii). After evaluating species in- terceptions enumerated in the two databas- es, I determined that at least 25 species should be added to the list of most com- monly intercepted thrips, most of them in the genera Thrips and Haplothrips. One species becoming increasingly common in interceptions is Neohydatothrips sama- yunkur (Kudo), which was established re- cently in Kenya. Two other recently estab- lished species—Pezothrips kellyanus (Bagnall) in the Mediterranean region and Echinothrips americanus Morgan in green- houses are dispersing and may become commonly intercepted in the near future. Morphological terminology follows Mound et al. (1976) and Nakahara (1994). Morphological features most often used in the keys include the following (Figs. I—4): Anteroangular setae (Fig. 4): one pair of elongated setae situated on anterolateral corners of pronotum. Anteromarginal setae: one pair of elongat- ed setae situated submedially on anterior margin of pronotum. Basantra: paired praepectal plates on pro- thoracic sternum of species of Phlaeoth- ripidae. Crespeda (Figs. 19-20): usually overlap- ping, scalloped scales or transverse plate on posterior margins of abdominal ter- gites. Ctenidia (Figs. 54, 56): comblike row of short microtrichia on distal margins of a lateral stria on tergite VIII either anterior or posterior of spiracles, also present (and usually somewhat less conspicuous) or absent on tergites ['V—VII. Fringe cilia (Figs. 1-3): elongated gracile hairlike setae arranged uniformly around costal, apical, and posterior margins of fore- and often hindwings. Metafurca (= metasternal furca = meta- 89 furca) (see 15-16): forked endosternal process formed from the inflection of the sternum, and with or without a median anteriorly directed thoracic Figs. process. Microtrichia: usually well defined short hairlike structure on body, antennae, and wings; those on posterior margin of ter- gite VIII being particularly useful in spe- cies (and sometimes generic) identifica- tion. Ocellar setae I, II, Ill (Fig. 4): pairs of elon- gated setae situated near ocellar triangle located anterior to fore ocellus; Il, located anterolaterad of fore ocellus, and III (or interocellar setae), lo- as follows: I, cated usually in triangle formed by ocelli, occasionally laterad of fore ocellus or be- tween hind ocelli. Posterocular setae (Fig. 4): 4—6 pairs of se- tae located on head in a curved row be- hind compound eyes or occasionally sub- medially in posterior part of head. Posteroangular setae (Fig. 4): |—2 pairs of elongated setae located on or near pos- terolateral corners of pronotum. Posteromarginal setae (Fig. 4): setae located on posterior margin of pronotum, usually between posteroangular setae. When setae on posteroangles are not developed, they are also defined as posteromarginal setae. Specimens used in scanning electron mi- crographs (SEMs) were obtained from re- cent incoming material for urgent identifi- cations. Some of the specimens were cleared and slide mounted in Hoyer’s for preliminary identifications, then removed from the slides, placed in 80% ethanol for later preparation for SEMs. Specimens to be prepared for SEMs were transferred overnight into 100% ethanol. They were then critical point dried using a Samway critical point dryer. Dried specimens were glued to paper points attached to SEM stubs and photomicrographed in a scanning elec- tron microscope. Images were digitally cap- tured and transferred to Adobe Photoshop 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 15-26. Morphological features of Thysanoptera, continued. 15, Dendrothrips ornatus, metafurca (lyre- shaped). 16, Limothrips cerealium, metaturca (U-shaped). 17, Neohydatothrips sp., pronotum, dorsal aspect. 18, Apterothrips secticornis, pronotum, showing wider than long. 19, Apterothrips secticornis, abdominal tergites. showing crespeda. 20, Chaetanaphothrips tergite VII, showing glandular area, posteromarginal craspedia, pos- teromarginal microtrichia, and posterolateral margins. 21, Dichromothrips corbetti, head and pronotum, showing 2 pairs of ocellar setae. 22, Odontothrips karnyi, foretibia (showing 1 or 2 clawlike processes. 23, Ceratothri- poides brunneus, sternite VU. 24, Anaphothrips obscurus, head and pronotum, showing 3 pairs of ocellar setae. 25, Ceratothrips ericae, sternite VII. 26, Megalurothrips sjostedti, sternite VII. VOLUME 105, NUMBER 1 9] Figs. 27-36. Morphological features of Thysanoptera, continued. 27, Elaphrothrips sp., head and tho- rax. 28, Bolothrips sp., head and pronotum. 29, Elaphrothrips sp., showing 2 pairs of wing-retaining setae on tergites. 30, Bolothrips sp., showing 1 pair of setae. 31, Gynaikothrips sp., showing absence of maxillary bridge. 32, Haplothrips sp., showing presence of maxillary bridge. 33, Karnyothrips sp., antennal segment IV. 34, Haplothrips sp., antennal segment IV. 35, Karnyothrips sp., basantra. 36, Haplothrips sp., basantra. 92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 5.0° tor MacIntosh, where they were edited for publication quality. Line drawings of specimens were ren- dered from images observed through a camera lucida attached to a Zeiss Axios- kop 2® microscope, using both transmit- ted light and phase contrast modes. RESULTS The 1983-93 database was evaluated to determine the range of species being inter- cepted from Africa, the Mediterranean Re- gion, and Europe over a ten-year period (Table 1). A total of 2,437 interceptions in- cluded 51 genera and 129 identified species distributed among three of the families of thrips: Aeolothripidae, Thripidae, and Phlaeothripidae. In addition, 206 specimens were identified only to genus. Species listed in Table | and occurring in the U.S. are also indicated. The second database (1994-1999) was used to evaluate recent trends in the trans- port of thrips in commerce from Africa and Europe (Table 2). This database was used to evaluate the most common thrips inter- cepted in agricultural and horticultural ship- ments since 1994. In this sample, 497 in- terceptions were represented by 43 identi- fied species in 18 genera distributed among Aeolothripidae, Thripidae, and Phlaeothri- pidae. An additional 67 specimens were identified to genus only. From among the 43 species intercepted between 1994 and 1999, 23 represented nearly 85% of the to- tal number of interceptions (Table 3) and comprised 9 genera. From the list of thrips in Table 1, an illustrated key to nearly all of the represented genera is herein provid- ed. KEY TO PERTINENT FAMILIES OF THYSANOPTERA (Merothripidae and Heterothripidae are omitted because they lack representation in the database sample.) 1. Forewing with veins, surface with microtrichia: terminal abdominal segment normally not tubular; OVIPOSItOMAPESEN Gye zaps noo spe tee oo aee ceed toes Ceratothripoides — Forewing scale with 4 or 5 veinal setae; discal seta present; tergite VIII with posteromarginal comb of microtrichia medially interrupted or absent; B2 and B3 setae on sternite VII on posterior margin (Figs. 25—26) UBileither onion anterior of pOstertOG Mar ow ana = yee ayes Waenen aie! «heise st ete) ete tol ove el =y-eieeatre 36 36. Forewing scale with 4 veinal setae; anterior vein of forewing with 2-3 setae in distal % of vein separated frompnearly, uninterrupted srOw sere ners temevte) cbeiaells) cyt neve neuen oi cements Megalurothrips (Figs. 26, 57-58) — Forewing scale with 5 veinal setae; anterior vein of forewing with 2—4 intermittent setae in distal % SPO TO Geno Ae Rois he pacha nC PEI EES COD CEen iy Ore een OULU ono irae Ret OrniG CeOIENCROES <5, © 37 37. All facets of compound eyes unpigmented; B1, B2, and B3 setae on sternite VII on posterior margin (STEAD) cg aidonG oe a Ao gain Robo OD UA BOM OOO UA Goes Gono Ge Cee OOOO EE Ceratothrips — Compound eye with 5 pigmented facets; B1 setae on sternite VII arising anterior of posterior margin, B2land!B3.0n) postenormmaroiny (ChEIoe26) rn rake @ /el-ueleletaalieekent tote elt Pat olny ted tnd f= Eee Be Tenothrips ACKNOWLEDGMENTS I thank the following individuals for their help in this project: Joseph Cavey, USDA/ APHIS, Hyattsville, MD, and Peter Touhey, SEL, Beltsville, MD, for providing infor- mation for the databases, Sueo Nakahara, SEL, USDA (retired), and Douglass R. Miller and Natalia J. Vandenberg, SEL, USDA, for their comments in reviews of the manuscript. LITERATURE CITED Bhatti, J. S. 1967. Thysanoptera Nova Indica. Publi- cation of author, 24 pp. . 1973. A preliminary revision of Sericothrips Haliday, sensu /at., and related genera, with a re- vised concept of the tribe Sericothripini (Thysan- optera: Thripidae). Oriental Insects 7: 403-449. . 1978. Systematics of Anaphothrips Uzel 1895 sensu latu and some related genera (Insecta: Thy- sanoptera: Thripidae). Senckenbergiana Biologica 59: 85-114. . 1990. The genera Ceratothrips and Tenothrips (Insecta: Thysanoptera: Thripidae). Zoology (Journal of Pure and Applied Zoology) 2: 205— 352% Dyadechko, N. P. 1977. Thrips or fringe-winged in- sects (Thysanoptera) of the European part of the USSR. [Translated for Agricultural Research Ser- vice, United States Department of Agriculture and National Science Foundation]. Amerind Publ. Co., Pvt. Ltd. New Delhi. 1977. 344 pp. Jacot-Guillarmod, C. EF 1970-1977. Catalogue of the Thysanoptera of the world. Annals of the Cape Province Museum 7(1): 1-216; (2): 217-515; (3): 517-976; (4): 977-1255. Kirk, W. D. J. 1996. Thrips. Naturalists’ Handbooks 25. The Richmond Publishing Co. Ltd. Slough, England, 70 pp. Lewis, T. ed. 1997. Thrips as Crop Pests. CAB Inter- national. Wallingford, UK, and New York, NY, USA, 740 pp. Moulton, D. 1948. The genus Frankliniella Karny, with keys for the determination of species (Thy- sanoptera). Revista de Entomologia 19: 55-114. Mound, L. A. and G. Kibby. 1998. Thysanoptera, an Identification Guide. 2nd Edition. CAB Interna- tional. Wallingford, Oxon, UK, 70 pp. Mound, L. A., G. D. Morison, B. R. Pitkin, and J. M. Palmer. 1976. Thysanoptera. Handbooks for Iden- tification of Insects, Vol. I, part 11. Royal Ento- mological Society London, 79 pp. Nakahara, S. 1988. Generic reassignments of North American species currently assigned to the genus Sericothrips Haliday (Thysanoptera: Thripidae). Proceedings of the Entomological Society Wash- ington 90: 480-483. VOLUME 105, NUMBER 1 1822, 183 pp. 389. ton 101: 458—459. Nakahara, S. and G. Vierbergen. 1998. Second instar larvae of Frankliniella species in Europe (Thy- sanoptera: Thripidae). Proceedings of the Sixth In- ternational Symposium on Thysanoptera. Antalya, Turkey. April 27—May 1, 1998. pp. 113-120. Pitkin, B. R. 1972. A revision of the flower-living ge- nus Odontothrips Amyot & Serville (Thysanop- . 1994. The genus Thrips (Thysanoptera: Thri- pidae) of the New World. United States Depart- ment of Agriculture Technical Bulletin Number . 1997. Annotated list of the Frankliniella spe- cies of the World (Thysanoptera: Thripidae). Con- tributions on Entomology, International 2: 355— . 1999. Validation of Neohydatothrips sama- yunkur (Kudo) (Thysanoptera: Thripidae) for a thrips damaging marigolds (Tagetes spp.). Pro- ceedings of the Entomological Society Washing- 99 tera: Thripidae). Bulletin of the British Museum (Natural History) 26: 373—402. Parker, B. L., M. Skinner, and T. Lewis, eds. 1991. Toward Understanding Thysanoptera. Proceedings International Conference on Thrips, February 21— 23, 1989, Burlington, Vermont, USA. United States Department of Agriculture, Forest Service, General Technical Report NE-147, 464 pp. Priesner, H. 1964. Ordnung Thysanoptera (Fransen- fliger, Thripse). Bestimmungsbiicher zur Boden- fauna Europas. Akademie-Verlag, Berlin, 242 pp. Solis, M. A. 1999. Key to selected Pyraloidea (Lepi- doptera) larvae intercepted at U.S. ports of entry: Revision of Pyraloidea in “Keys to some fre- quently intercepted lepidopterous larvae” by D. M. Weisman 1986. Proceedings of the Entomo- logical Society Washington 101: 645-686. Stannard, L. J. 1968. The Thrips, or Thysanoptera of Illinois. [linois Natural History Survey Bulletin 29, Article 4. pp. 215-552. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 100-107 STUDIES OF THE GENUS CULEX LINNAEUS IN FLORIDA I. REDESCRIPTION OF THE PUPAE OF CULEX NIGRIPALPUS THEOBALD AND CX. TARSALIS COQUILLETT, VECTORS OF ST. LOUIS ENCEPHALITIS, AND A KEY TO PUPAE OF CULEX SPECIES IN THE EASTERN UNITED STATES (DIPTERA: CULICIDAE) RICHARD FE. DarsIE, JR. AND JONATHAN E Day Florida Medical Entomology Laboratory. University of Florida, 200 9th Street SE, Vero Beach, FL 32962 (e-mail: rfd@mail.ifas.ufl.edu) Abstract.—The pupal stages of Culex nigripalpus and Cx. tarsalis are redescribed and illustrated. A key to the pupae of 15 species of Culex in the eastern United States is included. Key Words: Culex (Culex) nigripalpus Theobald and Culex (Culex) tarsalis Coquillett are both well known vectors of St. Louis encepha- litis virus (Monath 1980, Day and Curtis 1999; Day and Stark 1999). Cx. tarsalis is also an important vector of western equine encephalitis virus (Reeves and Hammon 1962). It is necessary that all stages of these important species be adequately described, illustrated and recognizable. To that end, the pupae are treated here. The pupa of Cx. nigripalpus was briefly described and figured by Lane (1953). It was illustrated by Belkin et al. (1970) but “not studied in detail.”” Lungstrom (1955) figured the abdomen and metanotum of Cx. tarsalis using the nomenclature of Knight and Chamberlain (1948). He gave no other description. White (1954) the chaetotaxy of the Cv. tarsalis pupa in gen- eral also employing the nomenclature of Knight and Chamberlain (1948) for the ab- domen and that of Penn (1949) for the cephalothorax. In this work, the pupae of both species are described in detail follow- ing the generally accepted nomenclature of Belkin (1962) and Harbach and Knight discussed pupa, Culex nigripalpus, Culex tarsalis (1980). Other life stages were described by Carpenter and LaCasse (1955), Belkin et al. (1970, nigripalpus), Wood et al. (1979, tar- salis). These pupae may be identified using the key to the pupae of the Culex species of eastern United States that appears below and includes 15 of the known 29 species of Culex in the United States. METHODS AND MATERIALS Specimens of Cx. nigripalpus were col- lected in Indian River and Manatee Coun- ties, Florida, reared individually, and larvae and pupal exuviae mounted in Canada bal- sam. For details of the techniques, consult Darsie (1998). Pupae of Cx. tarsalis ex- amined are listed following the description. In the key that follows the undescribed pupa of Culex (Melanoconion) peccator Dyar and Knab was included. Pupae of this species were collected as follows: Florida, Indian River Co., I-16-70, 11 Pe, J. S. Hae- ger, and will be described later. Abbrevia- tions used in the descriptions are br = branches, LePe = larval and pupal exuviae. VOLUME 105, NUMBER 1 DESCRIPTIONS Culex nigripalpus Theobald (Fig. 1) Description.—Positions and sizes of se- tae as figured; ranges and modal branchings in Table 1. Cephalothorax: Mostly tan, scuta, coxae and femora brown to dark brown. Setae 1,3-CT long, usually triple; 2.4-CT moderately long, usually double or triple 7,9-CT long, usually double or triple; 5,8-CT long, usually 4- or 5-branched, 6- CT short, usually double; 10,12 CT long, 10-CT with 8-12 br, 11,12 CT with 2-5 br; Trumpet: Length 0.75—0.82 mm, x 0.79 mm, index 5.35—6.75, < 6.24, tracheoid 0.20—0.31 of length * 0.26; pinna O.16— 0.19, * 0.17 mm, without narrow slit ex- tending from proximal aspect, 0.14—0.24 length of trumpet * 0.19. Abdomen: Length 3.66—4.14 mm, ¥ 3.88 mm; tan, segments I-IV darker, with reticulate pattern in dor- somedian 0.25 of I; sternum II with short, sharp spicules along posterior border. Setae O-II-VU, 9-I-VI, and 14-III-VUI minute, single. Seta 1-I dendritic, with 6—14 pri- mary br, each usually with 9 or more ter- minal br; 1-I[ short with 21—37 fine br: 1-ITI-VII long, III usually with 9 br: mostly quadruple on IV,V; and usually double on VI.VIL. Seta 2-H-VII short, single. Seta 3-I- III rather stout, long on I, moderately long on III, double; 3-IV-VI moderately long, 4—8 br on IV, usually single or double on V-VII. Seta 4-I-VI short, moderately long on VII,VIII, usually 7-branched on I, usu- ally 4- or 5-branched on II,III and V,VI; single to 4-branched on TV, VII, VIII. Seta 5-I short, usually triple, 5-II,III] moderately long, usually with 4—6 br; 5-IV-VI long to very long, stout, 5-IV usually quadruple, 5-V,VI usually double. Seta 6-I very long, single; 6-II-VI long, with 1—3 br on I, mostly double or triple on HI-VI; 6-VII small, with 4—6 br. Seta 7-I,I] long, mostly double; 7-III-V_ short, usually 4- or 5-branched on III,V, double or triple on IV; 7-VI,VIL long, single. Seta 8-II-VII short, pedunculate, usually 3- or 4-branched on 101 Ill, [IV and VI; double or triple on V, VII. Seta 9-VII moderately long, stout, with 3—5 primary br, each ending in 2-3 terminal short br; 9-VIII long, stout, usually acicu- late, with 6—10 primary br, some ending in 2,3 terminal short br. Seta 10-IH-VII long, usually double on III-IV; single (rarely dou- ble) on V-VI, 10-VII single. Seta 11-III-VII short, single on III-VI, 1—3 br on VIII. Fe- male genitalia lobe 0.19—0.24 of paddle, + 0.21; male genitalia 0.28—0.33 of paddle, + 0.30. Paddle: Ovoid, light tan, midrib reaching apex, length 0.43—1.03 mm,x 0.98 mm; index 1.17—1.51, * 1.34; setae 1,2-P short, single. Specimens examined.—FLORIDA, In- dian River Co., Vero Beach, II-29-00, 5 °, 6 do, LePe, from progeny rearings of fe- males collected by power sweeper in veg- etation in wooded area near orange groves (Hussain & Darsie); Manatee Co., Palmetto, VUI-7-2000, 6 2, 2 5, LePe, from progeny rearings of females collected in miniature light trap with CO, (R. Frommer). Culex tarsalis Coquillett (Fig. 2) Description.—Positions and sizes of se- tae as figured; ranges and modal branchings in Table 2. Cephalothorax: Light tan, pos- terior parts of scutum and mesothoracic wing darker brown. Setae 1,3-CT long, usu- ally triple or quadruple; 2,4-CT moderstely long, double or triple, rarely 4-branched; 5-CT prominent, long, triple or quadruple, rarely 5-branched; 6-CT moderately long, with 2-5 br; 7-9-CT long, 7-CT double; 8-CT 3-6 br; 9-CT double or triple; 10-CT long, aciculate, with 5—9 br; 11-CT long, rather stout, double; 12-CT long, mostly double or triple; an adventitious 4-branched seta near 12-CT on one specimen. Trumpet: Conical, dark brown, length 0.62—0.92 mm, ¥ 0.82 mm; index 4.7—6.2, * 5.4; pinna 0.11—0.32 mm, + 0.22 mm and 0.18—0.35 of total length, * 0.24 without narrow slit extending from proximal end, tracheoid 0.17—0.23, of total length, + 0.19. Abdomen: Length 3.47—4.02 mm, + 3.79 mm; light 102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON NmaRscet Fig. 1. by permission). CT—cephalothorax, Pa—paddle. tan, I-IV somewhat darker, without spicules on sternum I. Setae O-II-VII, 9-I-VI and 14-III-VIII minute, single. Seta 1-I dendrit- ic, with 8-13 primary branches, each branch with 5—8 terminal branches; 1-II moderately long, with 5—13 br; 1-II-VI long, with 5—12 br on III, with 4—8 br IV, Pupa of Culex nigripalpus. A, Cephalothorax. B, Metanotum and abdomen (from Belkin et al. 1970, with 3—7, (usually 4) br on V,VI, usually double or triple, seldom with 4 br on VIL. Seta 2-II-VII short, single. Seta 3-I long, rather stout, double; 3-I-VIH moderately long, double on II,III; with 2—8 br on IV; usually double or triple, rarely single or quadruple on V—VII. Seta 4-I-II short, 4-1 VOLUME 105, NUMBER 1 Table 1. branching). 103 Setal branching of the pupa of Culex nigripalpus (range is followed in parentheses by the modal Abdominal Segments Cephalo- Seta thorax I I Ill Vv VI VII Vill IX Paddle 10) — — | 1 | l ] l = — 1 14(3) 6 -14(9) 21-37(27) 5-10(9) 47(6) 3-5(4) 2-4(2) 2-3(2) == 1 | 2 24(2) I-41) l I 1 | | — — l ) 2-5(3) . 2 2 4-8(5) 2 1-2(1) 1-2(2) 1-3(2) — 4 2-6(3) 7-9(7) 2-6(4) 4-8(6) 1-4(1) 3-7(5) 2-4(4)_ 1-2(2) = —- — 5 3-6(5) 2-5(3) 3-7(5) 1-7(4) 2-5(4) 2-3(2) 2-3(2) 1-2(2) — -— — 6 1—3(2) 1-2(1) 1-3(1) 24(3) 2443) -2-3(3)_ 2-43) _—s 4 6(6) — —_= = 7 2A(2) 24(2) 1—3(2) 3-5(4) 24(2) 4-7(5) | | — —_—_ — 8 3-5(4) — — 4-6(4) 46(4) 244(2) 1-5(3) 2442) -— —_- — 9 24(3) | l l | l 3-5(4) 6-107) — — 10. 7-10(8) — — 1—3(2) 2-3(2) 1-2(1) 1 | — —_ — 1] 2 -= — 1 l l 1-3(1) = —_—- — 12 4-6(5) ae — - 14 | | | | l a — with 5—9 br, 4-IT, TI] usually with 5 br; 4-IV usually double or with 3—5 br; 4-V-VIII me- dium long, usually with 4,5 br on V,VI; 4-VIL, VIII single to triple, rarely quadruple. Seta 5-I short, usually with 3—5 br (2-7); 5-II-III moderately long, usually with 4,5 br, seldom with 6 or 7, 5-[V-VI long, stout, double or triple; 5-VII long, usually double. Seta 6-I,II very long, single; 6-[I-VI long, usually triple, 6-VII short, mostly with 6—8 br, rarely quadruple. Seta 7-[,II long, single or double; 7-III short, usually with 6 br: 7-IV,V short, usually with 4 or 5 br, 7-VI,VII moderately long or long, single. Seta 8-IH-VII short, pedunculate, usually with 3—6 br. Seta 9-VII long, stout, usually aciculate, with 4—6 br, 9-VIII long, stout, aciculate, with 7—9 br, rarely with 2 br api- cally. Seta long, usually double, 10-V-VII long, single. Seta 11-VII short, with 1—4 br. Paddle: Ovoid, length 0.95—1.35 mm, * 1.19 mm; index 1.35—1.56, ¥ 1.42; seta 1-P short, sin- gle, seldom double or triple; 2-P short, sin- gle. Specimens examined.—MONTANA: Phillips Co., Malta, VII-9-72, 2 9, 1 6, LePe, (Quickenden, Darsie); Ravalli Co., Hamilton, VIII-54, 1 2, 1 6, LePe, USS. Public Health Service Rocky Mountain 10-I11,.1YV moderately long or Laboratory colony (J. M. Brennan). CAL- IFORNIA, San Diego Co., V-7-49, 2 2, 1 3d, LePe; Otay R., V-8-49, 1 2, LePe; Lake Henshaw, V-7-49, 2 2; Lake Cuyamaca, V-7-49, 1 @ Pe (Heid); Los Angeles Co., Agoura, HI-22-49, 1 6, Pe (Heid). DISCUSSION The pupa of Cx. nigripalpus was briefly described by Lane (1953), employing the setal nomenclature of Baisas (1935) as modified by Penn (1949). He stated that the cephalothorax has small tufts, while our ac- count gives accurate branching of each seta. He described the “tube,” 1.e., three times basal width; in Florida Cx. ni- gripalpus pupal trumpets are 7.8—11.1 * 8.9 times longer than basal width. Perhaps Lane was not measuring the same species. In all, 13 abdominal setae were characterized by Lane. His setae 6-II-VI were triple; in our study seta 6-II was usually single, and setae 6-IH-VI double to quadruple. Lane called five other setae (1-II, 5-IV,V, 9-VII,VIIT) ‘multiple’? without precise branching which we include. Seta 5-VI,VII are listed as double; in our study they are single to triple, although modally double. The pad- dles in our pupae are 1.9—2.3, ¥ 2.1 times trumpet, as 104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mr \ Pa Prollie AponleR Tealdig » Zoo1 1g32 d af Fig. 2. Pupa of Culex tarsalis. A, Cephalothorax. B, Metanotum and abdomen. BU—external buttress, CT— cephalothorax, GL—genital lobe, Mr—midrib of paddle, Pa—paddle, T—respiratory trumpet. VOLUME 105, NUMBER | 105 Table 2. Setal branching of the pupa of Culex tarsalis (range is followed in parentheses by the modal branching). Abdominal Segments Cephalo- Seta thorax I u Il IV Vv VI VIL VIII IX Paddle 0 —- — 1 ] 1 1 i | ] — a 1} 2-5(3) 8-13(12) 5-13(7) 5-12(7) 4-8(5) 3-7(4) 3-6(4) 244(2) — 1-3(1) l 2 2-3(3) 1—2(1) 1-2(1) | 1 | 1 | — — | 3 3-6(4) 2 2 2 2-8(5) 1-3(2) 2-4(2) 1-3(2) — —- — 4 24(2) 5-9(5) 2-7(5) 4-6(5) 2-5(2) 3-6(5) 2-5(4) 1-3(2) 24(2) — — 5 (3=5(4) 2-7(5) 4-7(5) 4-8(5) 3-5(4) 2-3(2) 2-3(2) 2-3(2) — — — 6 2-5(4) 1 1 2—3(3) 5 3-4(3) 2473) 48(8) a — — 7! 2 1—2(2) 1—2(2) 3-7(6) 2-4(4) 3-70 1 ] -= _— — 8 3-6(5) — — 2-6(5) 2-4(3) 244(2) 1443) 3-5(3) — — — 9 1-3(3) 1—2(1) 1 1 1 1 1 4-6(4) 5-10(8) — — 10 5-9(6) —_— — 1-3(2) =: 1-2(2) ] 1 1 — — 1] 2 oo — | 1 1—-2(1) 1 1-4(1) — — —_— 12 2-5(3) 14 1 ] ] ] 1 ] -— — length of segment VIII, but Lane reported = — Seta 5-CT no more than 2.0 length of the paddle three times lengfh of VIII. 4-CT; abdominal tergum VIII with pos- terior lobe not overlying lateral part of On the other hand, Belkin et al. (1970) : : : i tergum IX; seta 1-I[X present ........ 2 includes a complete illustration of the pupa 2(1). Meatus of trumpet with narrow slit ex- and mentioned only the trumpet not flared, tending from proximal part of pinna; seta its pinna small, the spicules on the posterior 2-VI usually laterad of seta 1-VI (subge- margin of stermun II and seta 5-IV usually Bs WAETAUOCOMOR) oa. a asa she nter mee with 4 branches. Though few, all agree with =~ Mica Sie Semen onbria iow SUuer ee ia x tending from proximal part of pinna; seta our findings 2-VI mesad of seta I-VI ........... 9 Lungstrom (1955) and White (1964) il- 3(2). Seta 9-VIII very near posterolateral cor- lustrated the abdomen of Cx. tarsalis using MCS ee ee tate este cedecei Stone and Hair the Knight and Chamberlain (1948) nomen- — ~ Seta 9-VII distinctly removed from pos- clature. They are of questionable quality Seg STE CORREE are ate anes gs : an : 4(3). Seta 5-V double or triple and nearly as and neither author offered textual descrip- ae ae oe ~ ‘ ong as following tergum ........... 5 tion. - Seta 5-V usually with at least 4 branches, if fewer, then distinctly shorter than fol- KEY TO THE PUPAE OF CULEX IN THE Meonnetereuin ey eno oe 6 EASTERN UNITED STATES 5(4). Seta 1-II with 14 or fewer branches; pinna The key was formulated mostly using de- of trumpet, including slit, less than 0.3 of total length ..... pilosus (Dyar and Knab) scriptions by Darsie (1951), Foote (1954), Belkin et al. (1970) and Zavortink and - Seta 1-I with 25 or more branches; pinna of trumpet including slit 0.4—0.5 of total O’Meara (1999), and specimens from the length: 5... 2). erraticus (Dyar and Knab) Florida Medical Entomology Laboratory 6(4). Seta 6-IV,V at least 4-branched; seta 5-V collection. For assistance in using this key, usually 5-branched; trumpet index usually iS SOD, aR ebcia done atratus Theobald see Figs. 1-2. 2 - Seta 6-IV,V usually triple: seta 5-V usu- i Seta 5-CT very long, =5.0 length of seta ally 4-branched: trumpet index =8.0 ... 7 4-CT: abdominal tergum VII with pos- 7(6). Pinna including slit =0.35 length of trum- terior lobe overlying lateral part of ter- pet: seta 8-CT single .. mulrennani Basham gum IX; seta 1-IX absent (subgenus Mi- = Pinna including slit =0.4 length of trum- CTACAES) 9 Mitre the heehee 2 ee, Abe, See pet; seta 8-CT with 3 or more branches 106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Trumpet lighter in color between apex of tracheoid and base of pinna, flared api- cally iolambdis Dyar - Trumpet uniformly dark distal to apex of tracheoid, or if lighter, then remainder of trumpet also lighter, not flared apically oo SOS os Ope eee peccator Dyar & Knab 9(2).6 Trumpet index =7.7; seta 9-VII usually double; seta 9-VIII usually 4-branched (Subgenus Neoculex) .....- territans Walker - Trumpet index =7.0; seta 9-VII usually with 4 or more branches; seta 9-VIIT usu- ally 6-branched or more (subgenus Culex) 10(9). Abdominal tergum I with distinct short spicules posteriorly ins Brce ee bahamensis Dyar and Knab - Abdominal tergum I without short spic- ules posteriorly... tess 2 pean == 11 11(10). Seta 2-P absent on paddle; seta 1,6-VI usually double restuans Theobald - Seta 2-P present, rarely absent: seta 1,6- VI usually with 3 branches or more .. 12 12(11). Trumpet index rarely * 5.0; seta 1-VII usually with 4 or more branches ... - sfSades REN Per eds quinquefasciatus Say - Trumpet index =5.5; seta 1-VII usually doublevonitriplescyecctee poate eye 13 13(12). Seta 1-If with 20 or more branches; pos- terior border of sternum I with sharp, short spicules nigripalpus Theobald = Seta 1-II with 17 or fewer branches; pos- terior aspect of sternum II without spic- WES yh Mya otis cee adept Cet oe 14 14(13). Trumpet index about 7.0, pinna short, <0.13 of total length . . salinarius Coquillett - Trumpet index =6.0. pinna longer, =0.16 ofitotal lengthens. co. o.x-- tarsalis Coquillett ACKNOWLEDGMENTS We are indebted to W. J. Tabachnick and N. Hussain for their support and assistance. The manuscript was reviewed by W. J. Ta- bachnick and C. R. Rutledge. This is Flor- ida Agricultural Experiment Station Journal Series No. R-08423. LITERATURE CITED Baisas, F E. 1935. Notes om Philippine mosquitoes V. The pupal characters of anophelines under the Myzorhynchus series and group Neocellia, with further comments on the larvae and adults of si- nensis. Monthly Bulletin of the Philippine Bureau of Health 15: 291-339. Belkin, J. N 1962. The mosquitoes of the South Pacific (Diptera, Culicidae). Volumes I and II. University of California Press, Berkeley and Los Angeles XII + 608 pp., 412 figures Belkin, J. N., S. J. Heinemann and W. A. Page. 1970. Mosquito studies (Diptera, Culicidae) XXI. The Culicidae of Jamaica. Contributions of the Amer- ican Entomological Institute 6(1): 1—458. Carpenter, S. J. and W. J. LaCasse. 1955. Mosquitoes of North America (north of Mexico). University of California Press, Berkeley, 360 pp. Darsie, R. F Jr. 1951. Pupae of the culicine mosquitoes of the northeastern United States (Diptera, Culic- idae). Cornell Agricultural Experiment Station Memoir 304, 67 pp. . 1998. Description of the pupae of six species in subgenus Leicesteria, genus Armigeres, from Nepal (Diptera, Culicidae). Proceedings of the En- tomological Society of Washington. 100: 234— 246. y, J. EK and G. A. Curtis. 1999. Blood feeding and oviposition by Culex nigripalpus (Diptera: Culic- idae) before, during and after a widespread St. Louis encephalitis epidemic in Florida. Journal of Medical Entomology 36: 176-181. J. FE and L. M. Stark. 1999. Avian serology in a St. Louis encephalitis epicenter before, during and Day, after a widespread epidemic in south Florida, USA. Journal of Medical Entomology 36: 614— 624. Foote, R. H. 1954. The larvae and pupae of the mos- quitoes belonging to the Culex subgenera Melan- iconion and Mochlostyrax. Technical Bulletin, United States Department. of Agriculture No. 1091, 126 pp. Harbach, R. E. and K. L. Knight. 1980. Taxonomists’ Glossary of Mosquito Anatomy. Plexus Publish- ing, Inc., Marlton, 413 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. Lane, J. 1953. Neotropical Culicidae. Brazil, Sao Pau- lo, 2 Vols., 1,112 pp. Lungstrom, L. 1955. Additional observations on the recogition features of the Culex tarsalis larva and pupa (Culicidae: Diptera). Transactions of the Kansas Academy of Science 58: 68-74. Monath, T. P. (Ed.). 1980. St. Louis Encephalitis. American Public Health Association, 679 pp. Penn, G. H. 1949. The pupae of the mosquitoes of New Guinea. Pacific Science 3: 3-85. Reeves, W. C. and W. McD. Hammon. 1962. Epide- miology of the arthropod-borne viral encephaliti- des in Kern County. University of California Pub- lications on Health 4: 1-257. VOLUME 105, NUMBER 1 White, J. M. 1954. Chaetotaxy of the pupa of the mos- quito Culex tarsalis. Proceedings of the Oklahoma Academy of Science for 1954, pp. 30-32. Wood, D. M., P. T. Dang, and R. A. Ellis. 1979. The mosquitoes of Canada (Diptera: Culicidae). Se- ries: The insects and arachnids of Canada. Part 6. 107 Biosystematics Research Institute, Canada De- partment of Agriculture Publication 1686, 390 pp. Zavortink, T. J. and G. F O’Meara. 1999. Culex (Mi- craedes) biscaynensis n.sp. from Florida (Diptera: Culicidae). Journal of the American Mosquito Control Association 15: 263-270. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 108-119 SUBCORTICAL CAVITY DIMENSION AND INQUILINES OF THE LARVAL LOCUST BORER (COLEOPTERA: CERAMBYCIDAE) KRISTIN ANN LARSON AND DAN MYERS HARMAN Appalachian Laboratory, 301 Braddock Road, Frostburg, MD 21532, U.S.A. (e-mail: kranlars@ yahoo.com; harman @al.umces.edu) Abstract.—Subcortical, pre-tunneling cavities made by the locust borer (Megacyllene robiniae Forster) on the black locust (Robinia pseudoacacia L.) tree were studied in western Maryland on three site types: strip-mine, roadside, and old field-pasture. A num- ber of invertebrates, associated with sap, frass, and boring dust within the cavities, were collected and identified. The more common cavity taxa included two suborders of mites, three nitidulid genera, nematodes, and two dipteran families. Mean numbers of mites were significantly greater in strip-mine sites than in roadside and old field-pasture sites. Cavity size was not related to abundance or diversity of cavity taxa, and cavity taxa did not appear to affect locust borer survival. Key Words: The locust borer (Megacyllene robiniae Forster) is a severe and persistent pest of the black locust (Robinia pseudoacacia L., Leguminosae) tree throughout its natural and extended range in North America. Its original range was thought to have been limited to the Appalachian chain from Pennsylvania to Georgia and west to the Ozark Mountain region (Cuno 1930). The locust borer is not present in Europe, how- ever, where black locust has been planted extensively. The borer is univoltine (one generation per year), primary (invading only living trees), and host specific (invad- ing only black locust). Locust borers deposit eggs into the bark crevices of black locust tree trunks in later summer and fall. Tiny, newly hatched lar- vae penetrate the inner bark, where they overwinter. In the following spring, they re- sume their tunneling activities, and enlarge and sapwood forming cavities of variable size. From its cavity, a larva extends its tunnel subcortical excavations in cambium Megacyllene robiniae, inquilines, cavities, survival, Robinia pseudoacacia into sapwood and heartwood. The larva maintains the tunnel opening, through which it pushes excess boring dust and frass during development, and through which it emerges as an adult after pupating within its tunnel (Hopkins 1907). The host tree at- tempts to close a borer wound by depositing tissue around its perimeter; however, a healthy larva normally maintains and en- larges both its tunnel entrance and its cav- ity. Hall (1942) observed larvae restricting activities to cavity-making, without subse- quent tunneling, during a drought year. As an adult attraction to flowers is a be- havioral feature of many cerambycids, lo- cust borer adults are commonly seen con- gregating on blossoms of goldenrod, Soli- dago spp., particularly S. altissima. While on these flowers, in close association with stinging wasps and other vespids, borer adults appear to exhibit Batesian mimicry (Garman 1916, Blackwell and Powell 1981). Madden (1996) stated that when handled, borer stridulation also mimicked VOLUME 105, NUMBER | that of vespids. Similarly, we have noticed that when handled, borers stridulate rapidly in an apparent excitation mode, and emit an unpleasant odor, apparently in defense. We also have verified the persistent pres- ence of certain insect groups, particularly those in the Diptera and Coleoptera (Niti- dulidae), displaying an attraction to sap and frass exudation at trunk surfaces near locust borer entrance holes. McCann (1992) iden- tified adult insects collected from trunk sur- faces on or near (within 10 cm) the entranc- es of active borer mines, however, his stud- ies did not include the opening of cavities, or any inquiry into inquilines within cavi- ties. He reported nine coleopteran species, representing five families. Little other in- formation is available in the literature. In the present study, outer tree bark was chipped away to fully expose the cavities and analyze their contents. This procedure revealed a protected habitat with a sap and frass food source, inhabited by a guild of other invertebrate species. Predators and parasitoids appear to have only limited effects on locust borer popu- lations. Locust borer studies have docu- mented predation by a garden spider (Ar- giope aurantia Lucas) on adults (Van Tyne 1983, Harman and Harman 1987, Echaves et al. 1998), woodpeckers on larvae (Hall et al. 1938, Hall 1942, McCann 1992), and ants on eggs (Echaves et al. 1998, Van Tyne 1983). In an unpublished three-week study, from late August through early September, D. Harman and J. McCann observed no in- stances of avian predation in a field of gold- enrod, heavily used by locust borer adults. For this study, an observer sat at the edge of field with binoculars prepared to identity any birds approaching goldenrod blossoms in the field. Observations were conducted for two hours each, in morning (7—9 a.m.), midday (11-1 a.m.), and evening (6-8 p.m.) for 21 days (the last week in August through the second week in September) in 1989. During the borer larval and pupal stages, some predation by woodpeckers oc- curs. The predation is usually light and 109 varies locally (McCann 1992). The same was observed for predation by garden spi- ders and ants. The term inquiline has gained entry into recent biological literature, defined with slightly different slants. An ecological glos- sary describes the term as “*... a type of symbiosis in which one or more organisms exists in the burrow, nest, or abode of an- other without harming the host” (Lincoln et. al 1986). The term is used for various situations including abodes of mammals, birds, and insects. Inquiline biology ad- dresses a range of biological concepts in- cluding such items as phenological timing for nutritional resources (Shibata 2001, Ehason 2000), and the roles of predation and food limitation in species abundance (Kneitel and Miller 2000). This study ad- dressed inquiline response to continuous wounding of living tree tissue by a wood- boring larva, which created the food source. The cavity inhabitants reported in this study are added to the list of organisms known to live close to borer larvae and may potentially affect or influence larval devel- opment. The objectives of this study were: (1) to identify inquilines in locust borer cavities, (2) compare their populations among different habitats and cavity sizes, and (3) to determine whether these inqui- lines affect locust borer larvae. In over- view, this study addressed the subject of strip-mine reclamation and revegetation, as black locust is an important reclamation species. An extensive literature review, which narrowed the scope of the investi- gation, included few studies addressing bi- ological control of the locust borer. MATERIALS AND METHODS Sites that provided a variety of condi- tions were selected for sampling. Three site types differing in soil disturbance levels were in decreasing order of disturbance: strip-mine, roadside, and old field-pasture. A total of nine sites (three of each site type) were selected within a 50 km radius of Frostburg, Maryland. These habitat types 110 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON were selected because they were seen as the most prominent and distinct habitats sup- porting growth of black locust, whose ecol- ogy is typically that of a “pioneer” species on open lands. As a minimum size require- ment, each site contained an area large enough to accommodate a 15 m * 15 m study plot containing 50 or more trees of approximately 10—20 years of age. Plot cor- ners were established and tree positions were numbered and mapped. In each plot, black locust was the predominant woody species, and stocking densities were similar. A total of 288 samples of bark sections containing locust borer cavities were re- moved from each of the nine sites during the period of active larval tunneling, late May through July, 1995. Borer activity was identified by the presence of yellow sap, boring dust, or both, emanating from tun- nels and visible on trunk surfaces. Four ac- tive larval cavities per site were taken weekly from randomly selected trees. Each tree had two rectangles of bark, approxi- mately 2.5 * 5.0 cm, containing locust bor- er cavities removed. The bark was carefully removed using a chisel and ax handle, and then placed in a plastic bag. Bark adjacent to the selected cavities was removed, when necessary, to include all parts of larger cav- ities. Each cavity was placed in a petri dish and examined under a dissecting micro- scope within 24 hours. To locate all inver- tebrates, samples were broken into small sections after initial examination. Cavity 1n- vertebrates were counted, sorted, and placed in 80% ethanol for further identifi- cation. A second major approach involved trap- ping. Wire cages stapled over cavities, tun- nel entrances, or both were used to capture locust borer adults and associated cavity- dwelling invertebrates as they emerged later in the season. The traps were constructed by folding and stapling finely meshed cop- per screen into teepee-like forms of approx- imately 4.0 cm wide * 8.0 cm long. Traps were in place from late June through July, 1995, on trees 5-15 cm in diameter at breast height (dbh). Trap edges were caulked to prevent the escape of inhabi- tants. For trapping, each of the nine sites was subdivided into thirds, on each of which 15 traps were placed on randomly- selected trees, totaling 405 traps. The num- ber of traps per tree varied from one to five, depending upon numbers of active tunnels. Whenever visible, boring dust was removed from traps and examined for the presence of invertebrates, which were counted and preserved. Entomologists from the Smith- sonian Institution, the USDA Systematic Entomology Laboratory, and Frostburg State University assisted in identification of cavity and trap inhabitants. However, spe- cialists were not available for all families and for some, the other sex, life stage, or caste was needed for further determination. For each cavity, cavity length, width, depth, and adjacent bark thickness were measured at their greatest points, and vol- ume was calculated from these measure- ments. Field observations indicated that the larvae excavated the cavities prior to tun- neling, and that the cavities increased in size only slightly, if at all, after tunneling began. If a tunnel was present, the tunnel diameter was also measured. Success of borer attacks was evaluated using Mc- Cann’s (1992) criteria and larval status was recorded as “‘died as an early instar” if little or no tunneling occurred beyond the cavity; “died after tunneling” if tunneling extend- ed into the sapwood, but tunnel entrance diameter measured less than 4 mm; and “emerged as an adult” if tunnel entrance diameter exceeded 4 mm. All trees within the study plots were mapped and individually numbered. Height, dbh, and resistance to pulsed electric cur- rent (an index of vigor) were measured for all trees from which cavities were removed. Three electrical resistance readings per tree were taken in mid-September over a 1- week period, and averaged. The Shigome- ter® (Model OZ-67, Osmose® Wood Pre- serving Co. of America, Inc., Buffalo, NY) provided an estimate of tree vigor by deliv- VOLUME 105, NUMBER | ering a 0.5-A pulsed electric current to the cambium layer (Carter and Blanchard 1977). More healthy trees show lower elec- trical resistance readings than less healthy ones (Shigo 1982). Readings were taken | m above ground level. To obtain a mean site age, three trees from which cavities were removed were randomly selected for each of the nine sites. Ages were obtained from increment bor- ings. Site indices were calculated from a set of black locust curves (Kellog 1936) for each of the three site types using the heights of trees and mean site age. Site quality re- lates to the growth in height of most com- mercial tree species. For a specified age, sites of better quality produce taller trees and a higher index value (Wenger 1984). Statistical analyses.—The Statistical Analysis System (SAS) (SAS Institute 1988) was used for all statistical analyses. Fisher’s least significant difference (LSD) test was used to find possible differences among means of site type and cavity size, site type and tree measurements, and cavity size and borer survival. Cavities of insuf- ficient dimension were excluded from sta- tistical tests. Fisher's LSD test was also used to look for significant differences among means of site type and numbers of species. Only the four most frequent cavity taxa were included in the ANOVA. Chi- square tests were used to compare cavity size Class versus borer survival, and versus site type (SAS Institute 1988). Chi-square tests were also used to compare tree vigor, indicated by the classes of electrical resis- tance readings, and borer survival. Linear regressions compared cavity volume against total invertebrates, and against the four more frequent cavity taxa. Pearson’s correlation coefficient tested numerical re- lationships among the four more frequent cavity taxa. RESULTS Cavity taxa.—Invertebrates collected from tree cavities represented 12 orders, 20 families, an estimated 50 genera, and about the same number of species. Invertebrate inquilines starting with the most abundant taxon were mites (Astigmata (97% of all mites) and Mesostigmata), nematodes (Rhabditida and possibly others), nitidulid larvae (primarily Glischrochilus spp. and Crytptarcha spp.), odiniid larvae—with the preceeding two larval families found in all nine sites, three collembolan families, xy- lomid larvae (Solva sp.), Armadillidium sp. (Isopoda), Staphylinidae, ants (Formicidae), and Miridae (Table 1). Trap taxa.—Invertebrates collected from boring dust in traps on trunks represented 13 orders, 30 families, an estimated 53 gen- era, and about the same number of species. Invertebrate inquilines starting with the most abundant taxon were nitidulid larvae (primarily Glischrochilus spp. and Cryptar- cha spp.), xylomid larvae (Solva sp.), mites (Astigmata and Mesostigmata), dermapter- ans (Forficulid sp.), collembolans (Poduri- dae), Armadillidium sp., chloropid spp., clerid larvae, and nematomorphs (Gordioi- dea) (Table 2). Numbers of species.—Species per cavity ranged from zero to six. The four more fre- quent cavity taxa followed by the percent- age of cavities that they were found in were mites (66%), nitidulid larvae (47%), nem- atodes (23%), and odinid larvae (21%). Numbers of species, which included only the four more frequent cavity taxa, were not significantly different among site types (P > 0.05). Mean numbers of nematodes, ni- tidulid larvae, and odiniid larvae were also not significantly different among site types (P > 0.05). However, mean numbers of mites were greater on strip-mine sites (F = 3.70, df = 2,186, P = 0.027) than roadside and old field-pasture sites. Pearson’s Cor- relation Coefficient showed a weak positive relationship between mites and odiniid lar- vae per cavity (R = 0.36861, P = 0.0078). Cavity volume and taxa.—The regres- sion of the total number of insects per cav- ity versus cavity volume did not show a sig- nificant linear relationship (P > 0.05), nor did the regressions of the numbers of indi- 112 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. The more common invertebrate taxa extracted from subcortical cavities in black locust trees, show- ing numbers of individuals by site type, i.e., strip-mine, roadside, and old field-pasture. This table includes taxa occurring in numbers greater than five, represented in all three site types, or both. Other less common taxa, with their total occurrences, and specific ant spp. include Gastropoda, Stylommatophora, Haplotrematidae, Haplotrema concavum (Say) (1); Arachnida, Araneae, Thomisidae (2), Agelinidae (1), Salticidae, Habrocestum pulex (Hentz) (1); Acari, Oribatida (4), Prostigmata (1); Diplopoda (1); Insecta, Coleoptera, Cleridae, Enoclerus sp. (1), His- teridae, Hololepta fossularis Say (2), Coccinellidae (1), Trogositidae (1); Dermaptera, Forficulidae (2); Psocop- tera (3); Hemiptera, Anthocoridae, Calloidis temnostethoides Reuter (4); Hymenoptera, Encyrtidae (1), Formi- cidae, Solenopsis molesta Say (4). Crematogaster lineolata (Say) (2), Leptothorax ambiguus Emery (1), tetramorium caespitum (L.) (1), Tapinoma sessile (Say) (1). Possible Mesostimgata include Uropodidae, Ame- roseiidae, and Parasitidae. Immature Nitidulidae are primarily Glischrochilus and Cryptarcha spp., with some Soronia spp. Solva pallipes (Loew) identified as most likely sp. as it is more common than the other Solva sp. in eastern North America (S. crepuscula Hull). el Taxon Strip-Mine Roadside Old Field-Pasture Total Nematoda Secernentea, Rhabditida 1,378 770 890 3,038 Arthropoda Malocostraca, Isopoda, Armadillidium sp. 16 1 17 Arachnida, Acari Astigmata 4,879 627 2,124 7,630 Mesostigmata 123 44 81 248 Insecta, Collembola Entombryidae/Isotomidae 6 16 28 50 Poduridae fl 2 8 17 Hemiptera, Miridae Lopidea robiniae Ubler 4 1 | 6 Coleoptera Staphlinidae Immature 2 6 8 Adult 3 3 6 Nitidulidae Immature 182 171 115 468 Adult 8 5) 1 14 Carpophilus lugubris Murray 1 1 Erotylidae, Megalodacne heros (Say) ] 1 | 3 Diptera, Xylomidae Solva pallipes (Loew )—immature 9 4 9 22 Odiniidae—immature 30 27 ay 109 Hymenoptera, Formicidae 3 6 ©) viduals of the four more common cavity taxa versus cavity volume. Cavity size and borer survival.—Borers survived to the pupal stage and presumably emerged from 50% of cavities, tunneled into sapwood and died in 13% of cavities, and died before creating a tunnel in 37% of cavities. Some size variation occurs among locust borers, in relation to sex (females larger than males) and other unknown fac- tors, but this feature was not compared with cavity size. When comparing mean cavity measurements and survival of borers, the cavities of adult borers that emerged were significantly longer, wider, deeper, and greater in volume than those of larvae that VOLUME 105, NUMBER 1 113 Table 2. The ten more common taxa from screen traps placed over borer cavity entrances and the types of sites in which they were located. Other, less common taxa and numbers of each include Gastropoda, Haploter- matidae, Haplotream concavum (Say) (1); Arachnida, Araneae, Agelenidae (1), Agelenopsi sp. (1), Clubionidae, Clubiona pallens (Hentz) (1), Salticidae (1); Insecta, Coleoptera, Erotylidae, Megalodacne heros (Say) (3). Histeridae (1), Staphylinidae (5), Trogositidae (3), Nitidulidae, adults (6); Collembola, Entombryidae/Isotomidae (3); Hemiptera, Anthocoridae Calloides temnostethoides Reuter (3); Homoptera, Coccidae (1), Membracidae, immature (2), Psyllidae (1); Neuroptera, Chrysopidae, Chrysoperla rutilabris (Burm.) (2); Diptera, Anthomyiidae (1), Ceratopogonidae, Forcipomyia sp. (1), Loncheidae (1), Odiniidae, immature (4), Traginops sp. adult (1); Drosophilidae (8), Phoridae (1), Muscidae (1), Tachinidae (1); Lepidoptera, immature (4); Hymenoptera (1): Proctotrupoidea (2), Formicidae (1), Formica subsericea Say (1). Immature Nitdulidae are primarily Glischro- chilus and Cryptarcha spp., with some Soronia spp. Solva pallipes (Loew) identified as most likely sp. as it is more common than the other Solva sp. in eastern North America (S. crepuscula Hull). Gaurax sp. near G. pseudostigma Johnson, likely a new species. S = strip-mine, R = roadside, O = old field-pasture. Total Taxon Site Types No. Sites Individuals Nematomorpha Gordioidea S.R 2 5 Arthropoda Malacostroca Isopoda, Armadillidium sp. S.R 3 16 Arachnida Acari, Astigmata 8,0. 3 44 Mesostigmata S | 16 Insecta Collembola, Poduridae S 2 28 Coleoptera, Nitidulidae—immature S.R.O 9 184 Cleridae, Enoclerus spp.—immature S,R.O 4 11 Dermaptera, Forficulidae Forficula auricularia L. S.R,O 5 31 Diptera, Xylomidae Solva pallipes (Loew )—immature S.R,O 9 82 Chloropidae R,O 7 I4 (Fiebrigella catalpae Mallach—11) (Coniscella hinkleyi Malloch—2) (Gaurax sp.—1) died shortly after entering the xylem, and cavities 5O1—1,000 mm} (class 3). Cavities those that died after entering sapwood. Mean bark thickness was not significantly different among the three groups (Table 3). Chi-square test results revealed signifi- cant differences in survival of borers among cavity size classes (x? = 115.526, df = 8, P = 0.001). Over 95% of borers emerged from cavities =1,001 mm? (class 4 = 1,001—2,000 mm? and class 5 = 2,001-— 6,000 mm*), while only 13% of borers emerged from cavities =100 mm) (class 1). Sixty-eight percent of borers emerged from from 101-500 mm) (class 2) showed little differences among the 3 borer classes (Fig. I} Cavity size and site type-—When com- paring the means of cavity measurements by site type, cavities in strip-mine sites were significantly larger than roadside and old field-pasture sites in terms of length, width, depth, and volume. Cavities re- moved from old field-pasture sites were sig- nificantly greater in length than those from roadside sites; those from roadside sites 114 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Mean + standard error comparison of cavity measurements by adult borer status. Means not sharing the same lower case letter within columns are significantly different (P < 0.05, Fisher’s least significant differ- ence test). Cavity Length Width Depth Volume Bark Thickness Borer Status (mm) (mm) (mm) (mm?) (mm) Emerged as an adult 15.4 + 0.5a 17.0 + 0.6a 44+ 0.2a 1,193.4 + 80.7a 10.9 + 0.3a Died after tunneling IVP ea Les lho) 8.8 + 0.7b 258-025 285.2 + 43.9b 10.6 + 0.6a Died as an early instar TOF s087b 9.3 + 0.6b 1S!) 0b 226.0 + 34.1b 10.4 + 0.4a ANOVA F 9.63 49.32 61.3 Sill 0.56 df 2, 284 2, 284 25,270 2, 270 22 RP 0.0001 0.0001 0.0001 0.0001 >0.05 were significantly deeper than those from old field-pasture sites. Mean bark thickness was significantly less in old-field pasture sites than on roadside and strip-mine sites (Table 4+). Chi-square tests revealed signif- icant differences between cavity size clas- ses and site types (x? = 30.115, df = 8, P = <0.0001). Strip-mine sites had fewer cavities in the smallest cavity class (= 100 mm), and more cavities in the two larger cavity classes (1,001—6,000 mm‘) com- pared to roadside and old field-pasture sites (Fig. 2). Tree measurements and site type.—Mean tree heights (m) were significantly greater (F = 31.60, df = 2,285, P = 0.0001) in old field-pasture sites (7.7 + 0.2) than on strip- mine (6.1 + 0.1) and roadside sites (6.0 + 0.2); whereas, mean dbh (ranging from 5.9-6.6 cm) was not significantly different among site types (P > 0.05). Mean tree age (yr) was significantly greater (F = 3.41, df = 2,243, P = 0.035) in old-field pasture sites (19.0 + 0.8) than in roadside sites (16.5 + 0.7), but not strip-mine sites (17.5 + 1.1). Mean electrical resistance readings (kohms) on strip-mine (16.6 + 0.9) and roadside sites (16.0 + 0.6) were signifi- cantly lower (F = 6.23, df = 2,277, P = 0.002) than readings on old field-pasture sites (19.2 + 0.8). In contrast, old field-pas- ture sites had a higher site index value (37) than strip-mine (31) and roadside sites (32). A chi-square test indicated no significant differences in survival of borers among the four classes of electrical resistance readings (x? = 7.680, df = 8, P = 0.465). DISCUSSION Cavity inquilines did not appear to have a negative effect on locust borer popula- tions. Therefore, use of the term inquiline is qualified as no direct evidence of cavity inhabitants harming the abode-building host was found. However, little is known about the habits of many of these organisms, es- pecially larvae. Nitidulid larvae are sa- prophagous in general, but may feed on fungal fruiting bodies or spores, and may be predaceous. The three genera found in borer cavities, Cryptarcha, Glischrochilus, and Soronia, are among genera that occur most often under fermenting bark or in ran- cid sap (Stehr 1991). McCann (1992) found three Glischrochi- lus species in or within 10 cm of borer cav- ities. The species were G. samquinolentus (Olivier), G. quadrisignatus (Hay), and G. fasciatus (Olivier). Since nitidulid larvae in this study were not identified to species, they may have been represented by the Glischrochilus spp. McCann (1992) found. Cryptarcha species in this study may have been represented by C. ampla (Erichson), also found by McCann (1992). Cryptarcha spp. are known to eat scolytid eggs and lar- vae, but are not considered to be obligate predators (Stehr 1991). VOLUME 105, NUMBER | 115 100 90 fe Died as an early instar | |CDied after tunneling 80 ‘'WEmerged as an adult | 70 2 2 o 60 2a Oo O50 & a. S 40 Oo o 30 20 10 0 1 2 3 4 5 Cavity size class Fig. 1. Locust borer development with regard to cavity size class Odiniid larvae are thought to be scav- engers of insect frass and other decaying organic matter. They may attack other lar- vae (Stehr 1991). A Cryptarcha larva ap- peared to retreat upon encountering an odi- niid larva during one cavity dissection. Al- though odiniid larvae might be aggressive towards other cavity taxa, they seemed too small in size and low in number to have an overall negative effect on locust borers. Xylomid larvae are thought to be scav- engers on decaying organic matter or pred- ators of larvae (Stehr 1991). Al- though xylomid larvae may be predaceous, they did not appear to have an overall neg- ative effect on borers. Borer adults presum- insect ably emerged from cavities with xylomid larvae twice as often as they failed to emerge from cavities with them. Mesostigmatids, commonly found in the borer cavities are mostly free-living preda- tors. Many are external or internal parasites of invertebrates, reptiles, birds, and mam- mals (Krantz 1978). Because the mesostig- matids were most often found alone in the cavities, they did not appear to pose a threat to borer larvae. Nematodes (Rhabditida and possibly oth- er orders) were a common cavity taxon, but their affect on borer known. Adult borers presumably emerged larvae remains un- from cavities with hundreds of nematodes. Nematodes were found most often in moist within surrounding and were occasionally found on dead mites and nitidulid larvae. One of the most common cavities tissues, nematode inquilines collected from the sub- cortical cavities, Myctolaimellus robiniae n. sp. (Diplogasterida: Cylindrocorporidae), has been described (Harman et al. 2000), 116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON | @Strip-mine | Ol Roadside | BOld field-pasture Number of cavities Cavity size class Fig. 2. Number of cavities per site type with regard to cavity size class. and others are under investigation. The eco- appear to be predaceous. Conspecific staph- logical role of these nematodes was not de- — ylinid larvae and adults usually consume termined, but records stated that most Cy- the same foods (Borror et al. 1989). Nine- lindrocorporids are saprobes, often in sym- teen specimens of larvae and adults were biotic relationships with insects. collected from cavities and traps. Twelve Most species of rove beetles (Staphylin- checkered beetle larvae (Cleridae) were idae), also collected in the present study, collected primarily from borer traps. Most Table 4. Mean + standard error comparison of locust borer cavity measurements by site type. Means not sharing the same lower case letter within columns are significantly different (P < 0.05, Fisher's least significant difference test). Cavity Length Width Depth Volume Bark Thickness Site Type (mm) (mm) (mm) (mm*) (mm) Strip-mine 16.2 + 0.8a 15.4 + 0.8a 4.0 + 0.2a 1057.4 + 101.1a 11.4 + 04a Roadside 11.2 + 0.6b 11.4 + 0.8b 3.2, 4 102b 610.0 = 91.8b 10.9 + O.4a Old field-pasture 13:7, = .O'6c 12:5. +.0!7b 2-4 + 0.2¢ 547.0 + 68.6b 9.7 + 0.3b ANOVA ya" 13.66 7.02 15.43 10.38 5.92 df 2, 284 2, 284 2, 270 22:70 2, 271 P 0.0001 0.001 0.000 1 0.0001 0.003 VOLUME 105, NUMBER | checkered beetles are predaceous as larvae and adults. They are common on or within trees, where they prey on larvae of wood- boring insects, especially bark beetles (Bor- ror et al. 1989). Both rove beetles and checkered beetles were not thought to pose any threat to locust borers, as they occurred infrequently, and were not present in the three cavity samples containing live borer larvae. Earwigs (Dermaptera), most often found grouped together, feed primarily on dead or decaying plant matter. Some feed on living plants, and some may be predaceous (Bor- ror et al. 1989), Like rove and checkered beetles, earwigs were thought to occur too infrequently to pose any threat to borers. Surprisingly, Gordian worms (Nemato- morpha) were found in eight traps. Nema- tomorphs, as juveniles, live within the body cavity of a host, usually an insect, and a host dies after its worm emerges (Pearse et al. 1987). Although Gordian worms may have been killed some borers, adult borers emerged from four traps that contained these worms. Ants (Formicidae) were occasionally found in borer cavities. They generally ap- peared to be too small to pose any threat borer larvae and were probably feeding on tree sap and cavity taxa other than borer larvae. Most ant species found in cavities were considered subordinate according to Fellers’ (1987) ranking of ants by domi- nance. Subordinate ants tend to be less ag- gressive and avoid other ant species. Coleoptera and Lepidoptera appear to support the greater ranges of parasitoids (Mills 1994). Two parasitoid families (Tachinidae and Phoridae) that utilize co- leopteran larvae were collected in borer traps, and one family (Encyrtidae) was col- lected from a borer cavity. Some Encyrtidae and Tachinidae utilize coleopteran larvae while some phorid parasitoids utilize cole- opteran pupae (Mills 1994). Each parasitoid family was represented by only one speci- men in this study. Cavity size is related to advancement and Mg survival of borer larvae, although it appears that many other determinants of cavity size are unknown. Strip-mine sites had signifi- cantly larger cavities, indicating more suc- cess among borer larvae, and yielded more mites than roadside and old field-pasture sites. A major implication regarding black locust on strip-mine sites is that stress is greater on-mine than off-mine resulting in increased vulnerability to invasion by the borer (Harman et. al. 1985), and other prob- able organisms. Although the locust borer is a primary invader, its success is still re- lated to the health of its host. The astigmatid mites found in the cavi- ties appeared to be primarily fungivorous. Many Astigmata are saprophagous, fungiv- orous, or graminivorous, while some are parasitic (Krantz 1978). A colony of Astig- mata of the cavities collected from a strip-mined site was maintained for over a year, subsisting on a diet limited to yeast pellets (K. Larson, personal observa- tion 1996). Significantly greater numbers of fungivorous mites on strip-mine sites than from one off-mine may indicate greater fungal infes- tation, due to stress at the site. Hopkins (1906) stated dead areas sur- rounding cavities, or hibernation cells, pen- etrate into the wood and appear to assist larvae in boring through inner bark to the wood. He also stated if the surrounding area is healthy, the larvae may not advance their tunnels into the wood (Hopkins 1906). Fun- gal infestation in less vigorous trees may contribute to the success of borer larvae in creating larger cavities. Wounds made by borer larvae provide an opening for fungi to enter locust trees. Damage by a heart rot fungus Fomes rimosus (Berkeley) Cooke a parasite of black locust, causes heartwood to become lightweight and crumbly (Hof- fard and Anderson 1982). While borer lar- vae over-winter, fungal activity around their hibernation cells may weaken surrounding tissue to such an extent that larvae may be able to consume more tissue when they re- sume activity in spring, thereby increasing 118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cavity size and improving chances of sur- vival. Selection of varieties of black locust re- sistant to fungi may reduce damage in- curred by borers requiring the presence of fungi for greater survival. Unfortunately, past references to fungi in relation to locust borer ecology have been largely observa- tional. Although mycological aspects were also beyond the scope of this study, the re- sults indicate a need for further studies of the tree-insect-fungal relationship. ACKNOWLEDGMENTS We thank Gary Hevel (Collembola, Pso- coptera, Membracidae, Staphylinidae, Niti- dulidae, Drosophilidae, Phoridae, Antho- myiidae, Muscidae, Tachinidae, Trogositi- dae, Coccinellidae, and Gelechiidae), and O. S. Flint (Chrysopidae), Smithsonian In- stitution; D. R. Smith (Formicidae and En- cyrtidae), Natalia J. Vandenberg (larval Cleridae, Nitidulidae, and Staphylinidae), Peter Touhey (Proctotrupoidea and Lon- chaeidae), A. L. Norrbom (Odiniidae), N. E. Woodley (Xylomidae), the late Curtis W. Sabrosky (Chloropidae), Thomas J. Henry (Anthocoridae and Miridae), D. R. Miller (Coccidae and Psyllidae), and David A. Nickle (Forficulidae), Systematic Entomol- ogy Laboratory (SEL), Agricultural Re- search Service, USDA; William L, Grogan (Ceratopogonidae), SEL cooperating scien- tist, Salisbury University, MD; and Wayne A. Yoder (Araneae, Acari, and Haplotre- matidae), Frostburg State University, for as- sistance with identifications. Partial funding for the study was provided by the Maryland Agricultural Experiment Station. This manuscript is Science Series Con- tribution No. 3569 of the University of Maryland, Center of Environmental Stud- 1es. LITERATURE CITED Blackwell, W. H. and M. J. Powell. 1981. Observations on late season pollination of Solidago canadensis L. (Asteraceae) in Southwest Ohio. Castanea 2: 136-139. Borror, D. J., C. A. Triplehorn, and N. EF Johnson. 1989. An Introduction to the Study of Insects, 6th ed. Saunders College Publishing, Fort Worth, Tex- as, 875 pp. Carter, J. K. and R. O. Blanchard. 1977. Electrical re- sistance related to phloem width in red maple. Ca- nadian Journal of Forest Research 8: 90-93. Cuno, J. B. 1930. Utilization of black locust. United States Department of Agriculture Circular 131. Echaves, V., D. M. Harman, and A. L. Harman. 1998. Site quality in relation to damage by locust borer Megacyllene robiniae Forster in black locust. Journal of Entomological Science 33(1): 106-112. Eliason, E. A. and D. A. Potter. 2000. Biology of Cal- lirhytis cornigera (Hymenoptera: Cynipidae) and the arthropod community inhabiting its galls. En- vironmental Entomology 29(3): 551-559. Fellers, J. H. 1987. Interference and exploitation in a guild of woodland ants. Ecology 68: 1466-1478. Garman, H. 1916. The locust borer (Cyllene robiniae) and other enemies of the black locust. Bulletin of the Kentucky Agricultural Experimental Station 29: 99-135. Hall, R. C. 1942. Control of the locust borer. United States Department of Agriculture Circular 626, 19 PPp- Hall} R.G., W. iG. ‘Baker, H.C: ‘Secrest;/andsH 7Ae Waters. 1938. Report on investigation of the lo- cust borer problem conducted at the Columbus, Ohio Laboratory of the Bureau of Entomology and Plant Quarantine, 1931-1937 inclusive. Harman, D. M. and A. L. Harman. 1987. Distribution pattern of adult locust borers, (Coleoptera: Cer- ambycidae) on nearby goldenrod, Solidago spp. (Asteracae) at a forest-field edge. Proceedings of the Entomological Society of Washington 89: 706-710. Harman, D. M., M. A. Van Tyne, and W. A. Thomp- son. 1985. Comparison of locust borer Megacyl- lene robiniae Forster (Coleoptera: Cerambycidae) attacks on coal strip-mined lands and lands not mined. Annuals of the Entomological Society of America 78: 50-53. Harman, A. L., J. L. Winter, and D. M. Harman. 2000. sp. (Diplogasterida: Cylindrocorporidae) from larval cavities of the lo- Myctolaimellus robiniae n. cust borer, Megacyllene robiniae Forster. Journal of Nematology 32(4): 389-395. Hoffard, W. H. and R. L. Anderson. 1982. A guide to common insects, diseases, and other problems of black locust. United States Department of Agri- culture, Forest Service, Southeastern Area, Forest Pest Management, Forestry Report SA-FR 19. Hopkins, A. D. 1906. Some insects injurious to forests. The locust borer. United States Bureau of Ento- mology Bulletin 58: 1-16. . 1907. The locust borer and methods for its VOLUME 105, NUMBER | control. United States Bureau of Entomology Cir- cular 83, 8 pp. Kellog, L. F 1936. Site index curves for plantations of black locust, central states region. Central States Experimental Station, no. 36. Knietel, J. M. and T. E. Miller. 2002. Resource and top-predator regulation in the pitcher plant (Sar- racenia purpurea) inquiline community. Ecology 83(3): 680-688. Krantz, G. W. 1978. A Manual of Acarology, 2nd ed. Oregon State University Book Stores, Inc., Cor- vallis. 509 pp. Lincoln, R. J.. G. A. Boxshall, and P. FE Clark. 1986. A Dictionary of Ecology, Evolution, and System- atics. Cambridge University Press. 298 pp. Madden, R. C. 1996. Cerambycid beetle on goldenrod. Website: http://www.marymt.edu/~madden/ceram. html. McCann, J. M. 1992. Studies in locust borer (Mega- cyllene robiniae Forster) ecology: I. Intrastand variation in locust borer attack rate. Il. Rate and characteristics of woodpecker predation on locust borers, M.S. Thesis, Frostburg State University, Frostburg, Maryland. Mills, N. J. 1994. Parasitoid guilds: defining the struc- ture of the parasitoid communities of endoptery- 119 gote insect hosts. Environmental Entomology 23: 1066-1083. Pearse, V., J. Pearse, M. Buchsbaum, and R. Buchs- baum. 1987. Living Invertebrates. The Boxwood Press, Pacific Grove, California. 848 pp. SAS Institute. 1988. SAS/STAT user’s guide, release 6.03 ed., SAS Institute Inc. Cary, North Carolina, 1,028 pp. Shibata, E. 2001. Synchronization of shoot elongation in the bamboo Phyllostachys heterocycla (Mono- cotyledoneae: Gramineae) and emergency of the gall maker Aiolomormpus rhopaloides (Hymenop- tera: Eurytomidae) and its inquiline Diomorus ail- omorphi (Hymenoptera: Torymidae). Environ- mental Entomology 30(6): 1098-1102. Shigo, A. L. 1982. Tree health. Journal of Arboricul- ture 8: 311-316. Stehr, F W. 1991. Immature Insects, Vol. 2. Kendall/ Hunt Publishing Company, Dubuque, Iowa. Van Tyne, M. A. 1983. A comparison of the incidence and severity of the locust borer, Megacyllene ro- biniae, on reclaimed strip-mined areas vs. non- strip-mined areas in western Maryland, M.S. The- sis, Frostburg State University, Frostburg, Mary- land. Wenger, K. F [ed]. 1984. Forestry Handbook, 2nd ed. John Wiley and Sons, New York, 1,335 pp. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 120-126 SURVIVAL OF LARVAE AND NYMPHS OF IXODES SCAPULARIS SAY (ACARI: IXODIDAE) IN FOUR HABITATS IN MARYLAND J. EF CARROLL U.S. Department of Agriculture, Agricultural Research Service, Parasite Biology, Ep- idemiology and Systematics Laboratory, Beltsville, MD 20705, U.S.A. (e-mail: jcarrol] @ anri.barc.usda.gov) Abstract.—Host-seeking and fed larvae and nymphs of the blacklegged tick, /xodes scapularis Say, were placed in mesh packets and in vials in the leaf litter on the floor of mixed deciduous forest, Virginia pine-southern red oak forest, and white pine plantations with and without Nepal microstegium, Microstegium vimineum (Trinius) A. Camus. An introduced shade-tolerant grass, Nepal microstegium, is expanding its range northeastward into areas densely populated with /. scapularis. As determined by flag sampling, the density of host-seeking nymphs at the Virginia pine sites was much lower than in the other habitats. None of the four habitats appeared to be consistently more favorable or unfavorable for the survival of confined fed and unfed /. scapularis larvae and nymphs. More unfed nymphs survived in vials than in packets in Virginia pine and white pine with Nepal grass sites. Fed larvae and nymphs tended to survive the summer better than unfed ticks. Key Words: In the United States, most cases of Lyme disease occur in an area from southern New England through the mid-Atlantic states (Spielman et al. 1985) with Maryland as the southernmost state with a significant Lyme disease problem. The blacklegged tick, /y- odes scapularis Say, the principal vector of the agent causing Lyme disease, is also in- volved in the transmission of the agents of babesiosis and human granulocytic ehrlich- losis (Spielman et al. 1985, Dumler and Bakken 1995). Off-host survival of fed and untfed ticks is affected by a variety of biotic and abiotic factors, such as natural enemies (e.g., predators, pathogens) and microme- teorological conditions (e.g., relative hu- midity) (Daniel and Dusbabek 1994). These factors seem to be associated with micro- habit, but differences in survival of flat (un- fed) /. scapularis nymphs may vary from blacklegged tick, immature stages, Nepal microstegium one region to another independent of the type of microhabitat (Bertrand and Wilson 1997). Extreme temperatures and low rela- tive humidities are harmful to /. scapularis (Stafford 1994, Vandyk et al. 1996). Gins- berg and Zhioua (1996) found that /. sca- pularis nymphs had a greater survival rate in deciduous forest compared to pine forest on Long Island, New York. Lord (1993) re- ported high mortality of unfed /. scapularis nymphs in New York. In Maryland, 64— 70% of fed female /. scapularis placed in leaf litter in a deciduous forest survived to oviposit at least some eggs (Carroll 1996), but little is known about the survival of free-living larvae and nymphs. In Maryland, mixed deciduous forests are a common natural habitat, varying in composition of dominant plant species ac- cording to soil type, drainage and other fac- VOLUME 105, NUMBER | tors. Blacklegged ticks are typically found in these deciduous woodlands in Maryland (Schmidtmann et al. 1994, Carroll and Kra- mer 2001). Often, on well-drained soils practically contiguous with the deciduous forests there are stands of Virginia pine, Pi- nus virginiana Miller, and southern red oak, Quercus falcata Michaux. In central Mary- land, white pine, P. strobus L., does not occur naturally (Ehas 1980) but is some- times planted as ornamental or in mono- cultures as a form of reforestation. Black- legged ticks also occur in the pine-domi- nated habitats. White-tailed deer, Odoco- ileus virginianus (Zimmermann), and other important hosts of /. scapularis readily move among all three habitats. Nepal mi- crostegium, Microstegium vimineum (Tri- nius) A. Camus (Poaceae), is a shade-tol- erant, non-native species of annual (some- times perennial) grass. Since being intro- duced into Tennessee over 80 years ago, M. vimineum has attained the status of an in- vasive weed in the U. S., while expanding its range northeastward into Maryland, New York, and New England (Hunt and Zaremba 1992, Redman 1995, Ehrenfeld 1999). Nepal microstegium has experienced explosive distributional growth in Mary- land since the 1980s (Redman 1995). In ar- eas around Loch Raven Reservoir, Balti- more County, where there are extensive plantings of white pine, M. vimineum is the dominant understory plant species, cover- ing considerable surface area and attaining heights of ~0.3 m. /xodes scapularis is abundant in these white pine and adjacent deciduous woodlands (Carroll, unpublished data). Nepal grass forms sprawling colonies that might provide shaded, humid refugia for larvae and nymphs of /. scapularis and enhance their survival during late spring and summer. The grass dies and collapses in the fall. The purpose of this study was to obtain preliminary information on the survival of fed and unfed /. scapularis nymphs in the deciduous forest, Virginia pine-oak forest, white pine plantations with 121 Nepal grass, and white pine plantations without Nepal grass. MATERIALS AND METHODS The deciduous forest study sites were lo- cated at Loch Raven Reservoir, Baltimore County, Maryland, as were the white pine plantings with and without Nepal grass. The Virginia pine-southern red oak forests were located in the U. S. Fish and Wildlife Ser- vice’s Patuxent North Tract in adjoining Anne Arundel County. The deciduous for- est was dominated by tulip tree, Lirioden- dron tulipifera L., red maple, Acer rubrum L., hickory, Carya sp., and black cherry, Prunus serotina Ehrhart. Understory vege- tation was generally sparse in the deciduous woods, with some ferns and Nepal grass present. Leaf litter in these deciduous for- ests was notably shallow, and in some plac- es virtually gone by mid—late summer, per- haps due to the abundance of earthworms that were observed. In contrast, leaf litter in the Virginia pine-southern red oak forests remained ~10 cm deep. Few other species of trees were present in these woods, and blueberries, Vaccinium sp., and greenbrier, Smilax sp., were common in the understory. The white pine plantations were virtually monocultures, with ferns and Nepal grass being the principal understory species. For each of the four types of habitat, three sites (=0.8 km apart) were selected (Fig. 1). Each site was sampled for the pres- ence of /. scapularis by flagging with a 0.5 by 0.5 m flannel cloth (crib cloth containing a rubber laminate) 5 times for 30 sec while walking slowly. At each of the three sites, confined ticks were placed in the leaf litter (where loose litter adjoined compacted litter or soil) at each of three locations, with the exception of unfed larvae, which were placed at one randomly selected location at each site in 1999, At each site each group of ticks was placed =10 m from the nearest group of confined ticks. The location of each group was marked with a flag. Unfed and fed larvae and nymphs of /. scapularis were placed at these locations in 1999 and 122 Habitats — White pine/ White. pine Nepal grass PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Deciduous Virginia pine Sites [a B Cc Vials/) W233) 4-203) 12:3 packets Fig. 1. i i2ese dee Saati 2e3 | A B Cc 1.203, 423) 123) 2.3 Seas: For each of 4 types of habitat, unfed and fed larvae and nymphs confined in nylon packets or in vials were placed in leaf litter at 3 sites (designated here as A, B and C). At each site the packets and vials were placed at 3 marked locations (designated here as 1, 2000 when free-living ticks were in the same stage of development. Unted larvae and nymphs were collected from the field, placed in plastic snap-cap vials and main- tained at 24—25° C, R. H. ~97%, and nat- ural photoperiod. Fed larvae and nymphs were allowed to engorge on white rats in accordance with an approved USDA, ARS, Beltsville Agricultural Research Center, [A- CUC protocol. Within 4 d after dropping from a rat, the fed larvae and nymphs were placed in nylon packets to be distributed at the study sites. The packets were of 82 by 82 mesh per cm? folded once to form a rath- er flat ~2 by 5 cm rectangle sealed on two sides by adhesive applied with a glue gun. Once the ticks were placed inside the pack- et, the fourth side was folded over and closed as securely as possible with a bull- dog clip. In 1999, 10 unfed larvae, 5 fed larvae, 6 unfed nymphs, and 3 fed nymphs were placed in each packet. The following year 8 unfed larvae, 6 fed larvae, 3 unfed nymphs, and 3 fed nymphs were placed in each packet. In 1999, unfed nymphs (6 per vial) were placed in plastic snap-cap vials (3 dram) with a 0.8 cm diameter hole in the cap and nylon cloth covering the mouth of the vial. Except in 1999, when one packet containing unfed larvae was placed at one 2 and 3) =10 m apart. of the three locations (chosen randomly) at each of the three sites for each habitat, one packet or vial containing ticks was distrib- uted to each location at each site. Nymphs were placed out in late spring to early sum- mer, and larvae in August. Unfed ticks were removed from the field and checked for mortality near the end of the natural activity periods for larvae and nymphs (late August to September). Fed ticks were checked after the time that individuals of the same stage in nature should have molted into the next life stage. Standard errors were calculated for surviving ticks by habitat. Mean num- bers of ticks in surviving vials and packets were compared using Student’s t-test. RESULTS At least one nymph or adult of /. sca- pularis was found by flagging at all but one of the study sites where ticks were placed (Table 1). However, the Virginia pine-oak habitat contained few /. scapularis. The summer of 1999 and the fall of 2000 were unusually dry, whereas the summer of 2000 was wet. In 1999, unfed larvae survived poorly in all habitats (=43%), but especial- ly so in the deciduous forest (7%), which had significantly fewer (P = 0.004) survi- vors than the Virginia pine-southern red oak VOLUME 105, NUMBER 1 Table |. ~) Ww Numbers of /. scapularis nymphs and adults captured by flagging 5 times for 30 sec while walking slowly at each study site just before the first packets of ticks were placed in the leaf litter. Virginia Pine Deciduous White Pine Open White Pine Grass Nymphs Adults Nymphs Adults Nymphs Adults Nymphs Adults Site | 1 0) 9 0 Ci 0 7 | Site 2 0 0 7 10) 23 0 10 0 Site 3 0 | 6 (0) 10 (0) | 0 sites (Table 2). Survival of unfed larvae was even worse in 2000, with <10% sur- viving. However, despite overall low levels of survival of unfed larvae in 2000, larval survival was somewhat greater (P = 0.0496) in the white pine with Nepal grass sites than the deciduous forest and white pine without Nepal grass sites. In 1999 few- er unfed nymphs in packets survived at the Virginia pine sites than at the deciduous forest sites (P = 0.014) (Table 3). At the Virginia pine and white pine with Nepal sites, significantly more unfed nymphs survived in vials than in packets (P < 0.05). Very few unfed nymphs (=2%) survived in 2000. In general, substantial proportions of fed 7. scapularis larvae and nymphs molted to the next stage and no dif- ferences in survival of fed ticks among the habitats were detected. In 1999, >72% of fed larvae in all four habitats survived to become nymphs, but in 2000 only about half (36-59%) survived. Most fed nymphs grass Table 2. in all habitats survived to become adults, with >62% surviving in 1999 and >83% surviving in 2000. DISCUSSION Based on flag sampling of nymphs and adults, the Virginia pine-southern red oak sites in this study appeared to support very few /. scapularis. Because Virginia pine habitats are often contiguous with decidu- ous forests in Maryland, and share deer and other hosts of /. scapularis, further com- parative sampling of /. scapularis in these these habitats is warranted. Ginsberg and Zhioua (1996) also found lower densities of I, scapularis in pine (mostly pitch pine, P. rigida Miller) woods than deciduous thick- ets. In New Jersey Schulze et al. (1998) found /. scapularis nymphs much more nu- merous in pitch pine habitats than in three other habitats not dominated by pines at one study area. At a second study area where white pine was the dominant species, num- Numbers of unfed and fed larvae of /. scapularis surviving in each of the 4 habitats. Means per location on following line; percentage of ticks surviving in each habitat in parentheses. Virginia Pine Deciduous White Pine Open White Pine Grass Unfed Larvae 1999 13/308 (43%) 2/30 (7%) 4.3 + 0.9 OF O87 2000 3/90 (3%) 2/90 (2%) 03) =.02 O2 = '0:2 Fed Larvae 1999 15/20 (75%) 33/40 (83%) 3.8 = '0!5 47° ='012 2000 32/54 (59%) 23/42 (55%) 36225057. 3.324) 007 9/30 (30%) 10/30 (33%) 3.0' + 016 S25) 129 2/90 (2%) 8/90 (9%) 0.2 = 0.2 0.9 = 0.3 18/25 (72%) 22/30 (77%) 3.6015 3.8 + 0:5 15/42 (36%) 23/48 (48%) Del O'S: 2.9 + 0:4 *Denominators (total number of ticks) vary because not all packets containing ticks were recovered. Some recovered packets were not intact, and all ticks had escaped. 124 Table 3. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Numbers of unfed and fed nymphs of /. scapularis surviving in each of the 4 habitats. Means per location on following line; percentage of ticks surviving in each habitat in parentheses. Virginia Pine Deciduous White Pine Open White Pine Grass Unfed Nymphs In Packets 1999 11/54" (20%) 26/48 (54% 12, 10:5 333° )0!5 2000 1/54 (2%) 0/48 (0%) Om 2071 0 In Vials 1999 29/54 (54%) 23/48 (48% 3.2 + 0.6 2.9 = 0.6 Fed Nymphs 1999 9/9 (100%) 9/12 (75%) 3.0 +0 2 Set 1O'5 2000 21/24 (88%) 20/24 (83% 2.6 = 0:2 DES 20:2 19/54 (35%) 9/42 (21%) 221 =10:6 V3) = 10%7, 1/54 (2%) O/48 (0%) Os 081 0 20/48 (42%) 25/42 (60%) 25) =:06 Boy a (917/ 13/21 (62%) 10/12 (83%) Ae) ee (05) 25 )==103 8/9 (89%) 16/18 (89%) 21 = 03 De = OD * Denominators (total numbers of ticks) vary because not all packets or vials containing ticks were recovered. Some recovered packets and vials were not intact, and ticks had escaped. bers of /. scapularis nymphs were not high- er than in three other habitats not dominated by pines (Schulze et al. 1998). In the present study, Virginia pine sites did not stand out as inhospitable to the sur- vival of J. scapularis larvae and nymphs, rather survival of fed 1. scapularis larvae and nymphs did not appear to vary consis- tently among the four types of habitat. As the summers progressed, the leaf litter in the deciduous forests steadily diminished to the extent that in a few instances packets were found exposed on bare soil, at the ex- act locations where they had been covered with fallen leaves when originally placed. This unusual (in the author’s experience in Maryland deciduous forests) phenomenon may account for the comparatively poor survival of flat nymphs in packets in decid- uous woods in the drought year 1999. Staf- ford (1994) found that nymphs of /. sca- pularis require elevated relative humidities for extended survival. Lord (1993) reported high mortality (90% in 45 d) of unfed /. scapularis nymphs confined in mesh pack- ets (10 by 20 cm) partly buried in leaf litter and soil in deciduous woods in southern New York. With totally exposed packets subject to more desiccating conditions, it is not surprising that many ticks died. In the following year, which experienced regular summer rains, virtually all unfed nymphs in all habitats died. Only in 2000, when sur- vival of unfed larvae was extremely low at all sites, was there any indication that Nepal grass colonies might be more favorable for survival of either life stage of /. scapluris than the open white pine woods. In general, flat nymphs and larvae survived better in 1999 than in 2000 when rainfall was plen- tiful. Flooding or prolonged submersion of the confined ticks may have caused mortal- ity directly or created conditions favoring fungal or perhaps even bacterial pathogens harmful to the ticks. Using larger packets or cages which extend upward out of the litter would give mobile ticks the opportu- nity to move to more favorable microenvi- ronments when conditions change (Yuval and Spielman 1990, Lord 1993). However, in the case of Lord (1993), unfed nymphs survived as poorly (<10%) as the unfed nymphs did in this study in 2000. In 1999, 20-54% of unfed nymphs in all four habi- tats survived the summer. Host-seeking nymphs of /. scapularis can disperse 25 m, VOLUME 105, NUMBER | thereby avoiding some deleterious situa- tions (Carroll and Schmidtmann 1996). Al- though fed larvae and nymphs lack the mo- bility of their unfed counterparts, much higher percentages of fed than unfed ticks survived in both years. In the late summer of 1999, an unknown person removed all the flags marking the positions of the tick packets from one open white pine site and a nearby white pine site with Nepal grass. Despite written descrip- tions of the locations of the packets, some were not found. A few other packets, per- haps removed by rodents or birds, were never recovered. Some packets, which were recovered, were found to have holes through which all the ticks escaped. These losses may have reduced some of the antic- ipated discriminating power of the study, but enough packets were recovered to de- tect any gross differences and some more subtle differences in tick survival among the habitats. In conclusion, even though the study took place during a unusually dry summer and a wet summer, none of the habitats ap- peared obviously more favorable or unfa- vorable for survival of confined fed and un- fed larvae and nymphs of /. scapularis. Pur- ther comparative sampling of host-seeking I. scapularis in Virginia pine-southern red oak and other Maryland habitats is needed. ACKNOWLEDGMENTS I thank Gene Scarpulla, Reservoir Natu- ral Resources Office, Environmental Ser- vices Division, City of Baltimore, Depart- ment of Public Works, Bureau of Water and Waste Water, Eldersburg, MD, for his co- operation and for encouraging interest in the Nepal microstegium problem. I also ex- press my gratitude to Kenneth Young and Eli Miramontes, USDA, ARS, Parasite Bi- ology, Epidemiology and Systematics Lab- oratory, Beltsville, MD, for their assistance in making the packets used for confining the ticks. LITERATURE CITED Bertrand, M. R. and M. L. Wilson. 1997. Microhabitat- independent regional differences in survival of un- fed Lxvodes scapularis nymphs (Acari: Ixodidae) in Connecticut. Journal of Medical Entomology 34: 167-172. Carroll, J. EF 1996. Survivorship of engorged female Ixodes scapularis and their eggs in a leaf litter microhabitat in Maryland. Entomologia Experien- talis et Applicata 78: 349-351. Carroll, J. F and M. Kramer. 2001. Different activities and footwear influence exposure to host-seeking nymphs of /xodes scapularis and Amblyomma americanum (Acari: Ixodidae). Journal of Medical Entomology 38: 596—600. Carroll, J. F 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. Daniel, M. and F. Dusbabek. 1994. Micrometeoro- logical and microhabitat factors affecting main- tenance and dissemination of tick-borne diseas- es in the environment, pp. 91-138. Jn Sonen- shine, D. E. and T. M. Mather, eds. Ecological Dynamics of Tick-Borne Zoonoses. Oxford, New York. Dumler, J. S. and J. S. Bakken. 1995. Ehrlichial dis- eases of humans: emerging tick-borne infections. Clinical Infectious Diseases 20: 1102-1110. Ehrenfeld, J. G. 1999. A rhizomatous, perennial form of Microstegium vimineum (Trin.) A. Camus in New Jersey. Journal of the Torrey Botanical So- ciety 126: 352-358. Elias, T. S. 1980. The Complete Trees of North Amer- ica: Field Guide and Natural History. Van Nos- trand Reinhold, New York, 948 pp. Ginsberg, H. S. and E. Zhioua. 1996. Nymphal sur- vival and habitat distribution of /xodes scapularis and Amblyomma americanum ticks (Acari: [xod- idae) on Fire Island, New York, USA. Experimen- tal and Applied Acarology 20: 533-544. Hunt, D. M., and R. E. Zaremba. 1992. The north- eastward spread of Microstegium vimineum (Po- aceae) into New York and adjacent states. Rho- dora 94: 167-170. Lord, C. C. 1993. Mortality of unfed nymphal /rodes dammini (Acari: Ixodidae) in field exclosures. En- vironmental Entomology 22: 82-87. Redman, D. E. 1995. Distribution and habitat types or Nepal microstegium [Microstegium vimineum (Trin.) Camus] in Maryland and the District of Columbia. Castanea 60: 270-276. Schmidtmann, E. T., J. EF Carroll, and W. J. E. Potts. 1994. Host-seeking of black-legged tick (Acari: Ixodidae) nymphs and adults at the woods-pasture interface. Journal of Medical 291-296. Schulze, T. L., R. A. Jordan, and R. W. Hung. 1998. Entomology 31: 126 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Comparison of /xodes scapularis (Acari: Ixodi- humidities. Journal of Medical Entomology 31: dae) populations and their habitats in established 310-314. and emerging Lyme disease areas in New Jersey. Vandyk, J. K., D. M. Bartholomew, W. A. Rowley, and Journal of Medical Entomology 35: 64—70. K. B. Platt. 1996. Survival of /xodes scapularis Spielman, A., M. L. Wilson, J. F Levine, and J. Pies- (Acari: Ixodidae) exposed to cold. Journal of man. 1985. Ecology of /xodes dammini-borne hu- Medical Entomology 33: 6-10. man babesiosis and Lyme disease. Annual Review —_- Yuval, B. and A. Spielman. 1990. Duration and regu- of Entomology 30: 439-460. lation of the developmental cycle of Ixodes dam- Stafford HI, K. C. 1994. Survival of immature /xodes mini (Acari: Ixodidae). Journal of Medical Ento- scapularis (Acari: Ixodidae) at different relative mology 27: 196-201. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 127-130 A NEW SPECIES OF PLUMISPINA ALBUQUERQUE (DIPTERA: MUSCIDAE) FROM SOUTHERN BRAZIL NISE DO CARMO COSTACURTA, CLAUDIO JOSE BARROS DE CARVALHO, AND MARCIA SOUTO COURI (NdCC, CJBdC) Departamento de Zoologia, Universidade Federal do Parana, Curitiba, Parana, 81531-980, Brazil (e-mail: cjbcarva@bio.ufpr.br); (MSC) Departamento de En- tomologia, Museu Nacional, Quinta da Boa Vista, 20940-040, Rio de Janeiro, RJ, Brazil (e-mail: mcouri @attglobal.net) Abstract.—A new species of the genus Plumispina Albuquerque (Diptera: Muscidae), P. similis Costacurta and Carvalho, from Guarapuava, Parana, southern Brazil is de- scribed, diagnosed and figured. A diagnosis of the genus P/umispina and a key to its two known species are provided. Key Words: The Neotropical genus Plumispina was proposed by Albuquerque (1954) to include a single species, P. longipilis, collected in the city of Rio de Janeiro, Brazil. The genus was accepted by Pont (1972) as monobasic, but recently Carvalho et al. (1993) consid- ered Plumispina as a junior synonym of Neodexiopsis Malloch. In a key to the world genera of the Coenosiini, Couri and Pont (1999) revalidated Plumispina. In their cladistic analysis (Court and Pont 2000) they indicated that Plumispina belongs to a larger monophyletic group containing 17 genera, including Coenosia Meigen. In this paper, we describe a new species of Plumispina from Guarapuava which also was found in Ponta Grossa and Colombo, southern Brazil, present a diagnosis of the genus, and give a key to its two known spe- cies. MATERIAL AND METHODS This study is based on material deposited in the following institutions: Department of Zoology of “Universidade Federal do Pa- Plumispina, southern Brazil, systematics, taxonomy rand” (DZUP) and “Museu Nacional, Rio de Janeiro” (MNR3J). The terminology and abbreviations used for the external morphology, including ter- minalia, follow McAlpine (1981) and Car- valho (1989). SYSTEMATICS Plumispina Albuquerque 1954 Type-species.—Plumispina longipilis Al- buquerque 1954 (orig. desig.). Diagnosis.—Plumispina 1s a typical Coe- nosiini because of the absence of prealar; the proepimeral lower seta directed down- wards; and katepisternals with 1:1:1 setae forming an equilateral triangle. This genus belongs to a larger, monophyletic Coenosia group sensu Couri and Pont (2000). The main diagnostic characters of the genus are: male dichoptic; one pair of reclinate orbital setae; arista with short plumae; and hind tibia with long ciliation on anterodorsal, dorsal, and posterodorsal surfaces. This last character, together with the setulose sternite 1 and the shape of the hypandrium, gono- pod and paramere (Fig. 3; Albuquerque 128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1954: figs. 5, 7-8) could be considered as autapomorphic characters of the genus. KEY TO SPECIES OF PLUMISPINA (Based on males; females are unknown.) 1. Arista bare on apical third; scutellum with pair of apical setae two-thirds as long as basal one; hind femur and hind tibia with many long hairs on ventral surfaces (Albuquerque 1954: 179, fig. 2); sternite 5 without a median projection on posterior margin (Albuquerque 1954: 179, fig. 9) (Brazil Sy oDioero te oorola te P. longipilis Albuquerque Rio de Janeiro): = acy ichneumonids and tachinids — chalcids. Amydria larvae are the largest colonizers of the upper layers of the fungal dumps ex- amined. Further research is warranted to de- termine the actual nutritional needs of lar- vae, and their level of dietary specialization on fungi. No antagonistic interaction was observed between moth and Atta mexicana. The workers do not dig or tunnel larvae into the dump material to any significant extent. The predaceous ponerine ant, Pa- chycondyla villosa (Fabricius), which was very common at the collection localities, was observed digging a few millimeters into the Atta dumps in what appeared to be attempts to reach Amydria larvae. ACKNOWLEDGMENTS We are indebted to Vichai Malikul and George Venable, Department of Systematic Biology, Entomology Section, Smithsonian Institution, and John Chung for the illustra- tions. John Steiner of the Smithsonian Pho- tographic Services assisted with the photo- graphs. Norman Woodley, Systematic En- tomology Laboratory, U.S. Department of Agriculture, identified the tachinid para- sitoid. Specimens examined are deposited 194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in the following institutions: The Natural History Museum, London, England (BMNH); National Museum of Natural His- tory, Smithsonian Institution, Washington, DC, USA (USNM); and the University of Texas, Austin, TX, USA (UTA). LITERATURE CITED Anderson, C. and FE L. W. Ratnieks. 2000. Task parti- tioning in insect societies: novel situations. Insectes Sociaux 47: 198-199. Beutelspacher, C. R. 1977. Una nueva especie de Ac- rolophus Poey, 1932, de Chamela, Jalisco, México (Lepidoptera: Acrolophidae). Anales Instituto Biologia. Universidad Nacional Autonoma de México. 48, Serie Zoologia (1): 145-150. Bononi, V., M. Autori, and M. Rocha. 1981. Leuco- coprinus gongylophorus (MGéller) Heim, 0 fungo do formigueiro de Atta sexdens rubropilosa Forel. Rickia 9: 93-97. Davis, D. R. 1984. Neopseustidae, Nepticulidae, Opos- tegidae, Tischeriidae, Incurvariidae, Cecidosidae, Adelidae, Prodoxidae, Tineidae, Psychidae, Ar- rhenophanidae, pp. 16-25. /n Heppner, J. B., ed. Atlas of Neotropical Lepidoptera, 1 (Checklist, pt. 1). Dr. W. Junk, The Hague. . 2000. Tineoidea and Gracillarioidea, pp. 469— 482, figs 25.1-25.21. In Bousquets, J. L. et al., eds. Biodiversidad, Taxonomia, y Biogeografia de Artropodos de México: Hacia una Sintesis de su Concocimiento, Vol. 2. Davis, D. R., D. H. Clayton, D. H. Janzen, and A. P. Brooke. 1986. Neotropical Tineidae, II: biological notes and descriptions of two moths phorectic on spiny pocket mice in Costa Rica (Lepidoptera: Ti- neoidea). Proceedings of the Entomological So- ciety of Washington 88: 98—109. Davis, D. R. and G. S. Robinson. 1998. The Tineoidea and Gracillarioidea, pp. 91-117. /n Kristensen, N. P., ed. Lepidoptera, Moths and Butterflies. Hand- buch der Zoologie, Vol. IV, Arthropoda: Insects, Part 35: 91-117. Walter de Gruyter & Co., Berlin, New York. Fisher, P. J., D. J. Stradling, and D. N. Pegler. 1994. Leucoagaricus basidiomata from a live nest of the leaf-cutting ant Atta cephalotes. Mycological Re- search 98: 884-888. Hart, A. G. and FE L. W. Ratnieks. 2001. Task partitioning, division of labour and nest compartmentalisation col- lectively isolate hazardous waste in the leafcutting ant Atta cephalotes. Behavioral Ecology and Socio- biology online, __http://link.springer-ny.com/link/ service/journals/00265/contents/00/003 1 2/. Mueller, U. G., S. A. Rehner, and T. D. Schultz. 1998. The evolution of agriculture in ants. Science 281: 2034-2038. Pagnocca, EF, M. Bacci, Jr., M. H. Fungaro, O. C. Bueno, M. J. Hebling, A. Sant’anna, and M. Ca- pelari. 2001. RAPD analysis of the sexual state and sterile mycelium of the fungus cultivated by the leaf-cutting ant Acromyrmex hispidus fallax. Mycological Research 105: 173-176. Rogers, J. D., Y-M. Ju, and F San Martin-Gonzalez. 1995. Discoxylaria myrmecophila and its Hypo- creodendron anamorph. Mycologia 87: 41—45. Waller, D. and J. C. Moser. 1990. Invertebrate enemies and nest associates of the leaf-cutting ant Atta tex- ana (Buckley) (Formicidae, Attini), pp. 225-273. In Vander Meer, R. K., K. Jaffe, and A. Cedeno, eds. Applied Myrmecology: A World Perspective. Westview Press, Boulder, Colorado. Walsingham, Lord (Thomas De Grey). 1913-14. 42 (Lepidoptera-Heterocera, 4), pp. 225-392. In Godman, F. D. and O. Salvin, Biologia Centrali- Americana. Walter, C. 1938. The Texas leaf-cutting ant and its con- trol. United States Department of Agriculture Cir- cular 494, 18 pp. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 195-202 POLLEN PROVISION RECORDS FOR THREE SOLITARY BEE SPECIES OF MEGACHILE LATREILLE AND HERIADES SPINOLA (HYMENOPTERA: MEGACHILIDAE) IN SOUTHWESTERN MONTANA PETER D. JENSEN, KEVIN M. O’ NEILL, AND MATTHEW LAVIN (PDJ, KMO) Department of Entomology, Montana State University, Bozeman, MT 59717, U.S.A.; PDJ present address: Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (KMO e-mail: koneill@montana.edu); (ML) Department of Plant Sciences and Plant Pathology, Montana State University, Bozeman, MT 59717, U.S.A. Abstract.—We identified the pollen included in nest provisions by three species of solitary bees at four sites in the vicinity of Bozeman, Montana. Megachile relativa Cresson and Heriades carinata Cresson were studied in trap nests in natural populations, whereas Megachile rotundata (FE) were from a managed, introduced population adjacent to an alfalfa field being used for seed production. Over 90% of 186 cells examined in the three species contained more than a single type of pollen (and up to seven different types). The most intensively studied species, M. relativa, provisioned with pollen from eight families of dicot plants (particularly Asteraceae and Fabaceae), as well as two unidentified mono- cots that were also common in provisions of H. carinata and M. rotundata. Results include new pollen records for all three species. Key Words: Megachile, Heriades, trap nests, pollen provisions, alfalfa leafcutting bee data (F.). However, after identifying pollen from nest cells, Strickler et al. (1996) iden- tified two plant families used by M. relati- va, and Stubbs et al. (1994) found four fam- ilies used by M. rotundata. Although these discrepancies could be due to differences in pollen availability among sites, they may also reflect differences between nectar and pollen preferences. Thus, in order to deter- mine the types of pollen collected by bees during actual pollen-collecting trips, it may be necessary to examine either the pollen present in nest provisions or that found on the scopae of foraging bees (or on bees in museum collections). Here, we report the pollen identified from nest cells of three Pollen collected by adult female bees provides the major source of protein and other nutrients for their larvae, and the types of pollen collected can correlate with growth and survival during development (Guirguis and Brindley 1974, Schmidt et al. 1987, Horne 1995, Michener 2000). The types of pollen collected also affect the ef- ficiency of bees as pollinators of cross-pol- linated plants, a subject of particular inter- est to those managing bees for fruit or seed production (Free 1993). Knowledge of the types of pollen collected cannot always be determined from flower visitation records alone because bees may visit flowers solely to obtain nectar. For example, Hurd (1979) cited 16 families of plants whose flowers are visited by Megachile relativa Cresson and 8 families visited by Megachile rotun- megachilid bees, M. relativa and M. rotun- data, and Heriades carinata Cresson, at four sites in the vicinity of Bozeman, Mon- tana. 196 MATERIALS AND METHODS We identified pollen that we removed from the nest cells provisioned by bees dur- ing the summers of 1999 and 2000. Me- gachile rotundata nested within 0.5 * 9.5 cm deep tunnels in commercial polystyrene ““bee boards’? manufactured by Beaver Plastics (Edmonton, Alberta) for use in commercial seed alfalfa production (Rich- ards 1984). The shelter containing the boards was located between two plots of alfalfa (Medicago sativa (L.)) being grown for seed on the Montana State University Post Farm, 3 km west of Bozeman, Gallatin County, Montana. Bees nesting in these boards were purchased as overwintering prepupae from Mennie Bee Farms Inc. (Parkside, Saskatchewan). All M. rotundata cells were provisioned by bees during July and August 2000 at a time when alfalfa was in bloom. The M. relativa and H. carinata were from native populations that nested in two types of trap nests. The first type con- sisted of pine boards into which we drilled 15 cm long holes and inserted paper straws with internal diameters of 3.2, 3.7, 4.6, 5.9, 7.5, 8.0, and 9.0 mm. Megachile relativa nested in 4.6—9.0 mm tubes, and H. cari- nata in 3.7 mm tubes. The second trap nest type, used only by M. relativa, consisted of pine boards with 15 cm long grooves (6.3 and 9.5 mm diameter) routed in the sides, then fitted with removable plexiglass sheets (3 mm thick) to provide a transparent surface for viewing nest con- tents and removing pollen. Nest boards of both types were mounted on fence posts (at heights of 1.5—2.0 m) adjacent to trees and with the nest holes facing southeast. We placed the trap nests at three sites: 1) Bozeman (WB), located on the western end of Bozeman on the Montana State University Horticultural Farm (nests which were western placed within an abandoned ornamental tree farm surrounded by agricultural test plots and weedy fields); 2) southeastern Bozeman (SEB), located in a residential area 3.0 km from WB (nests placed within area that PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON contained ornamental flowering plants and which was 150 m from a weedy industrial storage yard and 300 m from a wooded stream); and 3) Rocky Creek Farm (RCP), just east of Bozeman and 5.6 km east of WB (nests placed along the weedy border of a cultivated field and shaded by lilac, Syringa vulgaris (L.)) which was not in bloom while the bees nested. We used two methods to obtain pollen samples from 186 nest cells, including 145 from 44 M. relativa nests (from WB and SEB), 26 from 19 M. rotundata nests (all from the Post Farm), and 15 from 11 H. carinata nests (5 from WB, 10 from RCF). The first method was to insert the wooden end of a cotton swab stick into the nest and twist it in the provision of the outermost cell while the female was away from the nest. The second method was to open nests in the lab, taking pollen either from uneaten provisions, or from frass left by the devel- oping larvae (Strickler et al. 1996). From mid-May through August 1999, we also collected flowers within 200 m of the nests at approximately three-week intervals. We used this pollen to create a reference col- lection following the methods described by Moore et al. (1991) and Sawyer (1988) with slight adaptations described below. We placed pollen extracted from each cell or plant into an Eppendorf tube with 2 ml of distilled water and one drop of saf- ranin. After 24 h, we centrifuged the sam- ples at 3,000 rpm for 5 min, poured the dye off, and resuspended the pellet in water for a second rinse. After a second centrifuga- tion, we poured off the supernatant and re- suspended the pellet in two drops of water. We then placed the sample on a slide where it was allowed to dry before mounting it in Euparal and sealing the slide with clear nail polish. To identify pollen, we first examined the entire slide under a Nikon phase con- trast light microscope (40). We then ex- amined each type of pollen at high power (100) for identification. By using pollen identification keys (Kapp 1969) and com- paring pollen from nests with pollen in ref- VOLUME 105, NUMBER 1 erence samples, we identified most dicot pollen grains to family and many to genus. We made no exact counts of each type of pollen in samples, but we did record gen- eral estimates of the proportions of different pollen types, which were sometimes un- evenly distributed on slides due to clump- ing. However, we roughly estimated the rel- ative frequencies of different pollen types on each slide, as 1%, 5%, or greater values to the nearest 10%. Rare pollen types rep- resented by only several grains on a slide containing thousands of pollen grains were excluded from counts to reduce the possi- bility of recording pollen incidentally pick- ed up by females on flowers or other sourc- ES: We did not compare the frequency dis- tributions of pollen types provisioned by different bee species because samples came from different sites. However, we did com- pare pollen types in different types of M. relativa samples: 1) WB vs. SEB samples and 2) samples from uneaten provisions vs. those from frass. We first determined if there was a significant (Pearson’s) correla- tion between sample types in the number of cells containing each type of pollen (a sig- nificant correlation indicating similarity of the two samples). Where the correlation was not significant, we used 2 * 2 chi- square contingency table analyses (each with d.f. = 1) to test the null hypothesis that the proportions of cells with and with- out a particular type of pollen (e.g., thistle) were the same in the two sets of samples (a significant difference indicating that a par- ticular type of pollen was over- or under- represented in a set of samples). RESULTS AND DISCUSSION Overall, the three species provisioned with pollen from nine families of dicot plants and two types of monocots (Fig. 1). Using reference samples, we distinguished at least three types of Asteraceae: 1) Cir- sium spp. (thistle); 2) Taraxacum spp. (dan- delion); and 3) unknown Asteraceae. Sim- ilarly, Fabaceae could be divided into 1) 197 Lotus sp. (probably birdsfoot trefoil, Lotus corniculatus L.); 2) Medicago sp. (all which was probably alfalfa, Medicago sa- tiva L.); and 3) unknown Fabaceae. Dicot pollen grains that could not be identified were grouped in an “unknown” category. We found two types of monocot pollen (based on pollen grain size), hereafter re- ferred to as the “‘small” and “‘large’” mono- cots. Both the small and large monocot pol- len grains were of a general type (i.e., pro- late and with a single sulcus), indicating that they were clearly neither grass (Po- aceae) nor cattail (Typhaceae) pollen (Kapp 1969). Pollen provisioned by Megachile relati- va. Fifteen of 16 types of pollen distin- guished occurred in M. relativa nests (Fig. 1). The 145 cells sampled contained a mean (+ SE) of 3.1 + 0.1 types of pollen (range 1-7), but there was considerable variation in the number of pollen types per cell. At one extreme, there were nine cells in which we were able to find just a single pollen type among thousands of grains present in each sample. In a few cases, entire nests contained relatively few pollen types. One nest, for example, averaged just 1.3 + 0.2 pollen types per cell (range 1—2) and two of its six cells each contained a single type (one with Taraxacum, the other with the large monocot). The other extreme was one particularly diverse nest with eight cells that averaged 4.8 + 0.5 pollen types per cell (range 3—7) and contained a total of 8 dif- ferent pollen types. Note that we cannot be sure that all cells in this nest were provi- sioned by the same female, because nest su- persedure is common in trap nesters (Krom- bein 1967). The M. relativa pollen samples came ei- ther from frass (N = 116) or uneaten pro- visions (N = 29). Thus, we were concerned that using different types of samples might bias results if maceration or digestion of some pollen types reduced their detection in frass. However, similarity in the preva- lence of different pollen types in samples from larval frass and uneaten provisions of 198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON M. relativa M. rotundata H. carinata Asteraceae (Cirsium) ee a iz Bis fata 1 unknown dicots Ba 1 ee | Asteraceae (unknown) ae 2 7 monocot (small) aa 3 cee | = Fabaceae (Lotus) ke 1 eS Asteraceae (Taraxacum) 1 = | Fabaceae (Medicago) a: tae | monocot (large) 1 RSIS 3 ee Caprifoliaceae | 1 3 Brassicaceae 5 | 4 | Fabaceae (unknown) ' 4 4 Rosaceae 1 5! Fagaceae | | 4 Oleaceae | | ] Hydrangeaceae | | Tiliaceae 4 | : | 0.0 02 ue 6 08 0.0 02 04 06 08 10 0.0 02 04 06 08 AG Proportion of cells containing each pollen type Fig. |. Pollen records for three species of Megachilidae. Pollen types are ranked from top to bottom based on their occurrence in Megachile relativa records. Numbers by bars indicate the number of cells that in which the pollen type made up 100% of the provision. M. relativa (r = 0.73, N = 15, P = 0.002) indicates that timing of sampling (1.e., pre- vs. post-ingestion) did not markedly affect our results. Therefore, we combined data from provision and frass samples. The three types of Asteraceae were among the five most prevalent types in M. relativa cells. Strickler et al. (1996), who collected pollen samples from M. relativa cells in northern Michigan, found Astera- ceae from a diversity of genera, including Cirsium, to be the most prevalent pollen types; they also found pollen of Hyperica- ceae, Onagraceae, and Rosaceae. with Along Asteraceae and Rosaceae, we found five additional dicot families (Caprifoli- aceae, Fabaceae, Fagaceae, Hydrangeaceae, Oleaceae), as well as the two types of non- grass monocots. Along with numerous re- cords of visitations of M. relativa to dicot flowers, Hurd (1979) reported M. relativa visits to only one monocot family (Irida- ceae). We found no Iridaceae near our field sites, so the identity of the monocot pollen in our records remains a mystery. Medler and Koerber (1958) listed flowers in 23 spe- cies in 7 families visited by M. relativa in Wisconsin, although some of these may represent nectar-collecting rather than pol- len-foraging trips. We found considerable variation in the prevalence of different pollen types. Faga- ceae and Hydrangeaceae pollen were found in small amounts in just a few cells. In ad- dition, for some pollen types that were found in a large proportion of the cells, VOLUME 105, NUMBER | there may be a few cells in which its pres- ence in the provision was incidental (per- haps because the provisioning female used the pollen in previous cells). For example, among the 100 cells in which we identified Cirsium pollen (many at proportions = 50%), were 7 cells in which we estimated that it made up about 5% of the pollen. Oth- er pollen types found in only a few nests, were present in significant proportions in at least one cell. Although Oleaceae pollen was found in just two cells, it made up a minimum of 25% of the provision mass in one of these. Rosaceae pollen, though pre- sent in just six cells made up about 50% of the pollen in three cells (and approximately 90% in one of these). Both Brassicaceae and the unknown Fabaceae, present in nine cells each, were found once as approxi- mately half of a provision mass. Caprifoli- aceae pollen was found in 19 nests, but as approximately 25-50% of the provision in each. Thus, very few of the pollen types that we found can be excluded as purely incidental inclusions in M. relativa provi- sions. The types of pollen collected in 1999 (N = 48) and 2000 (N = 97) were similar (r = 0.72, N = 15, P = 0.002), but when comparing WB (N = 101) and SEB (N = 44), we found no correlation between the number of cells containing particular pollen types (r = 0.20, N = 15, P = 0.48). This difference resulted from a higher proportion of the WB cells containing the unknown di- cots (0.52 vs. 0.14; x? = 18.3, P < 0.001), the small monocot (0.45 vs. 0.2; x? = 19.6, P < 0.001), Lotus (0.40 vs. 0.09; x? = 13.5, P < 0.001), Taraxacum (0.41 vs. 0.07; x? = 16.5, P < 0.001), and the large monocot (0.25 vs. 0.05; x? = 8.3, P = 0.04). In con- trast, the WB cells contained a lower pro- portion of the unknown Asteraceae (0.11 vs. 0.69; x? = 78.0, P < 0.001), Medicago (0.07 vs. 0.48; x? = 32.7, P < 0.001), and Caprifoliaceae (0.0 vs. 0.43; x? = 50.2, P < 0.001). The discrepancies in pollen prev- alence between the WB and SEB samples may simply be due to differences in pollen 199 availability between the two sites. WB is adjacent to agricultural land, whereas SEB is in a neighborhood with ornamental plants. The number of pollen types per cell at WB (mean = 3.06 + 0.12) did not differ from the number per cell at SEB (mean = 3.02) O17 ¢ = 0:17; 143: dif. P= 0:87). Pollen provisioned by Megachile rotun- data. In 26 M. rotundata cells, we identified pollen from six dicots (Asteraceae, Capri- foliaceae, Fabaceae, and Rosaceae), in ad- dition to the two groups of monocot pollen and one unknown pollen type (Fig. 1). The M. rotundata nests were only several me- ters from two large plots of flowering al- falfa, and females were commonly seen for- aging on alfalfa (Ruth P. O'Neill, personal communication). However, we found alfalfa pollen in a smaller proportion of cells (0.42) than we did the large (1.00) and small (0.69) monocot pollen; all 26 cells examined contained at least one type of monocot pollen. The cells contained a mean of 3.1 + 0.4 types of pollen (range 1-7) and three contained only the large monocot pollen. Although Caprifoliaceae and Rosa- ceae pollen were each found in just one cell, the former made up 25% of the pro- vision in the cell, whereas the latter made up approximately half of the provision. Each of the remaining pollen types made up = 25% of the pollen grains in at least one cell (and often in greater proportions in numerous cells). Although our data are based on a small sample, the results indicate a_ relatively wide range of pollen types provisioned by M. rotundata. Our records (Asteraceae, Ca- prifoliaceae, Fabaceae, Rosaceae, and the two monocots) partially overlap with those observed in a lowbush blueberry agroeco- system, where M. rotundata provisioned not only with blueberry (Ericaceae, Vaccin- ium spp.), but also Asteraceae, Rosaceae, and Salicaceae (Stubbs et al. 1994). The mix of pollen used by M. rotundata nesting near blueberry and alfalfa indicates that, even when presented with an overwhelming predominance of a single pollen type, this 200 bee includes large proportions of other pol- len in its diet. Whether this mix represents a strategy of diet diversification or simply reflects some interaction between the rela- tive availability of and preferences for dif- ferent flowers remains to be determined. Although Horne (1995) demonstrated that M. rotundata forages for pollen on a wide variety of Fabaceae, she found that pollen preference did not correlate with success in offspring production on different pollen types. In controlled preference tests with over 200 species of 52 families, M. rotun- data was attracted to 21 species in 7 fami- lies, with high preference for Fabaceae (in- cluding Medicago), Lythraceae, Crassula- ceae, and Labiatae (Small et al. 1997). Sev- eral species of monocot (Liliaceae, A//iium) were also visited, although they showed rel- atively low attractiveness. However, flower visitation preferences records may not nec- essarily coincide with pollen preferences of M. rotundata. Pollen provisioned by Heriades carinata. Heriades carinata provisions included pol- len from eight of the categories we distin- guished, including Asteraceae, Caprifoli- aceae, Fabaceae, Tiliaceae, and both types of monocot pollen (Fig. 1). Cells contained a mean of 2.1 + 0.2 types of pollen (range 1-3). Cirsium was the most common pol- len, occurring in 13 of 15 cells and as 100% of the pollen in one cell. Along with a larg- er amount of Cirsium pollen, one cell con- tained pollen of Tiliaceae (approximately 10% of the pollen) which was not found in nests of the other two bee species. Each of the other seven pollen types made up = 25% of the pollen in at least one cell (and often in greater proportions in some cells). Analysis of pollen from nest cells in Mich- igan revealed ‘“‘almost entirely” staghorn sumac (Anacardiaceae, Rhus typhina L.) pollen (Matthews 1965); Hurd (1979) cites 11 families of dicots visited by H. carinata. SUMMARY AND CONCLUSIONS The types of pollen used by M. relativa, M. rotundata, and H. carinata overlapped, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON which is to be expected because all three species used a variety of pollen types and had a similar local flora available. Among the 15 pollen types found in M. relativa nests, 9 were also found in M. rotundata nests and 7 in H. carinata nests. The larger number of pollen types found in M. relativa nests is likely related to the larger number of cells sampled and greater number of sites at which it was studied. For all three spe- cies, the known range of pollen in provi- sions is much less than the known range of flowers visited (Matthews 1965, Hurd 1979, Small et al. 1997). It is difficult to know for particular cells whether pollen types present in low pro- portions represent |) a small number of pol- len-collecting trips to a particular plant spe- cies, 2) trips to flowers containing few pol- len grains, or 3) incidental inclusion of a pollen type picked up during a nectar-for- aging trip. Further, the proportion that rep- resents an incidental inclusion could vary among flower types. Due to variation in flower morphology and pollen placement, some pollen types could be picked up in- cidentally in large quantities during nectar visits, Whereas others may be transferred to the foraging bee in small numbers. Some of the pollen we identified, such as Fagaceae and Hydrangeaceae in M. relativa cells, may well have represented incidental inclu- sions of pollen picked up by nectar-foraging females. Alternatively, some of these re- cords may represent opportunistic pollen foraging on primarily nectar-gathering trips or exploratory visits to flowers by females seeking new pollen sources. Overall, we feel that it is safe to conclude that all three species foraged for pollen on variety of plant species. A relatively wide diet breadth is especially evident for M. relativa, given that six of the 15 pollen types each occurred in at least one cell as pure samples, whereas two others (Cirsium and Rosaceae) were found as nearly pure samples in individual cells. The same can be said for the other two species where several pollen types clearly made up at least 50% of the pollen VOLUME 105, NUMBER 1 in individual cells: 1) Cirsium, Medicago, the large monocot, Rosaceae, and the un- known family in M. rotundata cells and 2) Cirsium, Lotus, Medicago, the large mono- cot, and Caprifoliaceae in H. carinata cells. Nevertheless, because of uncertainties re- lated to possible incidental inclusion of pol- len, it is premature to use our records to precisely define the host ranges of these bee species, even at our sites. In addition, a complete analysis of the importance of each pollen type to the nutrition in developing bees will require estimates of individual pollen grain volume of different host spe- cies. A potential pollinator must visit the flow- ers of the crop species with a degree of con- stancy adequate to effect high levels of pol- len transfer. Even in agricultural systems, when the flowers of fruit or seed crops such as blueberry (Stubbs et al. 1994) or alfalfa predominate in close proximity to nests, M. rotundata may direct a high proportion of its pollen foraging trips to non-crop plant species. Horne (1995) found that M. rotun- data exhibited only moderate preference for alfalfa relative to birdsfoot trefoil (Loris caniculata L.) and crown vetch (Coronilla varia L.). Pollen records for M. rotundata suggest that control of alternative pollen sources could increase pollination efficien- cy in alfalfa seed crops, perhaps reducing the number of bees needed for commercial purposes. However, the types of pollen gathered by bees must be of nutritional quality adequate to sustain populations of the pollinator. Horne (1995) showed that pollen preference did not always correlate with reproductive success for M. rotundata given access to || species of plants. Thus, its moderate success on alfalfa relative to sainfoin and red clover suggests that in- creasing pollen source diversity could in- crease bee populations in agroecosystems, a goal potentially in conflict with that of increasing pollination efficiency on alfalfa. In addition, control of alternative pollen sources may negatively impact native pol- 201 linators, as well as parasitoids and preda- tors, that depend on the flowers. ACKNOWLEDGMENTS We thank William Kemp (USDA-ARS Bee Biology and Systematics Laboratory, Logan, UT) for providing trap nesting ma- terials, Ruth O’ Neill for providing the cells from her Megachile rotundata nests, and Pete Fay and Michael Ivie for use of their properties for trap-nesting sites. We also thank Sue Blodgett, Gregory Johnson, and William Kemp for providing advice at var- ious stages during the research, and Jordi Bosch, Jim Cane, Bill Kemp, Ruth O'Neill, and Norm Weeden for comments on the manuscript. This research was done in par- tial fulfillment of requirements for an M.S. in Entomology by Peter D. Jensen. LITERATURE CITED Free, J. B. 1993. Insect Pollination of Crops. Second Edition. Academic Press, New York, xii + 684 pp. Guirguis, G. N. and W. A. Brindley. 1974. Insecticide suscepubility and response to selected pollens of larval alfalfa leafcutting bees, Megachile pacifica (Panzer) (Hymenoptera: Megachilidae). Environ- mental Entomology 3: 691—694. Horne, M. 1995. Pollen preference and its relationship to nesting success of Megachile rotundata. Annals of the Entomological Society of America 88: 862— 867. Kapp, R. O. 1969. How to Know Pollen and Spores. WM. C. Brown Company Publishers, Dubuque, Iowa. Krombein, K. V. 1967. Trap-nesting Wasps and Bees: Life Histories, Nests, and Associates. Smithsonian Press, Washington, D.C., vi + 570 pp. Hurd, P. D., Jr. 1979. Apoidea, pp. 1741-2209. In Krombein, K. V., P. D. Hurd, Jr, D. R. Smith, and B. D. Burks, eds. Catalog of Hymenoptera in America North of Mexico, Vol. 2, Apocrita (Acu- leata). Smithsonian Institution Press, Washington D:e: Matthews, R. W. 1965. The biology of Heriades car- inata Cresson, Contributions of the American En- tomological Institute 1: 1—33. Medler, J. T. and T. W. Koerber. 1958. Biology of Me- gachile relativa Cresson in trap-nests in Wiscon- sin. Annals of the America 51: 337-344. Michener, C. D. 2000. The Bees of the World. The Entomological Society of 202 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Johns Hopkins University Press, Baltimore, xiv + 913 pp. Moore, P. D., M. E. Collinson, and J. A. Webb. 1991. Pollen Analysis, 2" Edition. Blackwell Scientific Publications, Boston, xiii + 216 pp. Richards, K. W. 1984. Alfalfa leafcutter bee manage- ment in Western Canada. Agriculture Canada Pub- lication 1945/E. Ministry of Supply and Services, Ottawa, 53 pp. Sawyer, R. 1988. Honey Press, Cardiff, 115 pp. Schmidt, J. O., S. C. Thoenes, and M. D. Levin. 1987. Survival of honey bees, Apis mellifera (Hyme- noptera: Apidae), fed on various pollen sources, Annals of the Entomological Society of America 80: 176-183. Identification. Academic Small, E., B. Brooks, L. P. Lefkovich, and D. T. Fairey. 1997. A preliminary analysis of the floral prefer- ence of the alfalfa leafcutting bee. Canadian Field- Naturalist. 111: 445-453. Strickler, K., V. L. Scott, and R. L. Fischer. 1996. Comparative nesting ecology of two sympatric leafcutting bees that differ in body size (Hyme- noptera: Megachilidae). Journal of the Kansas En- tomological Society 69: 26—44. Stubbs, C. S., E A. Drummond, and E. A. Osgood. 1994. Osmia ribifloris biedermannii and Megach- ile rotundata (Hymenoptera: Megachilidae) intro- duced into the lowbush blueberry agroecosystem in Maine. Journal of the Kansas Entomological Society 67: 173-185. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 203-208 NEW SYNONYMS AND STAGE DESCRIPTION FOR THREE SPECIES OF LEPTOHYPHIDAE (EPHEMEROPTERA) Davip E. BAUMGARDNER Department of Entomology, Texas A&M University, College Station, TX 77843-2475, U.S.A. (e-mail: dbaumgardner@tamu.edu) Abstract.—Vacupernius paraguttatus (Allen) and Allenhyphes michaeli (Allen) are new junior subjective synonyms of V. packeri (Allen) and A. vescus (Allen), respectively. Larval characters such as abdominal maculation are shown to be variable and unreliable in separating these species. Maxillary palp segmentation is difficult to detect among immature larvae of both species and must be used with caution when identifying immature leptohy- phid larvae to species. The adult stage of Homoleptohyphes mirus (Allen) is described for the first time based upon reared and field-associated specimens from southern Arizona. Key Words: phes, taxonomy The mayfly family Leptohyphidae (Ephemeroptera) is a New World family of mayflies distributed in North, Central, and South America, and the Carribean. Al- though the family is common and widely distributed, the species-level taxonomy of this family suffers from many problems that include a lack of larval and adult associa- tions, larval characters that are variable within and among populations and discrep- ancies in original descriptions and figures of species (Baumgardner and McCafferty 2000). Allen (1978) provided keys and de- scriptions for species of Leptohyphes. Re- cent publications by Lugo-Ortiz and Mc- Cafferty (1995), Wang et al. (1998), Baum- gardner and McCafferty (2000) and Wier- sema and McCafferty (2000) provided new or revised descriptions of taxa, synonyms, and distributional data. Despite these recent advances, many species-level problems re- main in Leptohyphidae. As part of an ongoing revision and cla- distic analysis of the family Leptohyphidae in North and Central America, two new ju- Ephemeroptera, Leptohyphidae, A//enhyphes, Vacupernius, Homoleptohy- nior subjective synonyms are presented for the family. Vacupernius paraguttatus and Allenhyphes michaeli are shown to be syn- onyms of V. packeri and A. vescus, respec- tively. In addition, the adult stage of Hom- oleptohyphes mirus is described for the first time based upon reared and field-associated specimens from Sonoita Creek, Santa Cruz County, in southern Arizona, near the type locality of the species. Depositions (and their acronyms) of ma- terials used in this study include: The Cal- ifornia Academy of Science, San Francisco (CAS); Texas A&M University, College Station (TAMU); Florida A&M University, Tallahassee (FAMU): Southwest Texas State University, San Marcos (SWTS); and Wilbur R. Enns Entomology Museum, Uni- versity of Missouri, Columbia (UMC). Vacupernius packeri (Allen) (Bies; J; 2) Leptohyphes packeri Allen 1967: 350. Leptohyphes phalarobranchus Kilgore and Allen 1973: 328; Allen 1978: 552 (syn.). 204 Leptohyphes paraguttatus Allen 1978: 552 New synonym. Vacupernius paraguttatus: Wiersema and McCafferty 2000. Vacupernius packeri: Wiersema and Mc- Cafferty 2000. Vacupernius packeri was described by Allen (1967) from a series of larvae col- lected in Honduras, and has been shown to be widely distributed throughout the south- western United States and Central America (Allen 1978, Henry 1986, Allen and Mur- vosh 1987, Lugo-Ortiz and McCafferty 1995). Henry (1986) associated the adult stage through rearing. In the larval stage, this species was distinguished from all oth- er described species of Leptohyphes sensu lato by distinctive maculations on the oper- culate gills (see Allen 1967; fig. 14) and vertex of the head; the presence of a 3-seg- mented maxillary palp; and the absence of what Allen referred to as the basal spine on the operculate gill, which is actually a col- orless outgrowth of the outer ventral la- mellae of gill 2 (Baumgardner and Mc- Cafferty 2000). Vacupernius paraguttatus, also described by Allen (1978), was based upon a single, immature larval specimen from the Hill Country of central Texas. This species was distinguished from other species of Lepto- hyphes sensu lato based primarily upon ab- dominal terga 2-4 that formed a “V-shaped pattern. Pale body and the presence of a 2-segmented maxil- lary palp were also considered important characters separating this species from oth- ers within the genus. Critical packeri larval specimens showed that a few of these specimens have the distinctive “V"-shaped maculation that was believed by Allen (1978) to be diagnostic of V. par- aguttatus. Unfortunately, the holotype of V. paraguttatus has become badly faded and no longer has the distinctive “*V”’-shaped pattern on the abdomen. However, this ““V"’-shaped pattern has been observed on maculations on examinations of series of V. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON some immature larvae of V. packeri. This abdominal mark has only been observed on immature specimens of V. packeri, indicat- ing developmentally influenced color pat- tern change. Abdominal larval markings similar to this have been found to be highly variable in other leptohyphid mayflies, such as L. zalope Traver, some of which display distinctive but variable abdominal markings (Baumgardner and McCafferty 2000). The distinctive maculations on the operculate gills of V. packeri are also absent in im- mature larvae, but begin to become visible in mature larvae. A few larvae of V. packeri were found to have both the abdominal “V"-shaped pattern and operculate gill maculation, providing the most compelling evidence to support the synonymy. Although Allen (1978) indicated that V. paraguttatus had a 2-segmented maxillary palp, careful re-examination of the slide- mounted mouthparts of the holotype showed the maxillary palp to be 3-seg- mented, but the intersegmental suture be- tween the second and third segment is dif- ficult to detect (Fig. 1). Examination of im- mature specimens of V. packeri also showed that they have a maxillary palp that appears to be 2-segmented. However, under high magnification a very faint suture is visible on the middle of the apical maxil- lary palp segment, as it is in the holotype of V. paraguttatus. For mature larvae, this intersegmental suture is distinct. Other lar- val specimens having both the distinctive “V" maculation of V. paraguttatus and the operculate gill markings of V. packeri were observed to have this weak suture between the second and third maxillary palp seg- ments. In mature specimens of V. packeri this suture is readily visible and distinct (Fig. 2). The above observations clearly support the synonymy of V. paraguttatus with V. packeri. Also of importance is the wide var- iation that occurs in immature specimens of this species, such as abdominal maculation and maxillary palp segmentation, characters which have historically been used to sepa- VOLUME 105, NUMBER 1 Figs. 1-4. larva, maxillary palp (arrow indicates location of weak 1, Leptohyphes paraguttatus, holotype suture line). 2, Vacupernius packeri, larva, maxillary palp (mature larvae). 3, Allenhyphes vescus, larva, maxillary palp. 4, Homoleptohyphes mirus, male adult, genitalia. rate numerous species of leptohyphid may- flies. This indicates that these characters should be used with caution when deter- mining species limits within the family Leptohyphidae. Type material examined.—Leptohyphes paraguttatus Allen: HOLOTYPE larva: Geronimo Cr., Guadalupe Co., Tex., 18-v- 73, Michael Peters; three associated slides (CAS #13603). Other material examined.—(AIl larvae, unless otherwise indicated.) UNITED STATES: ARIZONA: Yavapai Co., Wet Beaver Ck. at Wet Beaver Ck. Camp- ground, 24-25-y-1999, D.E. Baumgardner, (TAMU). TEXAS: Bandera Co., Winans 205 Creek, Hwy. 16 crossing, 12-iv-1992, Moulton & Stewart, 6 d, 2 2, (TAMU). Comal Co., Sattler, Rio Raft Co., Guada- lupe River at 5.5 mi. below Canyon Dam below 4th crossing., 26-x-1996, N. Wier- sema, (TAMU). Comal Co., Guadalupe River at Hwy. 311, 20-vi-1993, J. L. Cook, (TAMU). Milam Co., Rockdale, San Ga- briel River at Hwy. 487 crossing, 16-xi- 1996, N.A. Wiersema, (TAMU). Hays Co., San Marcos R. at Co. Rd. 101 (Caners Crossing), | mi. below conf. with Blanco R., in San Marcos City Limits, at Hays/ Caldwell Co. Line, 21-11-1997, DE Baum- gardner and DE Bowles, (TAMU). Medina Co., Seco Creek, 6 mi. S. of D’Hanis, 13- v-1993, L. Gilpin, V. Castillo, (SWTS, TAMU); Same but, 27-vi-1993, (SWTS, TAMU); Same but, 25-iv-1993, (SWTS). Milam Co., Rockdale, San Gabriel R. at 487 crossing, 16-xi-1996, N. Wiersema. Williamson Co., Georgetown, San Gabriel Park, below little dam and bridge, O7-x- 1996, N. Wiersema, (TAMU). BELIZE: STANN CK. DISTRICT, North Stann Creek, 2.7 mi. SE Middlesex on Humming- bird Hwy., 11-i-1996, R. W. Sites, (UM). MEXICO: NUEVO LEON, Cabazones R. at Hwy. 85, 15 mi. N. Linares, 16-v-1995, D. E. Baumgardner and B. C. Henry, (TAMU). Allenhyphes vescus (Allen) (Fig. 3) Leptohyphes vescus Allen 1978: 555. Leptohyphes michaeli Allen 1978: New synonym. Allenhyphes michaeli: McCafferty 2000. Allenhyphes vescus: Cafferty 2000. 549. Wiersema_ and Wiersema and Mc- Allenhyphes vescus was described by Al- len (1978) from the Hill Country of Texas based on a larva. Henry (1986) described the adults through rearing. The thin, deli- cate body and 2-segmented maxillary palp were considered diagnostic for the larval stage. Although not mentioned by Allen 206 (1978), the maxillary palp also has a diag- nostic terminal seta. In the same publication that Allen (1978) described A. vescus he also described A. mi- chaeli, based on a single larva from the Hill Country of Central Texas. Allenhyphes mi- chaeli was distinguished from other larvae then placed in Leptohyphes sensu lato by having a dark, median longitudinal line on abdominal terga 1—6, and a 1-segmented maxillary palp with a distinct apical seta. A careful study of numerous specimens of Allenhyphes vescus from the Hill Coun- try of Texas has clearly shown much vari- ability in the median longitudinal line char- acter. Specimens both with and without the median longitudinal line on abdominal ter- ga 1-6 were observed. Moreover, this ab- dominal feature gradually disappears as the larvae develop and no mature larvae of A. vescus were found to have the longitudinal line on abdominal terga 1—6. Although Allen considered A. michaeli to have a l-segmented maxillary palp, a care- ful re-examination of the holotype maxillae under high magnification indicated that a small intersegmental suture is present, and the palp is actually 2-segmented, as in A. vescus. In summary, larvae of Allenhyphes ves- cus have 2-segmented maxillary palps with an apical seta. The morphological details of the palp can most easily be observed under high (400) magnification (Fig. 3). Al- though both species were described in the same publication, A. vescus was chosen to be the senior name as the prerogative of the first revisor. Allenhyphes vescus is known throughout much of central Texas and is of- ten associated with streams in the Balconian ecoregion. It is also known from Nuevo Leon and Tamaulipas, Mexico. Type material examined.—Leptohyphes vescus Allen: HOLOTYPE larva: Rio Sa- binal at Utopia, Uvalde Co., Texas., 2-viil- 68, R. K. Allen; 2 slides (CAS #13607). Leptohyphes michaeli Allen: HOLOTYPE larva: North Fork Guadalupe Riv., 4 mi W. from PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hunt, Kerr Co., Tex., 27-vii-73, Michael Peters; 3 slides (CAS #13602). Other material examined.—UNITED STATES: TEXAS: Kimble Co., Llano R. @ Texas Tech Field Station, 07-xi-1998, DE Baumgardner, | male (reared), (TAMU). Val Verde Co., Dolan Falls Preserve, Devils River; The Nature Conservancy, O8-xi- 1998, DE Baumgardner, | male (TAMU). Comal Co., Honey Ck. in Honey Ck. St. Natural Area in Guadalupe R. State Pk, 08- ii1-1997, DE Baumgardner & DE Bowles, I male (reared), larvae (TAMU). William- son Co., Georgetown, San Gabriel Park, rif- fles below the little dam on the San Gabriel River, 2-x-1996, N. Wiersema, larvae (TAMU). Val Verde Co., Devils River, Do- lan Falls, 19-x-1993, C. R. Nelson & S. M. Stringer, larvae (TAMU). MEXICO: TA- MAULIPAS, spring (at local park) (at Hi- dalgo Ave?) near town of Jaumave, off Hwy. 101; 16-v-1995, 2 males, BC Henry & DE Baumgardner. TAMAULIPAS, R. Guayaalejo (Tamasi) off Hwy 247 nr. San Ignacio, 26-v-1993, B. Henry, larvae. Homoleptohyphes mirus (Allen) (Fig. 4) Leptohyphes mirus Allen 1967: 353. Homoleptohyphes mirus: Wiersema and McCafferty 2000. Male adult.—Body length: 3.0—4.5 mm. Forewing length 4.5—5.0 mm. Hindwing ab- sent. Cerci length 8.0—-10.0 mm. Overall coloration dark reddish brown with pale lat- eral markings. Head dark brown, with black over shading. Compound eyes large; width of one eye greater than distance between eyes. Antenna brown at base, pale apically. Pro- and mesonotum reddish brown with pale lateral markings. Metanotum shiny reddish brown to dark brown. Forefemur pale with dark brown stippled longitudinal bands; apical segments pale. Meso- and me- tafemora pale with dark brown to black lon- gitudinal bands; apical segments pale with brown stippling. Foreclaws similar and blunt. Forewings translucent with darkened VOLUME 105, NUMBER 1 subcostal vein. Abdomen pale to dark red- dish brown, similar in color to thorax; cerci pale with grey over shading. Genitalia as in Fig. 4; penes with shallow emargination and scattered small spines; subgenital plate with shallow emargination, covered with minute spines (Fig. 4). Female adult.—Body length: 3.0—4.0 mm. Forewing length 4.5—5.0 mm. Hind- wing absent. Cerci missing. Body colora- tion and markings similar to male. Com- pound eyes small and widely separated. Comments.—In both the larval and adult stages, H. mirus is most similar to H. di- morphus (Allen), because male larvae and adults have large compound eyes. These are the only two known species of Leptohy- phidae in North or Central America that have this feature. In the larval stage, H. mi- rus can be differentiated from H. dimorphus by the presence of short spines on the fore- femur of H. mirus. In contrast, H. dimor- phus has long setae on the forefemur. Dif- ferentiating adults of H. mirus from H. di- morphus can be difficult without associated larvae. Adults of H. mirus apparently can be separated from those of H. dimorphus by the mostly solid dark red-brown coloration of the abdominal sternites, with limited black overshadowed stippling. In H. dimor- phus, coloration of the abdominal sterna is pale yellow to reddish brown, overshad- owed by extensive, fine black stippling. Homoleptohyphes mirus is known only from the southwestern United States (far west Texas and southern Arizona) and northwestern Mexico, from the states of So- nora, Chihuahua and Sinaloa. Larvae can be found in small, apparently permanent, spring-fed desert streams. Upon reexami- nation, the specimen identified by Allen (1978) as L. ferruginus Allen and Brusca (now a synonym of L. zalope) from the Rio Sonora, Sonora, Mexico is clearly that of H. mirus and not L. zalope. Type material examined.—Leptohyphes mirus Allen: HOLOTYPE: Rio Blanco, Ar- izona, 3-iv-37, J. G. Needham, | male larva (FAMU). ALLOTYPE: Rio Blanco, Ari- 207 zona, 3-iv-37, J. G. Needham, | female lar- va (FAMU). PARATOPOTYPES: Rio Blanco, Arizona, 3-iv-37, J. G. Needham, 2 larvae (CAS), 4 larvae (FAMU). Other material examined.—UNITED STATES: ARIZONA: Santa Cruz Co., Son- oita Cr., nr. Patagonia, 15-11-1997, J. Slu- sark and K. Byrnes, 38L, 4 males (TAMU); Santa Cruz Co., Sonita Cr. at Blue Haven Rd., ca. 1 mi. SW Patagonia (31°30'57’N; 110°47'35"W), 06-vi-2000, DE Baumgard- ner, 52 larvae 3 males (reared) and 2 fe- males (reared) (TAMU). TEXAS, Brewster Co., Calamity Cr. at TX Hwy 118, ca. 22 mi. S. Alpine, 24-vili- 1996, DE Baumgard- ner & DE Bowles, 2 larvae (TAMU); Jeff Davis Co., H.C. Espy Ranch, Farm Rd. 1832, 14-v-1973, RG McClure, 20 larvae (TAMU). MEXICO: CHIHUAHUA, Rio San Pedro at Meoqui on Hwy. 45, 14-viti- 77, R. K. Allen, 6 larvae (CAS); CHIHUA- HUA, Rio Satevo at Gral. Tris on Hwy. 16, 13-vili-77, RK Allen, 9 larvae (CAS); CHI- HUAHUA, Rio Papagochic, 4 mi. E. Ciudad Guerrero on Hwy 16, 13-vili-77, R. K. Allen, 2 larvae (CAS); SINALOA, Rio Baluarce at Rosarito, 13-i-83, Allen, Mur- vosh, | larva (CAS); SINALOA, stream | mi. N. El Viola, 18-I-83, Allen and Mur- vosh 6 larvae (CAS); SONORA, Rio So- nora 2 mi. SE Rte. 21 between Uras & Ma- zocahui, 14-1-83, Allen & Murvosh, | larva (1 slide) (CAS) (record previously pub- lished as L. ferruginus by Allen 1978); SO- NORA, Rio Bavispe, 3 mi. SW Colonia Moralia at dam, 2-1-1983, Allen, Murvosh, 12 larvae (CAS); SONORA, Rio Altas at Tubutama, 13-I-83, Allen and Murvosh, 5 larvae (CAS). ACKNOWLEDGMENTS I thank Vince Lee (California Academy of Sciences) and Janice Peters (Florida A&M University) for providing Leptohy- phes type specimens, Nick Wiersema (Aus- tin, TX), Jerry Cook (Sam Houston State University) and Robert Sites (University of Missouri, Columbia) for providing addi- tional specimens of Leptohyphidae, and 208 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Abigail Fuller for assistance with figures. Special thanks are due to Joe Slusark (Unit- ed States Geological Survey) for providing additional specimens of Leptohyphes mirus from Arizona and David Bowles (Texas Parks and Wildlife) for his field assistance on several collecting trips. Reviews by Steven Burian (Southern Connecticut State University), Jan Peters (Florida A&M Uni- versity) and two anonymous reviewers greatly improved the final manuscript. Par- tial funding was provided by the Entomol- ogy Graduate Student Organization of Tex- as A&M University. LITERATURE CITED Allen, R. K. 1967. New species of new world Lepto- hyphinae (Ephemeroptera: Tricorythidae). Cana- dian Entomologist 99: 350-374. 1978. The nymphs of North and Central American Leptohyphes (Ephemeroptera: Tricory- thidae). Annals of the Entomological Society of America 71: 537-558. Allen, R. K. and C. M. Murvosh. 1987. Mayflies (Ephemeroptera: Tricorythidae) of the southwest- ern United States and northern Mexico. Annals of the Entomological Society of America 80: 35—40. Baumgardner, D. E. and W. P. McCafferty. 2000. Lep- tohyphes zalope (Ephemeroptera: Leptohyphidae): A polytypic North American species. Entomolog- ical News 111: 49-59. Henry, B. C., Jr. 1986. Mayflies (Ephemeroptera) of the Concho River, Texas. The Southwestern Nat- uralist 31: 15-21. Kilgore, J. I. and R. K. Allen. 1973. Mayflies of the Southwest: New species, descriptions, and records (Ephemeroptera). Annals of the Entomological Society of America 80: 35—40. Lugo-Ortiz, C. R. and W. P. McCafferty. 1995. Con- tribution to the taxonomy of the Leptohyphidae (Insecta: Ephemeroptera) of Central America. Studies on Neotropical Fauna and Environment 30: 165-176. Wang, T. Q., R. W. Sites, and W. P. McCafferty. 1998. Two new species of Leptohyphes (Ephemeroptera: Leptohyphidae) from Ecuador. Florida Entomol- ogist 81: 68-75. Wiersema, N. A. and W. P. McCafferty. 2000. Generic revision of the North and Central American Lep- tohyphidae (Ephemeroptera: Pannota). Transac- tions of the American Entomological Society 126: 337-371. PROC. ENTOMOL. SOC. WASH 105(1), 2003, pp. 209-224 BIODIVERSITY AND BIOGEOGRAPHY OF MEXICAN BUTTERFLIES (LEPIDOPTERA: PAPILIONOIDEA AND HESPERIOIDEA) ARMANDO LUIS-MARTINEZ, JORGE LLORENTE-BOUSQUETS, ISABEL VARGAS-FERNANDEZ, AND ANDREW D. WARREN (ALM, JLB, IVF) Museo de Zoologia, Facultad de Ciencias [MZFC], Universidad Nacional Aut6noma de México, Apdo. Postal 70-399, México 04510, D.F, México (e-mail: alm@hp.fciencias.unam.mx); (ADW) Department of Entomology, Oregon State University, Corvallis, OR 97331-2907 (e-mail: warrena@science.oregonstate.edu). Abstract.—We present a brief history of the scientific study of butterflies in Mexico, which began in the 18th century, and provide an overview of the holdings of Mexican butterfly specimens in institutional collections worldwide. The current status of a massive database on Mexican butterflies, at present containing over 450,000 distributional records, is detailed. Around 1800 species of butterflies have been reported from Mexico, which equals roughly 10% of the world butterfly fauna, placing Mexico among the ten most butterfly-rich countries in the world. About 14.8% of Mexico’s butterfly species are en- demic, and six genera appear to be endemic to Mexico. For the first time, we present a list of all endemic Mexican butterfly taxa: 88 species and 150 additional subspecies of Papilionoidea, and 171 species and 14 additional subspecies of Hesperiidae. Patterns of species richness and endemism are not alike; the richest areas are in the southeastern part of Mexico, mostly in the tropical evergreen forests, whereas endemism is greater in the arid north, and the humid montane forest in the central and southern parts of the country. The twenty richest sites for butterfly diversity in Mexico are identified, and comparisons are made between these sites and regions. Key Words: Rhopalocera, Papilionidae, Pieridae, Nymphalidae, Lycaenidae, Hesperi- idae, endemism, species richness, biogeographical patterns, diversity Scientific study of Mexican butterflies began with the Royal Scientific Expeditions to New Spain in the late 18th and early 19th centuries (Llorente et al. 1994). Since then, most lepidopterological research has been aimed at efforts to document the biodiver- sity of the country (Llorente and Luis 1993). Despite these efforts, the knowledge gathered in the last two centuries on the biodiversity and biogeography of Mexican butterflies remains incomplete. Considering the vast biodiversity that exists in Mexico, the continued existence of completely un- explored regions is not surprising, and the scarcity of institutional collections contin- ues to be a hindrance to efforts at furthering our knowledge (Llorente et al. 1996). Worldwide, about 18,000 species of but- terflles are known, which represent about 13% of the approximately 150,000 species of Lepidoptera [Shields (1989) estimates 17,280 butterfly species, Heppner (1991) estimates 19,238 butterfly species, Robbins and Opler (1997) estimate 17,500 total but- terfly species]. Data from Shields (1989) and Heppner (1991) estimate that 1,800 species of butterflies inhabit Mexico, rep- resenting around 10% of the world total 210 Table 1. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Richness of Mexican butterfly species, by family, compared to major biogrographical regions. Numbers are taken from Shields (1989) and Heppner (1991). N = Nearctic; M = Mexico; NL = Neotropical; P = Palaearctic; E = Ethiopian; O = Oriental; A = Australia/Oceania. About 13.1% of the species of Lepidoptera are butterflies. Family N M NL P E oO A Total Hesperiidae 290, 800 2,016 155 437 569 19] 3,658 Papilionidae 33 56 120 84 87 178 70 572 Pieridae 64 90 323 167 174 307 187 E222 Lycaenidae 164 430 2.611 407 1,413 1,540 429 6,564 Nymphalidae 214 440 2,857 1,083 1,156 1,563 349 7,222 Total 765 1,816 7,927 1,896 3,267 4,157 1,226 19,238 (see Table 1). This richness may be due to two facts: 1) Mexico is located in an area of tectonic convergence (termed the Mexi- can Transition Zone by Halffter (1976), also see Llorente (1996)), where the Nearctic and Neotropical regions overlap; together these regions contain 40% of the world’s butterflies; and 2) Mexico has an extratrop- ical-intertropical situation, with various mountain ranges, which generate a wide va- riety of climates (ranging from arid to hu- mid), as well as many vegetation types ranging from xerophilous scrubs to tropical forests and various types of temperate for- ests (Morrone et al. 1999). The Hesperioidea is the largest group of butterflies; a single family (Hesperiidae) is included, with about 3,600 species. Accord- ing to Heppner (1991), the Papilionoidea include four families, which are, from larg- est to smallest: Nymphalidae, Lycaenidae, Pieridae, and Papilionidae (see Table 1) [following Weintraub and Miller (1987), we do not consider the Hedylidae to be butter- flies]. Considering all biogeographical re- gions, the Neotropical Region 1s the richest in butterfly species, with the exception of the Papilionidae, which are more diverse in the Oriental Region (Heppner 1991). Mex- ico harbors more than twice the number of species than the entire Nearctic Region north of Mexico. This figure is considerably larger than that of the Australian Region and similar to the number of species in the Palaearctic Region. Several groups have di- versified extensively in Mexico, and there are also a number of paleoendemic and re- lictual groups, such as Baronia Salvin, 1893; Eucheira Westwood, 1834; Prestonia Schaus, 1920; Chlosyne Butler, 1870; Cyl- lopsis R. Felder, 1869; Paramacera Butler, 1868; Lamphiotes Callaghan, 1982; Zobera H. A. Freeman, 1970; Piruna Evans, 1955; Paratrytone Godman, 1900; and all mega- thymine genera (especially Agathymus H. A. Freeman, 1959 and Stallingsia H. A. Freeman, 1959). MATERIALS AND METHODS Collections of Mexican butterflies.—The number of butterfly specimens in nine of the largest American collections exceeds 72,000 (see Table 2). Two of the three most important collections in Mexico are housed at UNAM, the collection of the Departa- mento de Zoologia of the Instituto de Biol- ogia (IBUNAM), and the collection at the MZEFC, which has specialized in the fauna of the Mexican humid montane forests (Na- varro and Llorente 1996). Together, both collections contain over 100,000 mounted specimens and another 250,000 specimens papered in envelopes (Table 2). The third most important collection in Mexico is pri- vate and is the product of research con- ducted by members of the de la Maza fam- ily, which, according to G. Lamas (personal communication) [information obtained orally], is composed of approximately 35,000 specimens. Three other important private collections that have been assem- bled during the last three decades are those VOLUME 105, NUMBER | Table 2. 211 Numbers of Mexican butterfly records from major institutional collections. All numbers are trom the database of butterfly localities maintained by the **Alfonso L. Herrera’? Museum of Zoology, at UNAM in Mexico City. Collection # of Records Allyn Museum of Entomology, Sarasota, FL American Museum of Natural History, New York, NY California Academy of Sciences, San Francisco, CA Carnegie Museum of Natural History, Pittsburgh, PA Los Angeles County Museum of Natural History, CA San Diego Natural History Museum, CA University of California, Berkeley, CA National Museum of Natural History, Washington, DC Nevada State Museum, Las Vegas, NV The Natural History Museum, London Museo de Zoologia **Alfonso L. Herrera,”’ Instituto de Biologia, UNAM, Mexico City Colegio de la Frontera Sur (Chetumal, QROO) TOTAL UNAM 9,035 11,530 5,330 14,278 9,484 8,322 2,394 7,804 3,850 15,000 300,000 55,000 30,000 472,027 of Diaz Francés, Jesus Saldana, and Gon- zalez Cota. include a wide variety of material from all the south- ern Mexican states, and contain many spe- cies endemic to Mexico. Fortunately, the These collections two former collections were recently ac- quired by IBUNAM, and the latter, by the MZEC, sponsored by the Comision Nacion- al para el Conocimiento y Uso de la Bio- diversidad (CONABIO), thus ensuring their preservation in academic institutions. Mexican butterfly database.—Over the past 15 years, one of the primary objectives of the Lepidoptera collection at the MZFC has been the computerization of Mexican butterfly records into a database. This da- tabase contains three groups of primary data, including data from specimen labels of Papilionoidea housed in Mexican insti- tutions, the same data from Papilionoidea housed in major institutions in the U.S.A. and England, and data of Papilionoidea and Hesperioidea from over 1,000 literature sources (Table 3). Data from specimens of Mexican Hesperioidea housed in_ institu- tional collections are being added to the da- tabase. The database is currently supported by the “Sistema de Informaci6n Bidotica,”’ a program developed by CONABIO to manage nomenclatural, geographic, biblio- graphic and curatorial information on all of Mexico’s biota. At present, the butterfly database con- tains over 450,000 records, of which almost 85,000 are from natural history museums outside of Mexico, mainly in the U.S.A. and England. About 55,000 of these records are from IBUNAM, a minimum of 10,000 are from literature reports, and the remain- ing records are from faunal studies or spe- cific collecting efforts aimed at determining the distributions of certain taxa (Luis et al. 2000). The records in this database have enabled us to examine the distributions of many of the species that make up the Mexican fauna (Llorente et al. 1997). One of the best stud- ied butterfly faunas in Mexico is that of the western state of Michoacan. Through stud- ies made during the last 10 years, over 100,000 records have been obtained from Michoacan, trom 144 localities (Luis et al., in preparation). In contrast, despite its long history of butterfly collecting tracing back to the [8th century, the state of Veracruz is represented by 50,000 records from 562 lo- calities. To supplement this database of lo- cality information, three additional compi- lations have been or are being prepared: a catalogue of taxonomic names of Mexican Table 3. Numbers of localities from which butter- flies have been recorded in each Mexican state. All numbers are from the database of butterfly localities maintained by the **Alfonso L. Herrera’’ Zoology Mu- seum, at UNAM in Mexico City. State Localities VERACRUZ 562 BAJA CALIFORNIA SUR 547 BAJA CALIFORNIA 520 CHIAPAS 472 OAXACA 409 GUERRERO 228 SAN LUIS POTOSI 215 SINALOA 198 SONORA 196 JALISCO 178 DURANGO 157 MICHOACAN 144 TAMAULIPAS 139 NAYARIT 139 CHIHUAHUA 137 NUEVO LEON 125 HIDALGO 117 PUEBLA 96 COLIMA 96 MORELOS 91 YUCATAN 72 QUINTANA ROO 65 ESTADO DE MEXICO 63 DISTRITO FEDERAL 56 CAMPECHE 40 TABASCO 38 COAHUILA 35 QUERETARO 19 ZACATECAS 19 AGUASCALIENTES 17 GUANAJUATO 15 TLAXCALA 4 TOTAL LOCALITIES 5,209 butterflies, one of geographical localities of Mexican butterflies (Luis et al. 1996), and One containing literature citations that deal with Mexican Papilionoidea (Luis et al. 2000). GEOGRAPHICAL DISTRIBUTION OF MEXICAN BUTTERFLY RECORDS Although Mexico is a large Latin Amer- ican nation, thought by some to be relative- ly well known and explored, this is not true for insects (Burke and Fryxell 1995). Ac- cording to the data from nearly 500,000 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON specimens held in nearly a dozen museums in the U.S.A., The Natural History Muse- um, London, Mexican collections, and re- ports from the literature, the number of Mexican localities that have been explored for butterflies since the beginning of the former century is just over 5,000. Such a representation is poor for a county like Mexico which contains a vast megadivers- ity of organisms (Llorente and Luis 1993). Published distributional records on Mex- ican butterflies are varied in their scope. Reports range from occasional collections made during a few hours (e.g., Comstock 1959; Diaz 1975; Guzman 1975; Gonzalez 1977, 1978), to comprehensive faunistic studies (e.g., Luis and Llorente 1990; Luis et al. 1991, 1996; Vargas et al. 1994, 1999), with many reports that fall somewhere in between (e.g., Clench 1968, Freeman 1969, de la Maza 1976, Routeledge 1977, Brown et al. 1992, Balcazar 1993). For the Papi- lionidae and Pieridae (Llorente et al. 1997), it has been observed that the best studied sites are near, or geographically associated with, classic historical collecting sites such as the regions of Jalapa and Los Tuxtlas in Veracruz, the Sierra de Juarez in Oaxaca, and the Sierra Madre del Sur in Guerrero (Sierra de Atoyac de Alvarez, Chilpancingo and Omiltemi). Our state of knowledge of the geograph- ical distribution of butterflies within Mexi- co’s political boundaries remains poor; of the 31 states comprising the Republic, six states contain 52.56% of the recorded 5,209 collecting localities (Veracruz, Baja Cali- fornia Sur, Baja California, Chiapas, Oa- xaca and Guerrero). States with fewer than twenty recorded collecting localities and no published faunal studies include Tlaxcala, Aguascalientes, Zacatecas and Querétaro (Table 3). However, given that a substantial number of Mexican species of papilionoids are known to have wide distributions, and several highly localized endemic species are known, it seems valid to assume that the primary general distributional patterns displayed by Mexican butterflies have been VOLUME 105, NUMBER | Table 4. Numbers of butterfly species known from various Mexican states. BC = Baja California; BCS = Baja Calfornia Sur; DGO = Durango; COL = Colima; JAL = Jalisco; GRO = Guerrero; VER = Veracruz; OAX = Oaxaca; CHIS = Chiapas; QROO = Quintana Roo. Family BC BCS DGO Col JAL GRO VER OAX CHIS QROO Papilionidae 8 6 12 28 28 32 40 a1 42 24 Pieridae 26 2] 29 36 44 44 Dy 64 70 27 Nymphalidae 34 23 61 135 176 209 300 371 379 116 Lycaenidae 53 30 45 125 151 192 219 285 333 64 Hesperiidae 39 35 104 221 258 300 435 416 462 118 Total 159 121 251 545 657 TARE es 1,187 1,276 349 identified. Nevertheless, a large number of faunal studies remain to be conducted in or- der to understand at a finer level the distri- bution of species and groups which are en- demic to Mexico (Llorente et al. 1994, 1997; Soberon et al. 2000; Onate et al. 2000). According to the literature, museum col- lections, and recent faunal studies, less than one third of the Mexican states have a but- terfly species checklist (Vargas et al. 1996, Warren et al. 1998). Additionally, most of the existing state checklists are based on one or two faunal studies centered in the most diverse parts of those states (leaving large parts of those states un-collected). Ex- ceptions to this are the states of Baja Cali- fornia and Baja California Sur, where no rigorous faunal studies have been made but for which Brown et al. (1992) compiled in book-form numerous data from the litera- ture, museum collections and field work conducted at several localities. Table 4 pro- vides numbers of butterfly species from states for which we have species checklists, and Map | shows Mexico’s ten richest states, in numbers of Papilionoidea and Hesperioidea species. PATTERNS OF SPECIES RICHNESS The nine richest regions in Mexico, in terms of butterfly diversity, are shown on Map 2. Each of these regions includes from three to more than 30 individual collecting localities that can arguably be classified as single biotic units. In general, these regions contain various altitudinal, climatic, and vegetational gradients, such as the Sierra de Atoyac de Alvarez in Guerrero (with 339 Papilionoidea species), and the Sierra de Juarez in Oaxaca (with 450 Papilionoidea species), which ranges from 100 to 3,100 m. and includes many forest types. Sadly, in recent years, we have observed a pro- found alteration of these habitats, due to hu- man activities, which has resulted in wide- spread ecological fragmentation and de- struction. Several measurements should be consid- ered when trying to determine the most di- verse of these areas; namely the size of the area, variations in elevation, environmental heterogeneity, and the biogeographical his- tory of the region. Luis et al. (1991) shows that the Sierra de Juarez in Oaxaca is the region showing the widest altitudinal range (100—3,100 m), contrasted with the follow- ing regions: a) Los Tuxtlas, Veracruz (with 516 Papilionoidea species), with elevational extremes from sea level to 900 m, but with over 30 sampled localities and various veg- etation types (Raguso and Llorente 1997), b) Chajul, Chiapas (with 396 Papilionoidea species), composed of a single forest type at 300 m. elevation (de la Maza and de la Maza 1985a, b), and c) Presidio, Veracruz (with 392 Papilionoidea species). Most of the records from the Presidio area are his- torical (up to 150 years old); forests in this area have been altered considerably, and only a few sites remain that still host native vegetation. The four richest single Mexican localities for butterflies are located on the Atlantic 214 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mexican states with greatest diversity of Papilionoidea and Hesperioidea | \ ‘ eee et \ id = = Lat & ¢ XN is ea y+ fh mY Ne She Sy \ 4 ne 4 ¢ ~— x - \ Ni ie \ \ oS on a = ~ - x \a rc =~ ) \ ee ; f paas aes ad as Lan > \ AS \ cs Map |. slope, which is the most diverse region in Mexico for several groups of organisms (Escalante et al. 1998, Flores 1998). Mex- ico’s richest sites on the Pacific slope in- clude humid areas with great physiograph- Mexico’s localities ical heterogeneity (Map 3). Along Pacific slope there are only three that have more than 200 species. These are: Mismaloya and La Calera in Jalisco, and Rio Santiago in Guerrero, all with tropical that contain ments from the montane cloud forest (Mon- teagudo et al. 2001). While Mismaloya does not appear on Map 3 (see Map 2), Warren and Llorente (1999) reported 315 butterfly species from that site (including Papilionoidea and Hesperioidea). semi-deciduous forests ele- PATTERNS OF ENDEMISM The patterns of species richness and en- demism among Mexican butterflies differ. States 1. Chiapas 2. Oaxaca 3. Veracruz 4. Guerrero 5. Michoacan Ss 6. Jalisco 7. San Luis Potosi 8. Puebla 9. Tamaulipas 10. Colima Numbers of Papilionoidea and Hesperioidea species known from Mexico’s ten richest states. The tropical evergreen forests of southeast- ern Mexico comprise the richest region in Mexico, in terms of total numbers of spe- cies. Over 50% of the species of Papilion- oidea that occur in Mexico (over 700 spe- cies, excluding Hesperioidea) are known to occur in these forests (Salinas 1999). De- spite this great diversity, only about 2% of the butterfly species associated with the tropical evergreen forests are endemic to Mexico; most species in these forests range south into parts of Central and South Amer- ica. According to Rzedowski (1978), trop- ical evergreen forests formerly occupied 12.8% of Mexico; the current remaining forest fragments cover at most between 10 and 15% of their original area (Granillo 1985, Toledo 1988). Mexico’s endemic fauna, in contrast, is associated both with the arid communities of northwestern Mexico, and with the humid VOLUME 105, NUMBER 1 Mexican regions with greatest diversity of Papilionoidea Regions - Species | 1. Onzaba-Cordoba-Fortin de las Flores | 537 2. Region de los Tuxtlas 516 puseso oe 3. Jalapa-Coatepec-Teocelo 486 | \ (i : 4. Sierra de Juarez 450 3 i a = | ae = | 5, Sierra de Atoyac 339 | , | y ian 6. Sierra de Manantlan 315 | : { 7. Mismaloya-Bahia de Banderas 281 ae 2) 8. Acahuizotla 262) =< YY 9. Sierra de San Juan 237 | £ Sy, a ; J WO 7 ae a -_ _— : e i > a ' , ee - x ie ae | ‘\ \ : y { ra en = 7 £6 & ‘4 werk A a, : a 5@@8 4@ 6)" Beans 7 aol my Map 2. Numbers of Papilionoidea species known from Mexico’s most diverse regions. Each region includes from three to more than 30 separate collecting sites montane forests (especially the montane cloud forests) of central and southern Mex- ico. The insular distribution of the montane cloud forests along various mountain chains in Mexico has resulted in the speciation of many taxa, a phenomenon evident in many plant and animal groups (Halffter 1987). Ac- cording to Llorente (1984), there are two al- titudinal barriers in Mexico which limit dis- persion and generally prohibit continuous distributions of taxa: one around 600 m, and the other around 2,000 m. Each of these al- titudinal barriers present dramatically differ- ent climatic and vegetational conditions. Throughout Mesoamerica, the lower altitu- dinal zone is composed of the tropical ev- ergreen and semi-deciduous forests on the Atlantic slope, with tropical deciduous and semi-deciduous forests on the Pacific slope. The middle elevational zone (roughly 600— 2,000), is significantly cooler than the lower zone and is occupied by various humid for- est types. The highest elevational zone is seasonally cold, often arid, and is dominated by fir, pine and oak forests. The insularity of the humid montane for- ests in Mexico has made them spots of high endemicity for several groups of butterflies at the specific and subspecific levels. The assemblages of subspecies scattered in a number of these submontane islands some- times group together with tropical Mesoam- erican species, whereas in other cases they are more closely related with montane Cen- tral American species. In Mexico, there are six discernible biogeographical “‘islands”’: a) the highlands of Chiapas [and Guate- mala]; b) the highlands of the Los Tuxtlas region in Veracruz; c) the Sierra de Juarez- Sierra Madre Oriental mountain chains in 216 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mexican localities with greatest diversity of Papilionoidea Map 3. Veracruz and Oaxaca; d) the Sierra Madre del Sur mountain chain in Guerrero and Oa- xaca; e) the Pacific slope of the Nueva Ga- licia region from Colima to southern Sina- loa; and f) in lesser degree, the Transmex- ican Volcanic Belt. Some of lands” may be further subdivided into two or three portions, such as the highlands of Chiapas and Guatemala (Llorente and Es- calante 1992). We know of 88 species and 150 addi- tional subspecies of Papilionoidea that are endemic to Mexico, with three endemic genera (Baronia, Prestonia and Eucheira). These endemic species and subspecies in- clude 11.7% of Mexican butterfly taxa. Papilionoid genera which have diversified extensively in Mexico include Chlosyne, Cyllopsis, and Calephelis Grote and Rob- inson, 1869. Currently, 171 species and 14 subspecies of Hesperioidea are known to be these “‘is- Localities S 1. Chajual, Chiapas 396 2 Presidio, Veracruz 392 | 3. Cordoba, Veracruz 340 4 Laguna de Catemaco, Veracruz} 332 | 5 Metates, Oaxaca 301 6 Tamazunchale, San Luis Potosi] 270 7. Barranca de Cayoapa, Veracrug 270 8 Dos Amates, Veracruz | 254 9 Fortin de las Flores 250 | 10. La Calera, Jalisco 240 | 11. Teocelo, Veracruz 238 4 | 12. Puerto Eligio, Oaxaca 229 \" | 13. Onzaba, Veracruz 228 { py \2 | 14. Rio Santiago, Guerrero 225) A | 15. Yaxchilan, Chiapas 7235\\ || Numbers of Papilionoidea species known from the twenty richest localities in Mexico. endemic to Mexico, with three endemic genera (Zobera, Aegiale and Turnerina). These endemic hesperioids include 9.1% of Mexican butterfly taxa. Hesperioid genera that have diversified extensively in Mexico include Piruna, Paratrytone and Agathy- mus. The number of endemic species of Hesperioidea exceeds that of all four papi- lionoid families combined. Considering all 423 taxa considered to be endemic in this work (including species and subspecies), 43.7% are hesperioids (Appendix). Thus, overall, 14.8% of Mexican butterfly species are endemic, and over 60% of these endem- ic taxa are distributed in the Mexican Pa- cific region. The Pacific region is environ- mentally discontinuous with other Mesoam- erican and Central American faunal ele- ments, and possesses two well delimited biogeographical islands: the Sierra Madre del Sur in Guerrero and Oaxaca, and the VOLUME 105, NUMBER 1 Pacific slope of the Nueva Galicia region from Colima to southern Sinaloa. The Te- huantepec Isthmus and the Balsas River Ba- sin, respectively, play the role of physio- graphic and climatic barriers to these is- lands (Llorente 1984). CONCLUSIONS We summarize our biogeographical knowledge of Mexican butterflies, acquired through the past 200 years, as follows: 1) Mexico possesses 10% of the butterfly species of the world and 14.8% of these species are endemic, making Mexico one of ten countries with the greatest butterfly diversity. 2) Mexico and the areas adjacent to its bor- ders possess relictual, palaeo- and neo- endemic butterfly groups of great sci- entific interest, mostly in the northwest- ern arid zones, and in the montane com- munities to the south. 3) Patterns of species richness and ende- mism are not alike; the richest areas are in the southeast, mostly in the tropical evergreen forests, whereas endemic taxa are mostly distributed in the arid north, and in the humid montane forests in the central and southern portions of the country. Areas richest in species numbers and en- demic taxa are those exhibiting physio- graphic, climatic and vegetational het- erogeneity, in a mosaic of preserved and partially altered environments, (such as the Los Tuxtlas region in Veracruz, Cha- jul in Chiapas, and the Sierra de Juarez in Oaxaca, each one possessing Over one third of Mexico’s butterfly species). In 4 contrast, whole states or physiographic provinces with less environmental het- erogeneity, such as the Peninsula of Baja California (which claims less than 9% of the butterfly fauna of Mexico), possess fewer total and endemic taxa. 5) Mexico’s complex biogeographical his- tory has resulted in several interconti- nental, insular, distributional patterns 217 that are evident among Mexican butter- flies; these are the product of a hetero- geneous and disjunct distribution of arid, humid and montane environments. 6) There are a number of relictual taxa that occur in unique biogeographical areas in Mexico. ACKNOWLEDGMENTS We thank Drs. Robert K. Robbins (Smithsonian Institution, Washington, D.C.) and Gerardo Lamas Miiller (Universidad Nacional Mayor de San Marcos, Lima, Peru) for their comments on and contribu- tions to this paper, as well as for access to the collections and libraries under their su- pervision. Additionally, we thank Paul A. Opler for comments on this paper. Thanks also to CONACyT for sponsoring project 32002, and to DGAPA-UNAM IN 209900 for funding. Special thanks is extended to the Comision Nacional para el Conocimien- to y Uso de la Biodiversidad (CONABIO) for funding research visits by three of the authors to institutional collections in the United States and to The Natural History Museum in London, England. LITERATURE CITED Balcazar, M. A. L. 1993. Butterflies of Pedernales, Mi- choacan, Mexico, with notes on seasonality and faunistic affinities (Lepidoptera: Papilionoidea and Hesperioidea). Tropical Lepidoptera 4(2): 93— 105. Brown, J. W., H. G. Real, and D. K. Faulkner. 1992. Butterflies of Baja California. Faunal Survey, Nat- ural History, Conservation Biology. The Lepidop- tera Research Foundation, Inc. Beverly Hills, Cal- ifornia, 129 pp. Burke, H. R. and P. A. Fryxell. 1995. Naturalists and their travels in Mexico: Annotated bibliography and roster of natural history collectors. Contribu- tions from the University of Michigan Herbarium 20: 37-128. Clench, H. K. 1968. Butterflies from Coahuila, Mexi- co. Journal of the Lepidopterists’ Society 22(4): 227-231. Comstock, J. A. 1959. Butterfly collecting in the Mex- ican tropics. Journal of the Lepidopterists’ Society 12(3—4): 127-129. de la Maza, J. and R. G. de la Maza. 1985a. La fauna de mariposas de Boca de Chajul, Chiapas, Mexico (Rhopalocera) Parte I. Revista de la Sociedad Mexicana de Lepidopterologia 9(2): 23-44. . 1985b. La fauna de mariposas de Boca de Chajul, Chiapas, México (Rhopalocera) Parte II. Revista de la Sociedad Mexicana de Lepidopter- ologia 10(1): 1-24. de la Maza, R. G. 1976. Notas sobre lepidopteros de Rancho Viejo y Tepoztlan, Morelos, México. Pri- mera parte: Papilionoidea. Revista de la Sociedad Mexicana de Lepidopterologia 1(2): 42-61. Diaz, A. F 1975. Papili6nidos del Valle de Tepoztlan, Morelos. Boletin Informativo de la Sociedad Mex- icana de Lepidopterologia 1(3): 5-7. Escalante, P., T. Peterson and A. Navarro. 1998. Un analisis geografico, ecologico e historico de la diversidad de aves terrestres de México, pp. 279— 304. In Ramammorthy, T., R. Bye, A. Lot, and J. Fa, eds. Diversidad Biol6gica de México: Orige- nes y distribuci6n. Instituto de Biologia, UNAM. Mexico City. Flores, O. 1998. Herpetofauna de México: distribucion y endemismo, pp. 251-278. Jn Ramammorthy, T., R. Bye, A. Lot, and J. Fa, eds. Diversidad Biol- 6gica de México: Origenes y distribucion. Insti- tuto de Biologia, UNAM, México City. Freeman, H. A. 1969. New species and records of Hes- periidae from Mexico (Lepidoptera). Journal of the Lepidopterists’ Society 23(suppl. 1): 1-59. Gonzalez, L. L. 1977. Reporte de la Colecta en la Cei- ba, Puebla. Boletin Informativo de la Sociedad Mexicana de Lepidopterologia 3(3): 6—7. 1978. Notas sobre la familia Papilionidae (Lepidoptera) en México. Barranca de Patla, Puebla y alrededores. Boletin Informativo de la Sociedad Mexicana de Lepidopterologia 4(3): 3— 15. Granillo, V. S. 1985. Uso y abuso de la selva. Infor- macion Cientifica y Tecnologica 7: 35-38. Guzman, E. P. 1975. Algunos hespéridos del Valle de México. Boletin Informativo de la Sociedad Mex- icana de Lepidopterologia 1(5): 6—8. Halffter, G. 1976. Distribucion de los insectos en la zona de Transicion Mexicana. Relaciones con la entomofauna de Norteamérica. Folia Entomolo- gica Mexicana 35: 1-64. . 1987. Biogeography of the montane entomo- fauna of Mexico and Central America. Annual Review of Entomology. 32: 95-114. Heppner, J. B. 1991. Faunal regions and the diversity of Lepidoptera. Tropical Lepidoptera 2(suppl. 1): 1-85. Llorente, J. B. 1984. Sinopsis sistematica y biogeo- grafica de los Dismorphiinae de México, con es- pecial referencia al género Enantia Huebner (Lep- idoptera: Pieridae). Folia Entomologica Mexicana 58: 1-207. . 1996. Biogeografia de artr6podos de México. {Hacia un nuevo enfoque?, pp. 41—56. Jn Lloren- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON te, J. B., A. N. A. Garcia, and E. S. Gonzalez, eds. Biodiversidad, Taxonomia y Biogeografia de Ar- tropodos de México: Hacia una Sintesis de su Conocimiento. UNAM, Mexico City. Llorente, J. B. and P. Escalante. 1992. Insular Bioge- ography of Submontane Humid Forest in Mexico, pp. 139-148. Jn Darwin, S. P. and A. L. Welden, eds. Biogeography of Mesoamerica. E. O. Painter Printing Company, Florida. Llorente, J. B. and A. M. Luis. 1993. A conservation- oriented analysis of Mexican butterflies: the Pap- ilionidae (Lepidoptera: Papilionoidea), pp. 147— 177. In Ramammorthy, T., R. Bye, A. Lot, and J. Fa, eds. The Biological Diversity of Mexico: Or- igins and Distributions. Oxford University Press. Llorente, J. B., A. M. Luis, I. EF Vargas, and J. Sob- eron. 1994. Biodiversidad de las mariposas: su conocimiento y conservacion en México. Revista de la Sociedad Mexicana de Historia Natural 44: 313-324. . 1996. Papilionoidea (Lepidoptera), pp. 531— 548. In Llorente, J. B., A. N. A. Garcia, and E. S. Gonzalez, eds. Biodiversidad, Taxonomia y Bio- geografia de Artropodos de México: Hacia una Sintesis de su Conocimiento. UNAM, Mexico City. Llorente, J. B., L. Onate, A. M. Luis, and I. F Vargas. 1997. Papilionidae y Pieridae de México: Distri- bucion Geografica e Ilustracion. UNAM, México City. Luis, A. M. and J. B. Llorente. 1990. Mariposas en el Valle de México: Introduccion e historia. 1. Dis- tribucion local y estacional de los Papilionoidea de la Canada de los Dinamos, Magdalena Con- treras, D. FE, México. Folia Entomologica Mexi- cana 78: 95-198. Luis, A. M., J. B. Llorente, I. EF Vargas, and A. L. Gutiérrez. 2000. Sintesis preliminar del conoci- miento de los Papilionoidea (Lepidoptera: Insecta) de México, pp. 275-285. Jn Piera, F, J. J. Mor- rone, and A. Melic, eds. Monografias Tercer Mil- enio, Bol. S.E.A., Vol. I. Zaragoza, Espana. Luis, A. M., I. E Vargas, and J. B. Llorente. 1991. Lepidopterofauna de Oaxaca: I. Distribucion y fenologia de los Papilionoidea de la Sierra de Jua- rez. Publicaciones Especiales del Museo de Zool- ogia, Facultad de Ciencias, UNAM 3: 1-121. . 1996. Sintesis de los Papilionoidea (Rhopal- ocera: Lepidoptera) del estado de Veracruz. Folia Entomologica Mexicana 93: 91-133. Monteagudo, D., A. M. Luis, I. FE Vargas, and J. B. Llorente. 2001. Patrones altitudinales de la div- ersidad de mariposas en la Sierra Madre del Sur (México). (Lepidoptera: Papilionoidea). Shilap Revista Lepidopterologica 28(115): 203-233. Morrone, J. J., D. Espinosa, C. Aguilar, and J. B. Llo- rente, 1999. Preliminary classification of the Mex- ican biogeographic provinces: A parsimony anal- VOLUME 105, NUMBER | ysis of endemicity based on plant, insect and bird taxa. Southwestern Naturalist 44(4): 508-515. Navarro, A. and J. B. Llorente. 1996. Museos, colec- ciones biolégicas y la conservacion de la biodiv- ersidad: Una perspectiva para México. Ciéncias em Museus 3: 27-49. Onate, L., J. J. Morrone, and J. B. Llorente. 2000. Evaluacion del conocimiento y de la distribucion de las Papilionidae y Pieridae mexicanas (Lepi- doptera: Insecta). Acta Zoologica Mexicana (n.s.) 81: 117-132. Raguso, R. A. and J. B. Llorente. 1997. Papilionoidea, pp. 257-291. In Gonzalez, E., R. Dirzo, and R. Vogt, eds. Historia Natural de los Tuxtlas. Insti- tuto de Biologia, UNAM, Mexico City. Robbins, R. K. and P. A. Opler. 1997. Butterfly diver- sity and a preliminary comparison with bird and mammal diversity, pp. 69-82. Jn Wilson, D. E., M. L. Reaka-Kudla, and E. O. Wilson, eds. Bio- diversity II, Understanding and Protecting Our Bi- ological Resources. Joseph Henry Press, Washing- ton, D.C. Routeledge, C. E. 1977. El suborden Rhopalocera (Lepidoptera) del estado de Tabasco. Su lista, fre- cuencia, diversidad y distribucion. Revista de la Sociedad Mexicana de Lepidopterologia 3(2): 57— 73: Rzedowski, J. 1978. La Vegetacion de México. Li- musa, Mexico City. Salinas, J. L. G. 1999. Los Papilionoideos (Lepidop- tera: Rhopalocera) de los Bosques Tropicales Per- ennifolios de México. Tesis de Licenciatura. Fa- cultad de Ciencias, UNAM, México City, 120 pp. Shields, O. 1989. World numbers of butterflies. Journal of the Lepidopterist’s Society 43(3): 178-183. Soberon, J., J. B. Llorente and L. Onate. 2000. The use of large specimens-labels data base for con- servation purposes: an example using Mexican papilionid and pierid butterflies. Biodiversity and Conservation 9(10): 1441-1466. Toledo, V. M. 1988. La diversidad biologica de Meéx- ico. Ciencia y Desarrollo 81: 17—30. Vargas, I. FE, J. B. Llorente, and A. M. Luis. 1994. Listado lepidopterofaunistico de la sierra de Atoy- ac de Alvarez en el estado de Guerrero: notas ac- erca de su distribucion local y estacional (Rho- palocera: Papilionoidea). Folia Entomologica Mexicana 86: 41-178. ——.. 1999. Distribucion de los Papilionoidea (Lep- idoptera: Rhopalocera) de la sierra de Manantlan (250-1650 m) en los estados de Jalisco y Colima. Publicaciones Especiales del Museo de Zoologia, Facultad de Ciencias, UNAM 11: 1-153. Vargas, I. F, A. M. Luis, J. B. Llorente, and A. D. Warren. 1996. Butterflies of the state of Jalisco, Mexico. Journal of the Lepidopterist’s Society 50(2): 97-138. Warren, A. D. and J. B. Llorente. 1999. Butterflies of 219 Mismaloya, Jalisco, and adjacent parts of Bander- as Bay and southern Nayarit, Mexico. Dugesiana 6(1): 17-32. Warren, A. D., I. E Vargas, A. M. Luis, and J. B. Llorente. 1998. Butterflies of the state of Colima, Mexico. Journal of the Lepidopterist’s Society 52(1): 40-72. Weintraub, J. D. and J. S. Miller. 1987. Review. The structure and affinities of the Hedyloidea: a new concept of the butterflies. Cladistics 3(3): 299— 304. APPENDIX Species and Subspecies of Endemic Mexican Butterflies = Taxon that has been reported from the USA by stray specimens or in error. “ = Taxon reported from Belize or Guatemala by rare strays. “= Taxon originally described from Arizona but now extinct in the USA. = Questionable taxonomic status, further study re- quired (this symbol is after names). Taxa listed in bold are endemic to Mexico at the species level. We note that some taxa listed as endemic herein may be found to have distributions that extend outside of Mexico with future fieldwork. However, tax- onomic revisions and new species discoveries also will increase the number of endemic species in the future. Endemic genera include: Baronia (Papilionidae), Prestonia, Eucheira (Pieridae), Zobera, Aegiale, and Turnerina (Hesperiidae). We list subspecies (geographical races) to aid in the identification of patterns of endemism. Distributions of many Theclinae and Hesperiidae are less well known, and most modern publications on these groups have not recognized subspecies extensively. PAPILIONIDAE Baroniinae Baronia brevicornis brevicornis Salvin, 1893 Baronia brevicornis rufodiscalis J. Maza & J. White, 1987 Papilioninae Battus philenor orsua (Godman & Salvin, 1889) “Battus philenor acauda (Oberthiir, 1880) Battus laodamas topas (Godman & Salvin, 1897) Battus eracon (Godman & Salvin, 1897) *Parides alopius (Godman & Salvin, 1890) Parides erithalion trichopus (Rothschild & Jordan, 1906) Parides panares panares (Gray, [1853]) Protographium agesilaus fortis (Rothschild & Jordan, 1906) Protographium epidaus tepicus (Rothschild & Jordan, 1906) 220 Protographium epidaus fenochionis (Salvin & God- man, 1868) Protographium thyastes occidentalis (R. G. Maza, 1982) Mimoides ilus occiduus (Vazquez, 1956) Mimoides thymbraeus aconophos (Gray, [1853]) Heraclides erostratus erostratinus (Vazquez, 1947) Heraclides erostratus vazquezae (Beutelspacher, 1986) “Heraclides rogeri rogeri (Boisduval, 1836) Heraclides torquatus mazai (Beutelspacher, 1974) Heraclides astyalus bajaensis (J. W. Brown & Faulk- ner, 1992) Pterourus esperanza (Beutelspacher, 1975) Pterourus glaucus alexiares (Hopffer, 1865) Pterourus glaucus garcia (Rothschild & Jordan, 1906) Pterourus palamedes leontis (Rothschild & Jordan, 1906) *Pterourus abderus abderus (Hoptter, 1856) Prerourus abderus baroni (Rothschild & Jordan, 1906) Pterourus garamas (Geyer, {1829]) Pterourus menatius morelius (Rothschild & Jordan, 1906) PIERIDAE Dismorphiinae Enantia mazai mazai L\orente, 1984 Enantia mazai diazi Llorente, 1984 Lieinix lala turrenti J. Maza & R. G. Maza, 1984 Lieinix neblina J. Maza & R. G. Maza, 1984 Lieinix nemesis nayaritensis Llorente, 1984 Dismorphia amphione lupita Lamas, 1979 Dismorphia amphione isolda Llorente, 1984 Dismorphia crisia alvarezi J. Maza & R. G. Maza, 1984 Dismorphia eunoe eunoe (Doubleday, 1844) Dismorphia eunoe popoluca Llorente & Luis, 1988 Dismorphia eunoe chamula Llorente & Luis, 1988 Coliadinae *Phoebis agarithe fisheri (H. Edwards, 1883) Prestonia clarki Schaus, 1920 Eurema agave millerorum Llorente & Luis, 1987 Pierinae Anthocharis cethura bajacalifornia J. Emmel, T. Em- mel & Mattoon, 1998 Anthocharis limonea (Butler, 1871) *Euchloe guaymasensis Opler, 1987 Hesperocharis crocea jaliscana (Schaus, 1898) Hesperocharis graphites avivolans (Butler, 1865) Eucheira socialis socialis Westwood, 1834 Eucheira socialis westwoodi Beutelspacher, 1984 Catasticta flisa oaxaca Beutelspacher, 1986 Catasticta undescribed species *Catasticta nimbice nimbice (Boisduval, 1836) Catasticta teutila flavifaciata Beutelspacher, 1986 Catasticta teutila teutila (Doubleday, 1847) Pereute charops charops (Boisduval, 1836) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pereute charops leonilae Llorente, 1986 Pereute charops sphocra Draudt, 1931 Melete polyhymnia serrana R. G. Maza, 1984 Perrhybris pamela chajulensis J. Maza & R. G. Maza, 1989 Perrhybris pamela mapa J. Maza & R. G. Maza, 1989 Ascia monuste raza Klots, 1930 *Ganyra howarthi (Dixey, 1915) NYMPHALIDAE Heliconiinae Actinote stratonice oaxaca (L. Miller & J. Miller, 1979) Actinote guatemalena guerrerensis J. Maza, 1982 Actinote guatemalena veraecrucis Jordan, 1913 Eueides isabella nigricornis R. G. Maza, 1989 Heliconius erato cruentus Lamas, 1998 Speyeria nokomis wenona dos Passos & Grey, 1945 « Speyeria nokomis melaena Mooser & Garcia, 1979 « “Speyeria nokomis coerulescens (Holland, 1900) Nymphalinae Nymphalis cyanomelas (Doubleday, {1848]) Polygonia g-argenteum (Doubleday, 1848) Polygonia haroldii (Dewitz, 1877) Anartia amathea colima Lamas, 1995 Chlosyne kendallorum Opler, 1999 *Chlosyne ehrenbergii (Geyer, [1833]) Chlosyne definita anastasia (Hemming, 1934) Chlosyne definita schaust (Godman, 1901) Chlosyne endeis endeis (Godman & Salvin, 1894) Chlosyne gaudialis wellingi L. Miller & Rotger, 1979 *Chlosyne eumeda (Godman & Salvin, 1894) Chlosyne marina (Geyer, 1837) *Chlosyne melitaeoides (R. Felder 1867) Chlosyne janais gloriosa Bauer, 1960 Chlosyne janais marianna Rober, [1914] Chlosyne rosita mazarum L. Miller & Rotger, 1979 Chlosyne rosita riobalsensis Bauer, 1961 Thessalia cyneas cynisca (Godman & Salvin, 1882) Thessalia leanira austrima Austin & M. J. Smith, 1998 Thessalia theona brocki Austin & M. J. Smith, 1998 Thessalia theona mullinsi Austin & M. J. Smith, 1998 Texola anomalus (Godman & Salvin, 1897) Texola coracara (Dyar, 1912) Texola elada elada (Hewitson, 1868) Phyciodes pallescens (R. Felder, 1869) Phyciodes mylitta mexicanus Hall, 1928 Anthanassa alexon (Godman & Salvin, 1889) Anthanassa ptolyca amator (Hall, 1929) Anthanassa sitalces cortes (Hall, 1917) « Anthanassa otanes cyno (Godman & Salvin, 1889) Anthanassa otanes oaxaca Beutelspacher, 1990 Castilia chinantlensis (R. R. Maza, 1978) Limenitidinae Myscelia cyananthe diaziana R. G. Maza & J. Maza, 1985 VOLUME 105, NUMBER | *Myscelia cyananthe skinneri Mengel, 1894 Myscelia cyananthe streckeri Skinner, 1889 Myscelia cyaniris alvaradia R. G. Maza & Diaz, 1982 Catonephele cortesi R. G. Maza, 1982 Eunica malvina almae Vargas, Llorente & Luis, 1998 Hamadryas amphinome mazai Jenkins, 1983 *Hamadryas atlantis lelaps Godman & Salvin, 1883 *Hamadryas glauconome grisea Jenkins, 1983 *Hamadryas guatemalena marmarice (Frihstorter, 1916) \Hamadryas honorina (Friihstorfer, 1916) Pyrrhogyra edocla paradisea R. G. Maza & J. Maza, 1985 Temenis laothoe quilapayunia R. G. Maza & Turrent, 1985 Epiphile adrasta escalantei Descimon & Mast, 1979 Bolboneura sylphis beatrix R. G. Maza, 1985 Bolboneura sylphis lacandona R. G. Maza & J Maza, 1985 Bolboneura sylphis veracruzana Draudt, 1931 Diaethria astala asteroidea R. G. Maza & R. R. Maza, 1985 Diaethria asteria (Godman & Salvin, 1894) Diaethria salvadorensis mixteca J. Maza, 1977 Callicore astarte casta (Salvin, 1869) Callicore texa grijalva R. G. Maza & J. Maza, 1983 Callicore texa loxicha R. G. Maza & J. Maza, 1983 Callicore tolima tehuana R. G. Maza & J. Maza, 1983 Adelpha milleri Beutelspacher, 1976 Adelpha boeotia oberthurti (Boisduval, 1870) Adelpha diazi Beutelspacher, 1975 Adelpha diocles creton Godman, 1901 Adelpha leucerioides leucerioides Beutelspacher, 1975 Basilarchia archippus hoffmanni (Chermock, 1947) Charaxinae Archaeoprepona amphimachus baroni J. Maza, 1982 Archaeoprepona demophon occidentalis Stoftel & Descimon, 1974 Archaeoprepona demophoon mexicana Llorente, Des- cimon & Johnson, 1993 Archaeoprepona phaedra aelia (Godman & Salvin, 1889) Prepona deiphile brooksiana Godman & Salvin, 1889 Prepona deiphile diaziana L. Miller & J. Miller, 1976 Prepona deiphile escalantiana Stotfel & Mast, 1973 Prepona deiphile ibarra Beutelspacher, 1982 Prepona deiphile lambertoana Llorente, Luis & Gon- zAlez, 1992 Hypna clytemnestra mexicana Hall, 1917 Consul electra undescribed subspecies Fountainea eurypyle glanzi (Rotger, Escalante & Cor- onado, 1965) Fountainea halice martinezi (J. Maza & Diaz, 1978) Fountainea halice maya (Witt, 1980) Fountainea halice tehuana (Hall, 1917) Fountainea nobilis rayoensis (J. Maza & Diaz, 1978) Memphis schausiana (Godman & Salvin, 1894) Memphis wellingi L. Miller & J. Miller, 1976 Morphiinae Morpho achilles guerrerensis Le Moult & Réal, 1962 Morpho achilles montezuma Guenée, 1859 Iphimedeia telemachus oaxacensis Le Moult & Réal, 1962 Satyrinae Cissia cleophes (Godman & Salvin, 1889) « Cyllopsis caballeroi Beutelspacher, 1982 Cyllopsis clinas (Godman & Salvin, 1889) Cyllopsis diazi L. Miller, 1974 Cyllopsis dospassosi L. Miller, 1969 Cyllopsis hedemanni tamaulipensis L. Miller, 1974 Cyllopsis henshawi hoffmanni L. Miller, 1974 « Cyllopsis jacquelinae . Miller, 1974 Cyllopsis nayarit Chermock, 1947 « Cyllopsis parvimaculata L. Miller, 1974 Cyllopsis perplexa L. Miller, 1974 Cyllopsis pertepida pertepida (Dyar, 1912) Cyllopsis pertepida intermedia L. Miller, 1974 Cyllopsis pseudopephredo Chermock, 1947 Cyllopsis suivalens suivalens (Dyar, 1914) Cyllopsis whiteorum L. Miller & J. Maza, 1984 Cyllopsis windi L. Miller, 1974 « Euptychia fetna Butler, 1870 Euptychia rubrofasciata L. Miller & J. Miller, 1988 “Megisto”’ pellonia (Godman, 1901) Megisto rubricata pseudocleophes L. Miller, 1976 Paramacera chinanteca L. Miller, 1972 Paramacera copiosa L. Miller, 1972 Paramacera xicaque rubrosuffusa L. Miller, 1972 Paramacera xicaque xicaque (Reakirt, |1867]) Splendeuptychia kendalli L. Miller, 1976 Taygetis mermeria griseomarginata L. Miller 1978 Taygetis uncinata Weymer, 1907 Taygetis weymeri Draudt, 1912 Pedaliodes undescribed species Ithomiinae Melinaea ethra flavicans Hoffmann, 1924 Oleria zea diazi J. Maza & Lamas, 1978 Callithomia hezia wellingi Fox, 1968 Episcada salvinia portilla J. Maza & Lamas, 1978 Preronymia artena praedicta J. Maza & Lamas, 1982 Pteronymia rufocincta (Salvin, 1869) Preronymia simplex timagenes Godman & Salvin, 1889 Greta annette moschion (Godman, 1901) LYCAENIDAE Theclinae Habrodais poodiae J. W. Brown & Faulkner, 1982 Eumaeinae Micandra tongida Clench, 1971 Laothus erybathis (Hewitson, 1867) Callophrys dospassosi Clench, 1981 Callophrys estela Clench, 1981 222 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Symbiopsis undescribed species Ministrymon undescribed species Erora undescribed species Thecla semones (Godman & Salvin, 1887) Ipidecla miadora Dyar, 1916 Polyommatinae Everes undescribed species Euphilotes bernardino garthi Mattoni, 1989 Lycaeides melissa mexicana (Clench, 1965) Riodininae Euselasia cataleuca (R. Felder, 1869) Euselasia hypophaea mexicana Lathy, 1926 Euselasia undescribed species Mesosemia gemina J. Maza & R. G. Maza, 1980 Napaea danforthi A. Warren & Opler, 1999 Rhetus arcius beutelspacheri Llorente, 1987 Calephelis acapulcoensis McAlpine, 1971 « Calephelis azteca McAlpine, 1971 « *“Calephelis dreisbachi McAlpine, 1971 « Calephelis huasteca McA\pine, 1971 « Calephelis nemesis bajaensis McAlpine, 1971 « Calephelis matheri McAlpine, 1971 « Calephelis mexicana McAlpine, 1971 « Calephelis montezuma McAlpine, 1971 « Calephelis perditalis donahuei McAlpine, 1971 « Calephelis sinaloensis sinaloensis McAlpine, 1971 « Calephelis sinaloensis nuevoleon McA\pine, 1971 « Calephelis yautepequensis R. G. Maza & Turrent, 1971 Caria melino Dyar, 1912 Caria stillaticia Dyar, 1912 Exoplisia undescribed species Melanis cephise huasteca J. White & A. White, 1989 Melanis cephise acroleuca (R. Felder, 1869) Emesis poeas Godman & Salvin, 1901 Emesis zela zela Butler, 1870 Apodemia hepburni remota Austin, 1991 Apodemia mormo dialeuca Opler & Powell, 1962 Apodemia mormo maxima A. G. Weeks, 1891 Apodemia murphyi Austin, 1988 Apodemia palmerii australis Austin, 1988 *Apodemia phyciodoides Barnes & Benjamin, 1924 Apodemia hypoglauca wellingi Ferris, 1985 Adelotypa eudocia (Godman & Salvin, 1897) Theope villai Beutelspacher, 1981 HESPERIDAE Pyrrhopyginae Chalypyge chalybea chalybea (Scudder, 1872) « Jonaspyge tzotzili H. A. Freeman, 1969 Melanopyge mulleri (Bell, 1934) Apyrrothrix araxes araxes (Hewitson, 1867) Mysoria wilsoni H. A. Freeman, 1969 Pyrginae Entheus crux Steinhauser, 1989 Epargyreus brodkorbi H. A. Freeman, 1966 (must also occur in GUAT) Epargyreus deleoni H. A. Freeman, 1977 Typhedanus salas H. A. Freeman, 1977 Typhedanus undescribed species Zestusa elwesi (Godman & Salvin, 1893) Zestusa undescribed species Codatractus cyledis (Dyar, 1912) Codatractus uvydixa (Dyar, 1914) Codatractus yucatanus H. A. Freeman, 1977 Ridens mercedes Steinhauser, 1983 Urbanus undescribed species #1 Urbanus dorantes calafia (R. C. Williams, 1926) Urbanus undescribed species #2 Urbanus viridis H. A. Freeman, 1970 “Autochton pseudocellus (Coolidge & Clemence, [1910]) Autochton siermadror Burns, 1984 Thorybes undescribed species Cephise mexicanus Austin & Mielke, 2000 Cogia aventinus (Godman & Salvin, 1894) Cogia hippalus peninsularis L. Miller & MacNeill, 1969 Telemiades choricus (Schaus, 1902) Mimia chiapaensis H. A. Freeman, 1969 Polyctor undescribed species Myrinia raymundo H. A. Freeman, 1979 Bolla cybele Evans, 1953 Bolla fenestra Steinhauser, 1991 Bolla guerra Evans, 1953 Bolla litus (Dyar, 1912) Bolla oriza Evans, 1953 Bolla solitaria Steinhauser, 1991 Staphylus undescribed species Staphylus tepeca (Bell, 1942) Staphylus tierra Evans, 1953 Zera eboneus (Bell, 1947) Quadrus francesius H. A. Freeman, 1969 Pythonides mundo H. A. Freeman, 1979 Pythonides rosa Steinhauser, 1989 Zobera albopunctata H. A. Freeman, 1970 Zobera marginata H. A. Freeman, 1979 Zobera oaxaquena Steinhauser, 199] *Antigonus emorsa (R. Felder, 1869) Antigonus funebris (R. Felder, 1869) Systasea microsticta Dyar, 1923 Doberes hewitsonius (Reakirt, [1867]) Doberes sobrinus Godman & Salvin, 1895 Anastrus luctuosus Godman & Salvin, 1894 Camptopleura oaxaca H A. Freeman, 1969 Chiomara georgina pelagica (Weeks, 1891) Erynnis brizo mulleri (Draudt, 1923) Erynnis mercurius (Dyar, 1926) Erynnis meridianus fieldi Burns, 1964 Erynnis tristis pattersoni Burns, 1964 Heliopyrgus sublinea Schaus, 1902 VOLUME 105, NUMBER | Celotes undescribed species Heteropterinae Piruna ceracates (Hewitson, 1874) Piruna cyclosticta (Dyar, 1920) Piruna dampfi (Bell, 1942) Piruna gyrans (Plotz, 1884) Piruna jonka Stainhauser, 1991 Piruna kemneri H. A. Freeman, 1990 Piruna maculata H. A. Freeman, 1970 Piruna microsticta (Godman, 1900) Piruna millerorum Steinhauser, 1991 Piruna mullinsi H. A. Freeman, 1991 Piruna purepecha A. Warren & Gonzalez, 1998 Piruna roeveri (L. Miller & J. Miller, 1972) Piruna undescribed species #1 Piruna sina H. A. Freeman, 1970 Piruna undescribed species #2 Dalla bubobon (Dyar, 1921) Dalla dividuum (Dyar, 1913) Dalla faula (Godman, 1900) Dalla kemneri Steinhauser, 1991 Dalla undescribed species Dalla mentor Evans, 1955 Dalla nubes Steinhauser, 1991 Dalla steinhauseri H. A. Freeman, 1991 Hesperiinae Synapte silna Evans, 1955 *Synapte syraces (Godman, 1901) Zariaspes mythecus Godman, 1900 Anthoptus macalpinei H. A. Freeman, 1969 Pheraeus covadonga covadonga H. A. Freeman, 1969 Pheraeus covadonga loxicha Steinhauser, 1991 Virga clenchi L. Miller, 1970 Monca jera Godman, 1900 Remella undescribed species Cobalopsis zetus (Bell, 1942) Turesis tabascoensis H. A. Freeman, 1979 Thoon wellingi H. A. Freeman, 1969 Enosis matheri H. A. Freeman, 1969 Ebusus ebusus nigrior L. Miller, 1985 Carystoides escalantei H. A. Freeman, 1969 Carystoides abrahami H. A. Freeman. 1969 Carystoides floresi H. A. Freeman, 1969 Carystoides mexicana H. A. Freeman, 1969 Cynea nigricola H. A. Freeman, 1969 Decinea rindgei H. A. Freeman, 1968 Decinea mustea H. A. Freeman, 1979 Oceonus pyste Godman, 1900 Oarisma era Dyar, 1927 Adopaeodes bistriata Godman, 1900 Stinga undescribed species Hesperia uncas gilberti MacNeill, 1964 Polites norae MacNeill, 1993 Polites pupillus (Plotz, 1883) Polites puxillius (Mabille, 1891) Polites sabuleti margaretae L. Miller & MacNeill, 1969 223 Ochlodes samenta Dyar, 1914 Poanes monticola (Godman, 1900) “Poanes”’ benito H. A. Freeman, 1979 Paratrytone aphractoia Dyar, 1914 Paratrytone decepta L. Miller & J. Miller, 1972 Paratrytone kemneri Steinhauser, 1996 Paratrytone undescribed species #1 Paratrytone raspa (Evans, 1955) Paratrytone undescribed species #2 Paratrytone omiltemensis Steinhauser, 1996 Paratrytone undescribed species #3 Paratrytone pilza Evans, 1955 Paratrytone polyclea Godman, 1900 Paratrytone rhexenor Godman, 1900 Paratrytone undescribed species #4 Quasimellana siblinga Burns, 1994 Quasimellana agnesae (Bell, 1959) Quasimellana mulleri (Bell, 1942) “Mellana”’ gala (Godman, 1900) Librita heras (Godman, 1900) Euphyes chamuli H. A. Freeman, 1969 (must also oc- cur in GUAT) Euphyes canda Steinhauser & Warren, [2002] Atrytonopsis frappenda (Dyar, 1920) Atrytonopsis zweifeli H. A. Freeman, 1969 Atrytonopsis undescribed species Amblyscirtes anubis (Godman, 1900) Amblyscirtes brocki H. A. Freeman, 1992 Amblyscirtes fimbriata pallida H. A. Freeman, 1993 Amblyscirtes fluonia Godman, 1900 Amblyscirtes folia Godman, 1900 Amblyscirtes novimmaculatus A. Warren, 1998 Amblyscirtes raphaeli H. A. Freeman, 1973 Lerodea similea Bell, 1942 « Vacerra undescribed species #1 Vacerra gayra (Dyar, 1918) Vacerra undescribed species #2 Vacerra undescribed species #3 Niconiades comitana H. A. Freeman, 1969 Halotus jonaveriorum Burns, 1992 Aides undescribed species Megathyminae Stallingsia smithi (H. H. Druce. 1896) Stallingsia undescribed species Stallingsia jacki D. Stallings, Turner, & V. Stallings, 1963 Megathymus beulahae beulahae D. Stallings & Turn- er, 1958 Megathymus beulahae gayleae D. Stallings, Turner & V. Stallings, 1963 Aegiale hesperiaris (Walker, 1856) (may represent a species complex) Turnerina mejicanus (Bell, 1938) Turnerina hazelae (D. Stallings & Turner, 1958) Agathymus belli (H. A. Freeman, 1955) Agathymus comstocki (Harbison, 1957) Agathymus dawsoni Harbison, 1963 Agathymus undescribed species #1 224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Agathymus undescribed species #2 Agathymus undescribed species #7 Agathymus undescribed species #3 Agathymus undescribed species #8 Agathymus undescribed species #4 Agathymus micheneri D. Stallings, Turner & V. Stall- Agathymus escalantei D. Stallings. Turner & V. Stall- ings, 1961 ' ince lOG6 Agathymus undescribed species #9 gs, 14 Agathymus undescribed species #10 Agathymus undescribed species #11 Agathymus undescribed species #12 Agathymus undescribed species #13 Agathymus rethon (Dyar, 1913) Agathymus undescribed species #5 Agathymus ricei D. Stallings, Turner & V. Stallings, Agathymus undescribed species #6 1966 Agathymus remingtoni (D. Stallings & Turner, 1958) Agathymus fieldi H. A. Freeman, 1960 Agathymus hoffmanni (H. A. Freeman, 1952) Agathymus juliae (D. Stallings & Turner, 1958) PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 225-237 HALYOMORPHA HALYS (STAL) (HETEROPTERA: PENTATOMIDAE): A POLYPHAGOUS PLANT PEST FROM ASIA NEWLY DETECTED IN NORTH AMERICA E. RICHARD HOEBEKE AND MAUREEN E. CARTER Department of Entomology, Cornell University, Ithaca, NY 14853, U.S.A. (e-mail: erh2 @cornell.edu; mec8 @cornell.edu) Abstract.—The Asian pentatomid Halyomorpha halys (Stal) (Heteroptera) is reported for the first time in North America based on collection of specimens or confirmed sightings in five counties in eastern Pennsylvania. Known in the Asian literature as the yellow- brown or brown marmorated stink bug, H. halys is considered a significant horticultural pest in Japan; it has a broad host range that includes plants of economic importance such as pea, soybean, apple, peach, cherry, and various other fruit and ornamental trees. In Japan’s northern regions it is regarded as a nuisance pest late in the season when adults aggregate and seek overwintering sites in and around houses and commercial establish- ments. Descriptions and photographs of the adult and immature stages are provided to help distinguish it from other North American pentatomids. Habits, seasonal history, and host plants, based on the Asian literature and our own observations, are summarized. Photographs of leaf-feeding damage are also provided. Halyomorpha halys may have been accidentally introduced with international commerce, i.e., via bulk freight containers from either Japan, Korea, or China. Key Words: species, Pennsylvania In late September of 2001, two speci- mens of a pentatomid stink bug, collected in Allentown, Pennsylvania (Lehigh Coun- ty), were submitted to the Cornell Univer- sity Insect Diagnostic Lab by Karen M. Bernhard (KMB), an extension entomolo- gist with the Lehigh County Cooperative Extension office. They proved to be Haly- omorpha halys (Stal), an Asian species not known previously to occur in North Amer- ica. Numerous log records of homeowners’ inquiries of stink bugs being found in and around homes in the greater Allentown re- gion over the past several years (compiled by KMB) indicate that populations of this exotic stink bug are well established in Le- high County and to a lesser degree in sur- Heteroptera, Pentatomidae, Halyomorpha halys, North America, adventive rounding counties in eastern Pennsylvania. Further surveys are needed to determine the full geographic range of this introduced plant pest in the eastern United States. The earliest confirmed U.S. sighting of H. halys is from Adams Island (Allentown) in the fall of 1996. We are also aware of stink bug sightings in Allentown in Septem- ber of 1998 and January of 1999. Since these initial sightings, homeowners of the region have made numerous telephone calls complaining of this bug to the extension of- fice of Lehigh County. After ERH identified the two specimens on October |, 2001, he joined KMB on October 3 in Allentown to make further observations and to collect ad- ditional specimens. On that unseasonably 226 warm day, adults of H. halys were extreme- ly numerous on the foundations, outer wall surfaces, eaves, and window and door frames of homes, and sidewalks. In this paper, we provide the first North American records for H. halys; review the literature on seasonal history, habits, eco- nomic damage, and host plants; provide notes On our own rearing data; and rede- scribe and give color photographs of the adult and immature stages that allow it to be distinguished from similar-appearing North American pentatomids. SYSTEMATICS Halyomorpha halys (Stal) (Figs. 1-7) Pentatoma Halys Stal 1855: 182. Poecilometis mistus Uhler 1860: 223. Syn- onymized by Josifov and Kerzhner 1978: ID Dalpada brevis Walker 1867: 226. Synon- ymized by Josifoy and Kerzhner 1978: 72: Dalpada remota Walker 1867: 227. Syn- onymized by Josifov and Kerzhner 1978: N72: Halyomorpha picus: auctorum (nec Fabri- cius 1794: 115). Halyomorpha halys: Josifov and Kerzhner 1978: 172. Note.—Josifov and Kerzhner (1978) de- termined that only one species of Haly- omorpha is found in Japan, Korea, and east- ern China. All citations pertaining to Hal- yomorpha species from these localities should be referred to H. halys (Rider et al. in press). There are currently 37 species of Halyomorpha recognized, of which 16 are African, 8 are Indian, and 13 are Asian. There is, however, considerable confusion surrounding several species; a generic re- vision is needed. Diagnosis.—The only eastern North America pentatomids that resemble Haly- omorpha halys in overall size (12-17 mm) and its dark “marbled” dorsal coloration are the Brochymena species of genus PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Amyot and Serville (tribe Halyini). Mem- bers of Brochymena, however, have the juga each with a tooth on the outer side subapically and the pronotum with the an- terolateral margins coarsely dentate. By comparison, H. halys lacks teeth on the out- er juga subapically and the anterolateral margins of the pronotum are not dentate, but entire. Among members of the tribe Pentatomini sensu Rolston and McDonald (1979) (to which H. halys belongs; many Old World workers place Halyomorpha in the tribe Cappaeini which at present has no known representatives in the New World), only the taxa Euschistus Dallas, Holcostethus Fieber, and Thyanta Stal might be confused with Halyomorpha. Euschistus species are rec- ognized by the conspicuous ostiole (of the metathoracic scent gland) usually with a distinct, well-developed auricle (similar to a small earlike or flaplike process or lobe). Also in Euschistus, the hind tibiae are dis- tinctly sulcate dorsally throughout their lengths. In Halyomorpha, the metathoracic scent gland ostiole is without a developed auricle but has an elongate tapering channel extended beyond the middle of the sup- porting or evaporative plate, and the hind tibiae are without a dorsal sulcus. Halyomorpha keys to Thyanta in Mc- Pherson (1982). Thyanta species, however, are generally smaller (7-11 mm) and the dorsal surface is chiefly or entirely green or pale tan (overwintering forms). In Slater and Baranowski (1978), Halyomorpha will trace to the genus Holcostethus. In Holco- stethus, the metathoracic scent gland is pro- duced into an elongate tapering channel as in Halyomorpha. In contrast, Holcostethus is smaller (7-9 mm); has the apices of the juga rounded, converging, and frequently contiguous in front of the tylus; and differs in general shape and coloration. Also in Halyomorpha, the connexiva are usually broadly exposed (hemelytra not covering them), revealing the distinctive alternating pattern of black and pale patches. The fourth antennal segment is annulated (pale VOLUME 105, NUMBER 1 white) at the base and apex, and the fifth segment is annulated at the base. The venter of the adult (thoracic sterna, abdomen, and leg bases) is distinctly pale luteous. Redescription.—Adu/t (Fig. 1): Length 12—17 mm.; width across humeral angles 7— 10 mm. Variable in color and size. Gener- ally brownish cinereous, but also greyish ochraceous, ochraceous, testaceous, or cas- taneous, densely and darkly punctate. Head broadly rounded in front, surface densely punctured with black, lateral margins nar- rowly reflexed and slightly sinuate, tylus and juga nearly equal in length (tylus per- haps slightly longer); eye large, sessile; ros- trum reaching second abdominal segment. Head, anterior and lateral areas of pronotum more or less heavily punctured with black, coarse punctures; basal angles of scutellum pale luteous or yellow, without punctures. Hemelytra mottled with brown, densely punctured, slightly tinged with reddish on apex of corium, apical membrane yellowish hyaline, with veins marked with dark brown. Body beneath and legs generally pale luteous; lateral areas of head, proster- num, and abdominal sterna more or less densely punctured; punctures black to somewhat metallic green. Legs pale reddish yellow; sparsely pubescent; distal half of femora and most of tibiae densely punc- tured with black. Detailed line drawings of eggs and all nymphal instars of H. halys are available in Kobayashi (1956, 1967) and Hoffmann (1931). The following key to instars and nymphal measurements and descriptions are taken, in part, from Kobayashi (1956, 1967). KEY TO INSTARS OF HALYOMORPHA HALYS 1. Anterior wing pads not developed (Figs. 3—5) a — Anterior wing pads well-developed (Figs. 6—7) POLS Da og GID EOE Oro De aOR Oe Rene 4 2. Middle of each tibia without a white band (igs 3-4) vere asus tess pctemeie eee Dele ashe. 3 — Middle of each tibia with a white band (Fig. 5) Third instar — Eyes spherically projecting (Fig. 4) Second instar 4. Posterior wing pads not developed (Fig. 6) . . Fourth instar — Posterior wing pads conspicuously developed (Fig. 7) Fifth instar Egg and egg cluster (Fig. 2): Length ap- proximately 1.6 mm, diameter 1.3 mm, el- liptical, chorion whitish with fine reticula- tion with minute spines. Micropylar projec- tions 30—32, capitate, white. Egg burster chitinized, blackish, T-shaped. Egg cluster consisting generally of 20-30 eggs, ar- ranged more or less regularly. First instar (Fig. 3): Length approxi- mately 2.4 mm. Body elliptical, not de- pressed. Head roundish, triangular. Head, thorax, dorsal plates, connexiva, and legs black, except lateral portions of each tho- racic tergum and each connexivum subhya- line or pale. Abdomen mostly yellowish red. Eye darkish red. Antenna reddish black. Second instar (Fig. 4): Length approxi- mately 3.7 mm. Body egg-shaped, more or less depressed. Head rectangular, with pair of hornlike projections in front of eyes. Head, thorax, dorsal plates, and connexiva mostly black, except lateral portions of each thoracic tergum and each connexivum sub- hyaline. Abdomen whitish with reddish spots and reddish junctions. Eye reddish black. Antenna reddish black except apical portion of third antennomere white. Legs blackish brown. Body above with large scattered punctures. Third instar (Fig. 5): Length approxi- mately 5.5 mm. Body pear-shaped, rather depressed. Head rectangular, with pair of hornlike projections in front of eyes. Head, thorax, dorsal plates, and connexiva mostly brownish black, except apical portion of median lobe near inner margins of lateral lobes, anterior portion of head in front of eyes, middle and some spots on thorax, lat- eral portions of anterior half of pronotum, middle of mesonotum and entire metano- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 228 VOLUME 105, NUMBER 1 tum, anterior portions of orifices of scent glands and middle portions of connexiva mostly yellowish brown. Abdomen whitish with reddish spots and junctions. Eye and antenna colored as in second instar. Legs blackish brown, except bases of femora and middle of tibiae white. Fourth instar (Fig. 6): Length approxi- mately 8.5 mm. Body as in previous instar, pear-shaped, rather depressed. Coloration of body almost same as third instar. Antenna reddish black, except apex of third anten- nomere and base of fourth antennomere yellowish white. Femora brown, scattered with black spots, except bases yellowish white. Tibiae and tarsi blackish brown ex- cept middle of each tibia yellowish white. Purplish black, shallow depression on inner surface of each connexivum at middle. Fifth instar (Fig. 7): Length approxi- mately 12 mm. Body as in previous instar, pear-shaped, rather depressed. Head, tho- rax, dorsal plates, and connexiva mostly brownish black with metallic luster, except some spots on head and thorax yellowish and some yellowish white spots on connex- iva. Anterior half of anterolateral margins of pronotum, lateral margins of mesonotum and middle of lateral margins of connexiva all whitish subhyaline. Abdomen luteous- white, with dense covering of black, metal- lic punctures, reddish junctions, and reddish spots. Eye reddish black. Antenna black, except apex of third and base of fourth an- tennomeres whitish. Femora mottled black- ish brown, bases whitish. Tibiae and tarsi mostly brown, except middle of tibiae white and apices of tarsi black. Thoracic sterna, including leg bases (coxae, trochanters, and basal areas of fem- ora) and abdominal venter of all instars, particularly the fifth (Fig. 8), whitish, sharp- ic Figs. 1-7. 229) Fig. 8. ventral aspect. Scale line = 5 mm. Fifth-instar nymph of Halyomorpha halys, ly contrasting with darker lateral margins of thorax and abdomen. SUMMARY OF BIONOMICS AND HABITS Aspects of the seasonal history, ecology, and developmental stages of H. halys (often cited as H. picus, H. brevis, or H. mista in the Asian literature) have been reported by Nozu and Sonoyama (1924), Suzuki (1924), Shizuoka Agricultural Experiment Station (1925), Takahashi (1930), Hoff- mann (1931), Abe and Mori (1942), Fukuda and Kitajima (1956), Kobayashi (1956, 1967, 1981), Saito et al. (1964), Yanagi and Hagihara (1986), Fujiie (1984, 1985), Ka- wada and Kitamura (1983a, b), and Chu and Zhou (1997). Overwintered adults emerge from their Halyomorpha halys. 1, Adult. 2, Egg cluster. 3, First-instar nymphs. 4, Second-instar nymph. 5, Third-instar nymph. 6, Fourth-instar nymph. 7, Fifth-instar nymph. Scale line for Fig. | = 10 mm; for Figs. 2— 4 = 1 mm; for Figs. 5—7 = 5 mm. 230 hibernation sites in early spring (probably by early to mid-April). Mating and egg lay- ing do not commence until about two weeks after adults emerge from overwin- tering sites. Under laboratory conditions, adults require 14—15 days from imaginal ecdysis to become sexually mature (Kawa- da and Kitamura 1983b). Sexually mature females usually mate on multiple occasions, as many as five times per day (Kawada and Kitamura 1983b). A female, mated only once, can lay eggs for nearly half her life span, but fecundity decreases in proportion to her age. The period of laying fertile eggs and fecundity increase with multiple copu- lations. Females deposit eggs on the lower leaf surfaces of host plants from May to late August. Egg clusters commonly contain 20 to 30 eggs, which hatch 4—5 days after de- position. As with other pentatomids, H. ha- lys has five nymphal instars (Kawada and Kitamura 1983b). Emerging adults of the first generation are generally observed in early to mid-Au- gust. Adults are most numerous in early August in parts of Korea (Chung et al. 1995). In Tsukuba, Japan (Ibaraki Prefec- ture), first generation adults are not found until early to mid-September. Light trap re- cords in Japan, from 1978 to 1986, indicate a single peak occurrence in most years from late July to early August (cited as H. mista) (Mortya et al. 1987). In the Allentown area (Lower Macungie Twp., The Rodale Work- ing Tree Center), fifth-instar nymphs and te- neral adults were common on foliage on August 23, 2002 (Gary Bernon, in litt.). Only one generation is produced annu- ally throughout most of the native range, but 1—2 generations have been reported (as H. picus) for central and southern Hebei Province, China (Zhang et al. 1993). Zhang et al. (1993) also noted that overwintered adults, with an average life span of 301 days, emerged from their overwintering sites from early April to mid-June. New generation adults seek overwintering sites from late August to late October. Hoffmann (1931) also reported multiple annual gen- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON erations in south China, near Canton. He noted that the first generation is completed before the middle of June, and that there are at least four, and possibly six, annual generations. There, eggs have been ob- served at the end of September and nymphs as late as mid-October. Duration of immature stages.—Nymphal development of H. halys apparently varies significantly depending on the location in Asia. For example, near Canton in south China, nymphs (cited as H. picus) devel- oped to the adult stage in as little as 27 days (Hoffmann 1931). In another laboratory study (Kadosawa and Santa 1981), nymphal development ranged from 29 to 54 days on soybean seed. In two separate rearing trials conducted in 1963 in the mountainous areas of the Tohoku districts in Japan, Saito et al. (1964) found that when eggs of H. halys (also cited as H. picus) were laid in mid- summer (July 11), presumably by overwin- tered females, adults appeared by August 19 (40 days for complete development from egg through five instars to adult). When eggs were laid in late summer (August 3), adults did not appear until October 3 (62 days for complete development). Develop- mental times from egg hatch through five instars in midsummer in the mountains of Japan’s Tohoku districts were as follows: egg (6 d), instar I (5 d), instar Il (6—7 d), instar IIIf (7-9 d), instar IV (6-8 d), and instar V (7-10 d). For eggs deposited later in the summer (August 3), developmental times increased moderately, particularly for fourth and fifth instars: eggs (5 d), instar I (4 d), instar I] (7-11 d), instar II (7-11 d), instar IV (15—19 d), and instar V (16—22 d) (Saito et al. 1964). One of us (MEC) reared H. halys in cap- tivity [at room temperature (20—22°C) and natural photoperiod (June—July)] to gain in- formation on numbers of eggs in an egg cluster and on developmental times for the immature stages (beginning with 10 over- wintered adults collected in Allentown on June 6, 2002). Egg clusters (n = 5; range of number of eggs/cluster = 25—28, mean VOLUME 105, NUMBER | = 26.8) were placed individually in small plastic boxes and supplied with cut stems (in aquapics) of leaves and fruits of com- mon chokecherry (Prunus virginiana L.) and changed every few days. Individuals from these five egg clusters were reared to adults. Our observations do not vary sig- nificantly from those reported in the Japa- nese literature, especially from localities in northern Japan. Developmental times were as follows: egg (5—9 d, mean = 7.2), instar I (4-6 d, mean = 5.4), instar I] (S—9 d, mean = 7.2), instar HI (5-8 d, mean = 6), instar TV (S—7 d, mean = 6.2), and instar V (9-11 d, mean = 10.25). Host PLANTS AND FEEDING DAMAGE Halyomorpha halys is a polyphagous horticultural pest of some importance in Ja- pan and elsewhere in its geographic range (Panizzi et al. 2000). In Japan, it severely injures various shade and fruit trees, vege- tables, pulses, and leguminous crops (Ko- bayashi 1967). In southern China, adults and nymphs cause considerable damage to flowers, stems, and pods of various beans, and also feed on flowers of Hibiscus rosa- sinensis L., stems of Celosia argentea L.., and fruits of Solanum nigrum L. and Bas- ella rubra L. (Hotfmann 1931). Host plants of economic significance include Citrus (citrus), Diospyros kaki L. (persimmon), Fi- cus (fig), Glycine max Merrill (soybean), Malus domestica L. (apple), Morus (mul- berry), Paulownia spp. (paulownia), Prunus avium L. (cherry), P. mume Siebold and Zuccarini (ume, Japanese apricot), P. per- sica Batsch (peach), Pyrus pyrifolia Nakai (pear), and reportedly some weeds (includ- ing Arctium spp.) (Shiraki 1952; JPPS 1966; Fujiie 1984; Yoshii and Yokoi 1984; Yuan 1984; Yasunaga et al. 1993; Chung et al. 1995; Funayama 1996, 2002; Watanabe 1996; Chu and Zhou 1997). Adults gener- ally feed on fruit, whereas nymphs feed on leaves, stems, and fruits. Halyomorpha halys is one of approxi- mately 50 insects that are considered major pests of various leguminous crops, and par- 231 ticularly soybean, in Japan (Kadosawa and Santa 1981, Kobayashi 1981, Kobayashi et al. 1972, Kobayashi and Oku 1976), Korea (Son et al. 2000), and China (Hoffmann 1931). Adults and nymphs of H. halys can cause serious yield loss by sucking sap from soybean seeds (Kadosawa and Santa 1981). The most serious damage to tree fruit (persimmon, for example) is observed from late August to late October in Korea (Chung et al. 1995). In Japan, most adult feeding damage to ‘Fuji’ apples is from ear- ly to mid-August, with the fruit most sus- ceptible to sucking (stylet) injury during the thickening period; the actual feeding injury appears as pitting and discoloration of the flesh (Funayama 1996). In non-astringent persimmon orchards in Korea, severe feed- ing symptoms by H. halys include physical changes of the fruit, such as concaving of the surface or its becoming dark blue in a “bull’s-eye” configuration and the flesh be- coming soft and “spongy” (Chung et al. 1995). Paulownia, which includes six species of deciduous trees native to China, is also a host for H. halys and it too can be severely affected by this stink bug. Species of this plant genus (at least two species are natu- ralized in the United States) have been planted as superior trees for more than 2000 years in eastern Asia. Paulownia species are highly suitable and popular as_ inter- cropping species in modern agroforestry and are used commonly for urban shelter- belts due to their rapid growth and attrac- tive high-canopy formation (Hiruki 1999). Paulownia witches’ broom, one of the my- coplasma diseases, is vectored or dissemi- nated primarily by H. halys (Jin et al. 1981, Zhu et al. 1982). Paulownia witches’ broom can greatly reduce the growth and vigor of trees, their severe decline causing prema- ture death (Shiozawa and Tsuchizaki 1992). In Japan, this disease represents a serious threat to the production of paulownia (P. tomentosa Steudel) timber, which is much used in the Orient for cabinet wood. To determine host plant preferences and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 9. magnification of leaf damage at feeding site; upper right insert: fifth-instar nymph feeding. feeding damage potential in North America, periodic surveys were conducted through- out the greater Allentown area in the spring and summer of 2002 by KMB and James F Stimmel (Pennsylania Dept. of Agriculture, Harrisburg) (JFS, personal commumuncia- tion). In late May and early June, adults (presumably overwintered) were beaten from the foliage of various trees and shrubs, including honeysuckle (Lonicera sp.), wal- nut (Juglans sp.), shadbush (Amelanchier sp.), butterfly-bush (Buddleia sp.), paulow- nia (Paulownia tomentosa), persimmon, and maples (Acer spp.). By early to mid- July, nymphs of several instars were beaten from some of the same foliage, as well as from basswood (Tilia sp.) and catalpa (Ca- talpa sp.) seedlings (KMB, personal com- munication). According to JFS (personal communication), fourth- and fifth-instar nymphs were abundant on persimmon and paulownia foliage at the Rodale Working Tree Center in the western part of Allen- town. Nearby, at a private residence, a large Feeding damage to Paulownia leaf (Paulownia tomentosa) by Halyomorpha halys. Upper left insert: g g ) 3 P ) population (perhaps hundreds) of nymphs, mostly fourth instars, were observed on a small and isolated group of butterfly-bushes (Buddleia). The nymphs fed on the leaves of Buddleia and Paulownia. Feeding dam- age on these hosts appeared as “stippled” areas, roughly circular and one-eighth inch in diameter. Under magnification, this “stippled”’ area is seen to consist of areas where the plant cells have been depleted in a “‘crow’s-foot’’ pattern (see Fig. 9). As in- jury progresses, the previously “‘stippled”’ areas become brown and scablike. STATUS AS A NUISANCE PEST In Asia, H. halys overwinters as adults and aggregates, sometimes in large num- bers, on the outside of buildings when it 1s seeking hibernation sites in the fall (Kobay- ashi and Kimura 1969). The invasion of homes, commercial establishments, and schools makes this stink bug a very serious nuisance pest in Japan (Inaoka et al. 1993, Watanabe et al. 1994a, b). The flight of H. VOLUME 105, NUMBER 1 halys to wall surfaces of buildings starts at the end of September and peaks around the third week in October. Adult flight ceases by the end of November. Similar aggregation and flight behavior have been observed in Pennsylvania for H. halys in residential areas in the fall of 2001. Wherever H. halys becomes established in the United States, it likely will become a serious household nuisance in a manner similar to the western conifer seed bug, Leptoglossus occidentalis Heidemann: the boxelder bug, Boisea trivittata (Say); and the multicolored Asian lady beetle, Har- monia axyridis (Pallas) (Mallis 1997). DISTRIBUTION Halyomorpha halys is reported from Ja- pan (Honshu, Shikoku, Kyushu), Korea, China (Anhui, Hebei, Heilongjiang, Henan, Jiangsu, Jilin, Liaoning, Neu Monggol, Shaanxi, Shanxi, Fujian, Guangdong, Gu- angxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Xizang, and Yunnan), and Taiwan (Kobayashi 1967; Rider et al. in press). Pennsylvania (Fig. 10), with 5 confirmed counties (Bucks, Lehigh, Monroe, North- ampton, and Pike) to date, is a new record for the United States and the Western Hemisphere. Confirmed sightings, based on credible descriptions provided by home- owners and, in some cases, specimens pro- vided, validate the establishment of H. ha- /ys in numerous communities in and around the Allentown/Bethlehem area, and from surrounding counties. These include: Pike Co.: Peck’s Pond. Monroe Co.: Saylors- burg, Swiftwater. Northampton Co.: North- ampton, Easton. Lehigh Co.: Orefield, Al- lentown/Bethlehem (including the follow- ing areas: Adams Island, Butztown, Cata- sauqua, Coopersburg, Coplay, Center Valley, Cetronia, Dorneyville, East Texas, Emmaus, Freemansburg, Fountain Hill, Greenawalds, Hellertown, Lanark, Macun- gie, Neffs, Springhouse Farms, Stiles, Wal- nutport, West Catasauqua, Westgate Hills, Whitehall, and Zionsville). These records 233 are not listed below under **Material ex- amined,” but are mapped in Fig. 10. Material examined.—All in the Cornell University Insect Collection (CUIC), Itha- ca, NY, except as noted. PENNSYLVA- NIA: Bucks Co.: Perkasie, 26 January 2002, R. Miller [1]. Lehigh Co.: Allentown, Albright Avenue (private residence), 21st Street (apartment complex), 3 October 2001, ERH and KMB [27, including 4, Na- tional Museum of Natural History (Wash- ington, DC); 2, North Dakota State Univer- sity (Fargo); 2, Florida State Collection of Arthropods (Gainesville)]; Girard Avenue (private residence), 3 October 2001, S. Yoder, coll. [4]; Muhlenberg College cam- pus and Trexler Memorial Park, 6 June 2002, ERH [10]. INTERCEPTIONS AT PORTS-OF-ENTRY North American interceptions of this ex- otic stink bug have been infrequent. For the period 1973-1987, only two interceptions of a species of Halyomorpha at U.S. ports- of-entry were recorded in the USDA’s “‘List of Intercepted Plant Pests”’; both were iden- tified as Halyomorpha picus (probably H. halys). One was intercepted in an aircraft from Japan in 1983 and the other in bag- gage from Korea in 1984. For 1989 to 1998, the USDA-APHIS’s Port Information Network (PIN) database lists eight intercep- tions of Halyomorpha from China, Korea, or Japan from aircraft, machinery and woodenware crating, machinery crating and miscellaneous cargo, and tractor soil (Peter Touhey, personal communication). Established populations of H. halys that were found in eastern Pennsylvania might have originated from aggregations of over- wintering adults that were accidentally in- troduced with international commerce, i.e.. via bulk freight containers from Japan, Ko- rea, or China. ACKNOWLEDGMENTS We thank Karen M. Bernhard (Lehigh County Cooperative Extension, Allentown, PA), James E Stimmel (Pennsylvania De- 234 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Farmersville 2 Gooes = ee" |Hellertown Fig. 10. County distribution of Halyomorpha halys in eastern Pennsylvania (upper left). Confirmed sightings of H. halys in the greater Allentown/Bethlehem area (bottom). partment of Agriculture, Harrisburg), and Gary Bernon (USDA-APHIS, Otis ANGB, MA) for sharing with us their field obser- vations on host plant records and feeding damage in the Allentown area; David A. Rider (North Dakota State University) for sharing information from his unpublished catalogue of world Pentatomidae on the synonymy of H. halys; Toshio Shono (Cor- nell University) for providing some English translation of Japanese; Robert A. Schall (USDA-APHIS-PPQ) for assisting in find- ing literature references on H. halys: Peter Touhey (Systematic Entomology Laborato- ry, USDA-ARS, Beltsville, MD) for shar- ing interception data; Kent Loeffler (Cor- nell University) for providing the photo- graphs used in Figs. 1-8 and for facilitating the figure reproductions; and A. G. Wheel- er, Jr. (Clemson University) and D. A. Rider for reviewing and giving critical comments on a draft of the manuscript. We are in- VOLUME 105, NUMBER 1 debted to J. Stimmel for providing the dig- ital images used in Fig. 9 and to him and K. Bernhard for also reviewing a draft of the paper. This research was supported by the Cor- nell University Agricultural Experiment Station federal formula funds, Project No. NYC-139413 to ERH, received from Co- operative State Research, Education, and Extension Service, U.S. Department of Ag- riculture. Any opinions, findings, conclu- sions, or recommendations expressed in this publication are those of the authors and do not necessarily reflect the view of the U.S. Department of Agriculture. LITERATURE CITED Abe, T. and Y. Mori. 1942. On the food plants and the infestation of Halyomorpha picus Fab. Nogyo- kenkyu 12: 139. [In Japanese. ] Chu, FE J. and Z. F Zhou. 1997. Study on control and observation of the bionomic characteristics of Halyomorpha picus Fabricius. Journal of Hebei Agricultural University 20(2): 12-17. [In Chi- nese. | Chung, B. K., S. W. Kang, and J. H. Kwon. 1995. Damage, occurrence, and control of hemipterous insects in non-astringent persimmon. orchards. RDA Journal of Agricultural Science, Crop Pro- tection 37(2): 376-382. [In Korean.] Fabricius, J. C. 1794. Entomologia systematica emen- data et aucta, secundum classes, ordines, genera, species, adjectis synonymis, locis, observationi- bus, descriptionibus. Hafniae, Proft. 8. Bd. IV: 1472. Fujiie, A. 1984. Stinkbug injury to pears. International Congress of Entomology Proceedings 17: 577. . 1985. Seasonal life cycle of Halyomorpha ha- /ys. Bulletin of the Chiba Experiment Station 26: 87-93. [In Japanese. ] Fukuda, J. and H. Kitajima. 1956. Halyomorpha picus Fab. Kaju-byogaichu-zusetzu, pp. 30-33. [In Jap- anese. | Funayama, K. 1996. Sucking injury on apple fruit by the adult of brown marmorated stinkbug, Haly- omorpha mista (Uhler). Annual Report of the So- ciety of Plant Protection of North Japan 47: 140— 142. [In Japanese.] . 2002. Oviposition and development of Haly- omorpha halys (Stal) and Homalogonia obtusa (Walker) Heteroptera: Pentatomidae) on apple trees. Japanese Journal of Applied Entomology and Zoology 46: 1—6. [In Japanese. | Hiruki, C. 1999. Paulownia witches’-broom disease important in East Asia. Acta Hort. (ISHS) 496: 63-68. 6.htm] Hoffmann, W. E. 1931. A pentatomid pest of growing beans in south China. Peking Natural History Bul- letin 5(3): 25-26. Inaoka, T., M. Watanabe, Y. Kosuge, and T. Kohama. 1993. Biology of house-invading stink bugs and some control trials against them in an elementary school in Hokkaido: 1. Species composition and seasonal fluctuations of house-invading activity. Japanese Journal of Sanitary Zoology 44(4): 341— 347. [In Japanese.] Japanese Plant Protection Society (JPPS). 1966. List of Important Diseases and Pests of Economic Plants in Japan (Appendage: Weeds). Nihon To- kushu Noyaku Seizo K. K., Tokyo. Jin, K. X., C. J. Liang, and D. L. Deng. 1981. A study of the insect vectors of witches’ broom in Pau- lownia trees. Linye Keji Tongxun 12: 23-24. [In Chinese. | Josifov, M. and I. M. Kerzhner. 1978. Heteropteren aus Korea. 2. Teil (Aradidae, Berytidae, Lygaeidae, Pyrrhocoridae, Rhopalidae, Alydidae, Coreidae, Urostylidae, Acanthosomatidae, Scutelleridae, Pentatomidae, Cydnidae, Plataspidae). Fragmenta Faunistica 23(9): 137-196. Kadosawa, T. and H. Santa. 1981. Growth and repro- duction of soybean pod bugs (Heteroptera) on seeds of legumes. Bulletin of the Chugoku Na- tional Agricultural Experiment Station, E 19: 75— 97. [In Japanese.] Kawada, H. and C. Kitamura. 1983a. Bionomics of the brown marmorated stink bug, Halyomorpha mista. Japanese Journal of Applied Entomology and Zo- ology 27(4): 304—306. [In Japanese.] 1983b. The reproductive behavior of the brown marmorated stink bug, Halyomorpha mista (Uhler) (Heteroptera: Pentatomidae). I. Observa- tion of mating behavior and multiple copulation. Applied Entomology and Zoology 18(2): 234— 242. Kobayashi, T. 1956. The developmental stages of some species of the Japanese Pentatomoidea (Hemip- tera), V. Transactions of the Shikoku Entomolog- ical Society 4(8): 120-130. . 1967. The developmental stages of some spe- cies of the Japanese Pentatomoidea (Hemiptera). XVI. Homalogonia and an allied genus of Japan (Pentatomidae). Applied Entomology and Zoolo- gy 2(1): 1-8. . 1981. Insect pests of soybeans in Japan. Mis- cellaneous Publication of the Tohoku National [http://www.actahort.org/books/496/496_ Agricultural Experiment Station 2: 1—39. Kobayashi, T. and S. Kimura. 1969. The studies on the biology and control of house entering stink bugs. Part I. The actual state of hibernation of stink bugs in houses. Bulletin of the Tohoku National 236 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Agricultural Experiment Station Morioka 37: 123-138. [In Japanese.] Kobayashi, T. and T. Oku. 1976. Studies on the distri- bution and abundance of the invertebrate soybean pests in Tohoku District, with special reference to the insect pests infesting the seeds. Bulletin of the Tohoku National Agricultural Experiment Station 52: 49-106. [In Japanese. ] Kobayashi, T., T. Hasegawa, and K. Kegasawa. 1972. Major insect pests of leguminous crops in Japan. Tropical Agriculture Research Series, No. 6: 109— 126. Mallis, A. 1997. Handbook of Pest Control: The Be- havior, Life History, and Control of Household Pests. Eighth Edition. Mallis Handbook & Tech- nical Training Company. 1,456 pp. McPherson, J. E. 1982. The Pentatomoidea (Hemip- tera) of Northeastern North America with Empha- sis on the Fauna of Illinois. Southern Illinois Uni- versity Press, Carbondale. 241 pp. Moriya, S., M. Shiga, and M. Mabuchi. 1987. Analysis of light-trap records in four major species of fruit- piercing stinkbugs with a special reference to body size variation in trapped adults of Plautia stali Scott. Bulletin of the Fruit Tree Research Sta- tion, Series A, 14: 79-94. Nozu, R. and I. Sonoyama. 1924. Fruit pest insects in Shimane Prefecture (9). Journal of Plant Protec- tion 11: 341-343. Panizzi, A. R., J. E. McPherson, D. G. James, M. Ja- vahery, and R. M. McPherson. 2000. Chapter 13. Stink Bugs (Pentatomidae), pp. 421-474. In Schaefer, C. W. and A. R. Panizzi, eds. Heterop- tera of Economic Importance. CRC Press, Boca Raton, Florida. 828 pp. Rider, D. A., L.-Y. Zheng, and I. M. Kerzhner. In press. Checklist and nomenclatural notes on the Chinese Pentatomidae (Heteroptera). I]. Pentatominae. Zoosystematica Rossica. Rolston, L. H. and FE J. D. McDonald. 1979. Keys and diagnoses for the families of Western Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae (Hemiptera). Journal of the New York Entomological Society 87(3): 189-207. Saito, Y., S. Saito, Y. Ohmori, and K. Yamada. 1964. Studies on bionomics of the bean bugs occurring in mountain regions, with particular reference to that of Halyomorpha picus and to the insecticidal tests in laboratory and field. Japanese Journal of Sanitary Zoology 15: 7-16. [In Japanese.] Shiozawa, H. and T. Tsuchizaki. 1992. Distribution of Paulownia witches’ broom and a survey of suck- ing insects occurring on Paulownia plants in Ja- pan. Proceedings of the Kanto-Tosan Plant Pro- tection Society 39: 259-260. [In Japanese. ] Shiraki, T. 1952. Catalogue of Injurious Insects in Ja- pan (Exclusive of animal parasites). Economic and Scientific Section, Natural Resources Divi- sion, Preliminary Study No. 71, Vol. III. General Headquarters, Supreme Commander for the Allied Powers, Tokyo, Japan. 162 pp. Shizuoka Agricultural Experiment Station. 1925. Rear- ing examination of stink bugs. Report Ope. Shi- zuoka Agricultural Experiment Station, Taisho 13: 247-249. [In Japanese. ] Slater, J. A. and R. M. Baranowski. 1978. How to Know the True Bugs (Hemiptera Heteroptera). Wm. C. Brown Company Publishers, Dubuque, Iowa. 256 pp. Son, C. K., S. G. Park, Y. H. Hwang, and B. S. Choi. 2000. Field occurrence of stink bug and its dam- age in soybean. Korean Journal of Crop Science 45(6): 405-410. [In Korean.] Stal, C. 1855. Nya Hemiptera. Ofversigt af Kongliga Svenska Vetenskaps-Akademiens Foérhandlingar 12(4): 181-192. Suzuki, M. 1924. On the stink bugs very injurious to the Citrus fruit. Journal of Plant Protection 11: 35-42, 95-100. [In Japanese.] Takahashi, S. 1930. Halyomorpha picus Fab., pp. 617— 620. Jn Insect Pests of Fruit Trees, Vol. 2. 1,225 PP- Uhler, P. R. 1860. Hemiptera of the North Pacific Ex- ploring Expedition under Com’rs Rodgers and Ringgold. Proceedings of the Academy of Natural Sciences of Philadelphia 12: 221-231. Walker, F 1867. Catalogue of the specimens of Het- eropterous-Hemiptera in the collection of the Brit- ish Museum. Part 1: 1—240. Watanabe, K. 1996. Characteristics of damage of Ly- gocoris lucorum (Meyer-Dur) (Heteroptera: Miri- dae) and Halyomorpha halys (Stal) (Heteroptera: Pentatomidae) on cherry. Annual Report of the Society of Plant Protection of North Japan 47: 143-144. [In Japanese.] Watanabe, M., R. Arakawa, Y. Shinagawa, and T. Oka- zawa. 1994a. Overwintering flight of brown mar- morated stink bug, Halyomorpha mista, to the buildings. Japanese Journal of Sanitary Zoology 45(1): 25-31. [In Japanese. ] 1994b. Anti-invading methods against the brown marmorated stink bug, Halyomorpha mista, in houses. Japanese Journal of Sanitary Zoology 45(4): 311-317. [In Japanese. ] Yanagi, T. and Y. Hagihara. 1980. Ecology of the brown marmorated stink bug. Plant Protection 34: 142-148. [In Japanese. ] Yasunaga, T., M. Takai, I. Yamashita, M. Kawamura, and T. Kawasawa. 1993. A Field Guide to Japa- nese Bugs—Terrestrial Heteropterans. Zenkoku Noson Kyoiku Kyokai, Publishing Co., Ltd., To- kyo. 380 pp. [In Japanese. ] Yoshii, T. and N. Yokoi. 1984. The damage on the mulberry shoot caused by the brown marmorated stinkbug, Halyomorpha mista. Journal of Sericul- tural Science of Japan 53(1): 81—82. [In Japanese. ] VOLUME 105, NUMBER | Yuan, T. L. 1984. Some studies on witches’-broom dis- ease of Paulownia in China. International Journal of Tropical Plant Diseases 2(2): 181-190. [In Chi- nese. | Zhang, C., D. Li, H. Su, and G. Xu. 1993. A study on the biological characteristics of Halyomorpha pi- cus and Erthesina fullo. Forest Research 6(3): 271-275. [In Chinese.] Zhus BeiMeseZ, ¥.'Chen, Ji*¥. Shen, I. 'Chien} Rik Hong, P. X. Shao, D. Q. Tong, and C. H. Peng. 1982. Mycoplasma-like organism and virus in Paulownia trees infected with witches” broom dis- ease. Acta Biochimica et Biophysica Sinica 14(4): 393-396. [In Chinese. ] Note added in proof.—In late August 2002, specimens of H. halys were retrieved 2317 from light traps in Phillipsburg (Warren Co.) and Little York (Hunterdon Co.), New Jersey that were operated by Rutgers Uni- versity Cooperative Extension (Barry Emens, USDA-APHIS-PPQ, Trenton, NJ, personal communication). These collections represent a new state record for New Jersey. While conducting a delimiting survey near Allentown, Pennsylvania, James Stim- mel, Gary Bernon, and ERH collected H. halys at Stewartsville (Warren Co.), New Jersey on October 2, 2002, and at Mertz- town (Berks Co.), Pennsylvania on October 3, 2002. Berks County is a new county re- cord for Pennsylvania. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 238-239 NOTE A New Species of the Asian Predaceous Midge Genus Pseudostilobezza Wirth and Ratanaworabhan (Diptera: Ceratopogonidae) from Hainan, China Pseudostilobezza Wirth and Ratanawor- abhan of the predaceous midge tribe Cera- topogonini was proposed for the type spe- cies from Vietnam, P. macclurei (Wirth and Ratanaworabhan 1973). The following new species from Hainan Island is only the sec- ond known species of the genus and the first record of the genus from China. For an explanation of ceratopogonid ter- minology, see Tokunaga and Murachi (1959) and Das Gupta and Wirth (1968); for terms of the tribe Ceratopogonini, see Wirth and Grogan (1988). Pseudostilobezzia wirthi Yu and Yan, new species (Figs. 1-8) Diagnosis.—The female is distinguished from that of the only other species in the genus, P. macclurei, by the presence of ma- crotrichia at apex of the wing, the costa pro- duced beyond the end of R,,;, and segment 3 of the palpus without a sensory pit. Female.—Wing length, 1.32 mm, wing breadth, 0.44 mm. Head: Dark brown in- cluding antenna and palpus. Eyes separated by space equal to diameter of | facet, with interfacetal pubescence (Fig. 3). Antenna with lengths of flagellomeres in proportion of 34:19:18:17:20:20:22:23:34:32:34:34: 38; antennal ratio 0.99 (Fig. 2). Palpal seg- ments with lengths in proportion of 8:17: 16:10:15; third segment with length to breadth ratio of 3.4, without sensory pit but with several elongate sensilla on mesal sur- face of distal portion (Fig. 4). Clypeus (Fig. 5) shield-shaped, with row of 4 dorsal setae and ventral row of 2 setae. Mandible with 7 well-developed teeth (Fig. 6). Thorax: Dark brown; scutum with sparse, fine, hair- like setae; scutellum with 4 bristles. Wing membrane (Fig. 1) covered with distinct microtrichia, macrotrichia present at wing tip. Costa produced beyond R,,; and slight- ly curved; costal ratio 0.85. Two well-de- veloped radial cells, second about 1.5 lon- ger than first. Legs with femora and tibiae brownish; hind tibial comb with 7 spines; claws large and subequal on foreleg, small and equal on mid- and hind legs (Fig. 8): tarsal ratio of fore-, mid-, and hind legs 2.66, 4.06, 2.94. Abdomen: Brown. Sper- mathecae (Fig. 7) elongate, oval, unequal, largest measuring 0.083 mm x 0.038 mm, plus a small, round rudimentary third. Male.—Unknown. Holotype.—Female, Qionzhong, Hainan Province, China, 13-III-1995. Deposited in the Medical Entomology Collection Gal- lery, Beijing 100071, China. Etymology.—Named for the late Dr. W. W. Wirth, in recognition of his important contributions to the study of world Cera- topogonidae. Discussion.—This new species is very similar to P. macclurei; however, P. mac- clurei differs from P. wirthi by the lack of macrotrichia on the wing tip, the first radial cell much shorter than the second, the costa not produced beyond R,,;, and palpal seg- ment 3 with a small sensory pit. Acknowledgments.—We are grateful to Art Borkent, Enderby, British Columbia, Canada, and William L. Grogan, Jr., Salis- bury University, Salisbury, MD, U.S.A., for reviewing earlier drafts of the manuscript. LITERATURE CITED Das Gupta, S. K. and W. W. Wirth. 1986. Revision of the Oriental species of Srilobezzia Kieffer (Dip- tera: Ceratopogonidae). United States National Museum Bulletin 283: 1-149. VOLUME 105, NUMBER 1 239 IRAN el Re 3 4 265) Figs. 1-8. Pseudostilobezzia wirthi. 1, Wing. 2, Antennal flagellum. 3, Interocular space. 4, Palpus. 5, Clyp- eus. 6, Mandible. 7, Spermathecae. 8, Tarsi and claws of fore-, mid-, and hind legs (right to left). 2 Tokunaga, M. and E. K. Murachi. 1959. Insects of Mi- idae). Florida & Fauna Handbook No. 4, E. J. cronesia Diptera: Ceratopogonidae. Insects of Mi- Brill, xv + 160 pp. cronesia 12(3): 103-434. Wirth, W. W. and N. C. Ratanaworabhan. 1973. Pseu- dostilobezzia, a new genus of biting midge from Yi-xin Yu and Ge Yan, /nstitute of Mi- Viet Nam (Diptera: Ceratopogonidae). Proceed- crobiology and Epidemiology, 20 Dongda ings of the Entomological Society of Washington 75: 177-179. Jie, Fengtai, Beijing 100071, People’s Re- Wirth, W. W. and W. L. Grogan, Jr. 1988. The preda- PUDlic of China (e-mail: yuyixin@ btamail. ceous midges of the world (Diptera: Ceratopogon- —et.cn). PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 240-242 NOTE First report of Epitraninae (Hymenoptera: Chalcididae) in the Nearctic The single cludes 64 described species (Noyes 2001). Boucek (1988) summarized the world dis- tribution of the genus as Africa, South Asia, and Australia, and considered the subfamily endemic to the Old World (Boucek 1992). He reported, however, that Epitranus cla- vatus Fabricius had been found in the New World on scribed from Guyana (Fabricius 1804) and subsequently reported from St. Vincent (Walker 1834) and Brazil (Ashmead 1904) as E. fulvescens Walker, and from Hawaii as E. lacteipennis Cameron (1883), with both names subsequently synonymized un- der E. clavatus (Bouéek 1982). This species also occurs in India, Malaysia, Taiwan, and Seychelles (Boucek 1982). Based on the distribution of Known species of Epitranus, Boucek (1982) suggested that the presence of Epitranus clavatus in the New World was the result of multiple introductions from the Old World into **... parts of trop- ical America before effective quarantine subfamily Epitraninae contains the genus Epitranus Walker, which in- several occasions. It was de- measures Were introduced.” Epitranus cas- taneus Cresson (1865) was described from Cuba, but because no one has examined its type, Boucek (1982) suggested that it might also be a synonym of E. clavatus. Burks (1936) suggested that the genus Epitranus might eventually be found in Florida based on the proximity of E. fulvescens in St. Vin- cent and E. castaneus in Cuba. In this note we report the first record of Epitranus clavatus (Fig. 1) as well as the subfamily Epitraninae (Chalcididae) in the Nearctic Region. A single female specimen was collected with the following informa- tion: USA, West Virginia, Hardy County, 3 miles NE Mathias, 38°55'N, 78°49'W, 17.V—3.VI-2002, ca. 2,200’ elevation, Mal- aise trap, D. R. Smith. The trap was set in disturbed areas within an eastern deciduous forest that is interspersed with relatively distantly-spaced “*weekend” cabins. Our identification of Epitranus clavatus was based on the key in Boucek (1982) and on voucher specimens in the National Col- lection of Natural History, Washington, D.C. identified by B. D. Burks. It is a mem- ber of the clavatus-group, including only one other taxon, E. ramnathi (Mani and Dubey). The species can be identified based upon the following set of characters: The marginal vein is distinct and the forewing pilosity is sparse and virtually absent be- neath the vein; the propodeum is matte (not shiny) with granulate sculpture and has a distinct median cell delimited by two sub- median carinae; the metafemur has fewer than 15 teeth and a distinct subbasal tooth (or angle); the tarsal sulcus of the metatibia is traversed for its entire length by an oblique carina; and the metasomal petiole is 3 (female) to 4 (male) times as long as broad. Several factors are remarkable about this discovery. All known specimens of New World EF. clavatus 1901 (Boucek 1992), thus the specimen dis- covered in West Virginia represents the first reported in over one hundred years from the New World from the Nearctic. It is the only specimen from thousands of Chalcidoidea collected and examined from 20—21 Malaise traps run in the area each year for 3 years. Thus, the species does not appear to be common either locally or temporally. Finally, its dis- covery in a sparsely populated area, miles from any port of entry, suggests that the species is more widely distributed than this single record indicates. Host records are known tor only a few species of Epitranus; generally they are were collected before as well as the only specimen VOLUME 105, NUMBER | Figw 1: Epitranus clavatus, female, habitus larval or pupal parasitoids of Pyralidae and Tineidae (Lepidoptera), some of which de- velop in stored products, and one species was reportedly reared from a Camponotus nest (Formicidae) (Boucek 1992). Epitran- us clavatus has been reared from the pupa of a “case bearing cotton moth” (in India) and from Tinea palaechrysis Meyrick or Crypsithyris sp. in Malaysia (Boucek 1982). According to Don Davis (personal communication), both moth genera are closely related and have a wide range of hosts including stored cotton and fabrics. It is likely that E. clavatus is transported via its moth host in shipping bags, cloth- ing, and carpeting all made from natural fibers. We thank Don Davis, Department of En- tomology, Smithsonian Institution, for in- formation pertaining to the hosts of Epi- T. Henry and M. Gates, Entomology Laboratory, and Gary Gibson, Agriculture and Agri-Food tranus Clavatus: Systematic Canada, for reviewing the manuscript; Ca- thy Apgar and Terry Nuhn, Systematic En- tomology Laboratory, the former for pro- ducing the photograph (Fig. 1) and the lat- ter for preparing the specimen; and the fol- lowing individuals who allowed placement of traps on their property in Hardy County, West Virginia: R. Gordon, R. Gray, S. Hen- derson, T. and K. Henry, D. and J. Miller, G. Wood, and N. Booth. LITERATURE CITED Ashmead, W. H. 1904. Classification of the chalcid flies. Memoirs of the Carnegie Museum 1(4): 1x1, 225-551 Boucek, Z Epitranus 62 9 1982. Oriental chalcid wasps of the genus Journal of Natural History 16: 577 1988 Australasian Chalcidoidea (Hymenop tera). A biosystematic revision of genera of four teen families, with a reclassification of species CAB International, Wallingford, UK, 832 pp 1992. The New World genera of Chalcididae (Hymenoptera). Memoirs of the American Ento mological Institute 53: 49-117 242 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Burks, B. D. 1936. The Nearctic Dirhinini and Epi- tranini (Hymenoptera, Chalcididae). Proceedings of the National Academy of Sciences Washington, Zoology 22: 283-287. Cameron, P. 1883. Descriptions of new genera and spe- cies of Hymenoptera. Transactions of the Ento- mological Society of London 1883: 187-197. Cresson, E. T. 1865. On the Hymenoptera of Cuba. Proceedings of the Entomological Society of Phil- adelphia 4: 1—200. Fabricius, J. C. 1804. Systema Piezatorum. C. Rei- chard, Brunsvigae [Braunschweig], 440 + 30 pp. Noyes, J. S. 2001. Interactive Catalogue of World Chalcidoidea (2001—second edition). CD-ROM. Taxapad and The Natural History Museum, Lon- don. Walker, E 1834. Monographia Chalciditum (contin- ued). Entomologist’s Magazine 2: 13-39. E. E. Grissell and D. R. Smith, System- atic Entomology Laboratory, PSI, Agricul- tural Research Service, U.S. Department of Agriculture, %c National Museum of Natu- ral History, Smithsonian Institution, Wash- ington, DC 20560-0168, U.S.A. (e-mail: egrissel@sel.barc.usda.gov; drsmith@sel. barc.usda.gov) PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 243-245 NOTE Discovery of Monodontomerus osmiae Kamijo (Hymenoptera: Torymidae) in the New World Monodontomerus osmiae Kamijo is a gregarious ectoparasitoid of megachilid bees. It was first discovered and named in Japan where it was reared from Osmia tau- rus Smith and O. excavata Alfken (Kamo 1963). Subsequently it was reared from O. cornifrons Radoszkowski in Japan (Kamijo 1965). Since then it has been reported only from the “Russian Far East” (ex taurus) by Zerova and Romasenko (1986), who keyed and figured the species in a pa- per on the parasitoids of megachilid bees in the former USSR. Zerova and Seryogina (2002) reviewed the entire Old World fauna of Monodontomerus but added no addition- al information; they did include M. osmiae in a key to the known species of that region. I recently discovered Monodontomerus Osmia osmiae attacking Osmia spp. nesting in straw bundles set out in my Silver Spring, Maryland, garden (39.076°N, 77.002°W). This is the first report of the parasitoid in the New World. A bundle of 88 paper nest- ing straws (4 * 8”) was placed in a 39 ounce coffee can and set out in the spring of 2001 under the protection of an overhang on the north side of my house. The straws and can were oriented horizontally and placed on a ledge about 5 feet above ground. In spring of 2001 many of the straws were used by several species of Os- mia and Megachile. A few adult bees were collected to determine the genera involved, but no attempt was made to determine the species. These straws overwintered outside, and in April and May of 2002 the same set of straws were reused, this time entirely by Osmia (based on the entrance plugs all be- ing made of mud). In early to mid-June, 2002, large num- bers of Monodontomerus were seen flying about the area and landing on the exposed outer edges of the straws. Adults were seen entering the interstices between straws, but no observations of oviposition were made. One collection of adult wasps was made in mid-June by aspirating specimens from the straw surfaces and by sweeping the air space near the nests and aspirating the re- sultant catches. These specimens were placed in alcohol but not identified until July 15. They consisted of two species: M. osmiae (20 females) and M. obscurus West- wood (21 females, 7 males). Another col- lection was made on 10 July. This consisted of 17 females and | male of M. osmiae and 116 females and 32 males of M. obscurus. These two random samples and the speci- mens collected represented only a small portion of the total specimens seen flying around the nest site. They were collected in an attempt to reduce parasitism rates on the nesting Osmia and to determine which of the common local species of Monodonto- merus were attacking the bees. When all specimens were identified in mid-July, and M. osmiae was detected, an attempt was made to collect additional par- asitoids at the nest site, but the numbers of specimens had fallen off. On 21 July, 4 fe- males and 7 males of the common species M. aeneus (Fonscolombe) were collected. On 30 July, 2 females of M. osmiae were collected as well as 5 females and 4 males of M. aeneus. On | August, the straws were placed in a plastic rearing box and brought indoors. Between 5—23 August, 33 females and 12 males of M. aeneus emerged. By | September emergence stopped and the straws were placed in the freezer. No spec- imens of M. osmiae were actually reared from the straws. The New World species of Monodonto- merus were revised by Grissell (2000), who 244 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2 Monodontomerus osmiae, male head. 1, Figs. 1-2 Face, 2, Dorsum, antennae removed examined over 5,000 specimens, reared mostly from solitary bees. A large number of specimens were reared from areas in Maryland, and no ©. osmiae were found up to that time. Monodontomerus species are generally difficult to identify, but /. osmiae is unusually distinct, especially in males, which have the face abnormally concave (Figs. 1— 2). Females of M. osmiae, unlike other Monodontomerus, have virtually no speculum (bare area) on the which makes it almost entirely setose. It would have been difficult to overlook such forewing, an anomalous species in either sex among all the specimens examined for the study. This suggests that M. osmiae may have moved into the area relatively recently. Little is known of the biology of Mono- dontomerus osmiae. Iwata and Tachikawa (1966) reported a preponderance of females for rearings of this species from Osmia tau- cocoons 87 males and 726 rus. From 61 females emerged. The number of parasit- oids per host (counted for 4 cocoons only) varied from 14 to 26. The gregarious nature of the parasitoid larvae and the highly fe- male-skewed sex ratio make this parasitoid particularly effective at rapidly increasing its numbers at the expense of its host. It is of more than passing interest that osmiae was reared in the Old World from Osmia cornifrons (the Japanese hornfaced bee), a bee that has been widely naturalized in the United States since the 1970’s and 1s widely available commercially (MAAREC 2001). It would be instructive if keepers and retailers of solitary bees were to sample the parasitoids at their nest sites for the presence of M. osmiae and determine the Some attempt should also be made to determine if M. os- extent of its distribution. miae attacks indigenous solitary bees. If so, it would be a case of an introduced para- sitoid being detrimental not only to its in- troduced beneficial host, but also to the na- tive fauna of its adopted homeland. I thank Mike Gates, Systematic Ento- mology Laboratory, and Gary Gibson, Ag- riculture and Agri-Food Canada, for re- viewing the manuscript. I also thank Mike Gates for providing the illustrations. LITERATURE CITED Grissell, E. E. 2000. A Revision of New World Mon- odontomerus Westwood (Hymenoptera: Chalci- doidea: Torymidae). Contributions of the Ameri can Entomological Institute 32(1): 1-90. Iwata, K. and T. vations on 53 species of the supertamilies, Chal- Tachikawa. 1966. Biological obser- cidoidea and Proctotrupoidea, from Japan (Hy- menoptera: Transactions of the Shikoku Entomological Society 9: 1-29. Kamijo, K. 1963. A revision of the species of the Mon- Apocrita) [sic] odontomerinae occurring in Japan (Hymenoptera: Chalcidoidea) [Taxonomic studies on the Tory- midae of Japan, If]. Insecta Matsumurana 26: 89 OS 1965. A new host-record of Monodontomerus osimae Kamijo. Insecta Matsumurana 28: 78. MAAREC 2001. Basic biology and management of the Japanese horntaced bee. Mid-Atlantic Apicul- tural Research & Extension Consortium. Publi cation 5.5, 3 pp Zerova, M. D. and L. P- Romasenko. 1986. Chalcidoids (Hymenoptera, Chalcidoidea), parasitoids of leat- VOLUME 105, NUMBER 1 245 cutting bees (Hymenoptera, Megachilidae), in the E. E. fauna of the USSR. [In Russian]. Zoologicheskii Laboratory, PSI Agricultural Research Zhurnal 65: 1631-1643. ; ‘ Zerova, M. D. and L. Ya. Seryogina. 2002. A revision are, eS: Oa eae OF PA STRCULUTS, i of Old World Monodontomerus (Hymenoptera: National Museum of Natural History, Chalcidoidea: Torymidae). National Academy ot Smithsonian Institution, Washington, DC Sciences of Ukraine, I. I. Schmalhausen Institute 20560-0168, U.S.A. (e-mail: egrissel@ sel. of Zoology, Special Issue, Kiev, 74 pp. barc.usda.gov) Grissell, Systematic Entomology NEW PUBLICATION “Holcocerini of Costa Rica (Lepidoptera: Gelechioidea: Coleophoridae: Blastobasinae)”’ by David Adamski Entomological Society of Washington Memoir No. 24, 147 pp. See inside back cover for ordering information. PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 246-248 NOTE Expanding Ranges for Four Species of Invasive Sawflies (Hymenoptera: Tenthredinidae) in Eastern United States Malaise trap collecting in the mid-Atlan- tic states has resulted in several significant southern range extensions in the eastern United States for four species of adventive sawflies. These collections indicate that these species continue to expand their rang- es in North America since they were first discovered. All four species are apparently of European origin and follow a similar pat- tern of introduction. All were originally re- corded from the northeastern United States or southeastern Canada, and three were in- dependently introduced into British Colum- bia. Here I report the first records for three species south of the state of New York and one species new to Maryland. All four spe- cies are known now from as far south as about the latitude of Washington, DC. No specimens were found in 14 years (1986— 1999) of extensive Malaise trap collecting approximately 100 miles south of Washing- ton, DC, in Louisa and Essex counties Vir- ginia. Specimens are deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC. Hoplocampa testudinea (Klug) (Tenthre- dinidae: Nematinae) (European apple saw- fly).—This pest of apple was first recorded from Long Island, NY, in 1939 and British Columbia in 1940 (Pyenson 1943). It has since been recorded from CT, MA, NH, NJ, VT, and RI (Smith 1979), and more recently from PA (anonymous 2002a) and the Kear- nysville area of WV (anonymous 2002b). Some records may be based on larvae or the characteristic damage to apple fruits which the young larvae leave a winding feeding scar on the surface and older larvae bore deeply into the core, usually causing mid-season fruit abortion (Weires 2001). Though the within larvae live and feed; damage attributed to this sawfly is probably correct, I prefer to base records on adults even though they are short-lived and more difficult to find than the damage or larvae. The following, based on one female, is the first record for Maryland and the most southerly record known. Records: MARYLAND: Montgomery Co., 4 mi SW of Ashton, 39°06'30"N, 77°01'30"W, Malaise trap, 18 April 2002, GE Hevel: Allantus viennensis (Schrank) (Tenthre- dinidae: Allantinae).—This species feeds on Rosa spp., and occurs on cultivated ro- ses. It was first reported from the area of Ithaca, NY (Smith 1975), and from the same locality by Smith (1979). The second collection is from my residence in Fairfax Co., VA, where I reared this species from larvae feeding on cultivated roses in 1982. It has subsequently been collected in a trap in back of the house as well as from other collection sites in northern Virginia and West Virginia, indicating that it is well es- tablished and generally distributed in this area. There are several generations a year; specimens have been taken from April through August. Records: VIRGINIA: Clarke Co., Uni- versity of Virgina Blandy Experimental Farm, 1 mi. S Boyce, 39°05’N, 78°10'W, Malaise trap, D. R. Smith (many specimens each year, 1990-1995); Fairfax Co., near Annandale, 38°50'N, 77°12'W, D. R. Smith (first reared in 1982 and subsequently taken from roses and in a Malaise trap for a num- ber of years from 1983-1997); Loudoun Co., near junction of Goose Cr. and Sycolin Road, Malaise trap, C. J. Anderson (col- lected each year, 1998-2000); 12969 Tay- lorstown Road, Malaise trap, C. J. Ander- son (2000). WEST VIRGINA: Hardy Co., VOLUME 105, NUMBER 1 3 mi NE of Mathias, 38°55'’N, 78°49'W (collected in 2000 and 2001). Nesoselandria (Fabricius) (Ten- thredinidae: Selandriinae).—This species, the larvae of which feed on mosses (Vik- berg and Nuorteva 1997), was first recorded in North America from Ontario, Quebec, and British Columbia (Smith 1967), and subsequently recorded from NH (Smith 1969), and NY (Smith 1979). It also occurs in Maine (Franklin Co., 1967) and Michi- gan (Dickinson Co., 1982, 1983; Baraga Co., 1981). The earliest North American re- cord I have seen is “Sclaterville Wild Flower Preserve, New York, May 27, 1938.” Collections now establish this spe- cies in Maryland and Virginia. The trap in Fairfax Co., VA, has been run in the same place since 1981, but it wasn’t until the nineteenth season that the species was col- lected. Records: MARYLAND: Garrett Co., Finzel Swamp, 2 km S Finzel, 39°38’N, 79°00'W, V-21—30-1992, VI-30—VII-10- 1993, Malaise trap, E. Barrows and D. R. Smith. VIRGINIA: Fairfax Co., near An- nandale, 38°50'N, 77°12’W, VIII-5—11- 2001, VI-16—22-2002, D. R. Smith. Heptamelus (Stephens) (Tenthredinidae: Selandriinae).—Benson (1962) first recorded this species from Brit- ish Columbia from specimens collected in 1953. Smith (1969) saw only the specimens Benson recorded, and Smith (1979) record- ed it only from British Columbia. Accord- ing to Benson (1952) larvae bore down- wards in the stems of ferns of the genera Athyrium, Polypodium, and Blechnum, but the species appears to prefer Athyrium (Shaw and Bailey 1991). The first eastern North America record I have seen is from New York, Westchester Co., Armonk, Cald- er Center, V-30—VI-5-1974. I have collected specimens from Maryland and Virginia. Most records are April, May, and June, with a few from August and September. Records: MARYLAND: Prince Georges’ Co., Beltsville Agricultural Research Cen- ter, 39°02’N, 76°52'W, D. R. Smith (1992 morio ochroleucus 247 and 1993). VIRGINIA: Fairfax Co., near Annandale, 38°50'N, 77°12’W (many spec- imens, first collected in 1984 and in most years to the present). Acknowledgments.—I thank the follow- ing for allowing collections on their prop- erties: M. Bowers, University of Virginia Blandy Experimental Farm, Clarke Co., Virginia; T. J. Henry, D. R. Miller, R. Gray, S. Henderson, and R. D. Gordon, Hardy Co., West Virginia; V. Power and T. Carlow, Loudoun Co. Virginia; J. and B. Kloke, Louisa and Essex counties, Virginia; and Maryland Nature Conservancy, Finzel Swamp, Garrett Co., Maryland. LITERATURE CITED Anonymous. 2002a. European apple sawfly. Pennsyl- vania Tree Fruit Production Guide. Part II—Dis- eases, pests and natural enemies. http://tfpg.cas. psu.edu/part2/part22bk.htm. . 2002b. European Apple Sawfly, Hoplocampa testudinea (Klug). http://www.ento.vt.edu/fruittiles/ EAS.html. Benson, R. B. 1952. Hymenoptera (Symphyta). Family Tenthredinidae, pp. 51-137. Jn Royal Entomologi- cal Society of London, Handbooks for the Identi- fication of British Insects, Vol. 6, pt. 2(b). . 1962. Holarctic sawflies (Hymenoptera: Sym- phyta). British Museum (Natural History) Ento- mology Bulletin 12: 281—409. Pyenson, L. 1943. A destructive apple sawfly new to North America, Journal of Economic Entomology 36: 218-221. Shaw, M. R. and M. Bailey. 1991. Parasitoids (Hy- menoptera: Braconidae, Ichneumonidae, Pterom- alidae) and notes on the biology of the fern-boring sawfly Heptamelus ochroleucus (Stephens) (Hy- menoptera: Tenthredinidae) in the English Lake District. The Entomologist 110: 103—109. Smith, D. R. 1967. Two sawflies new to North Amer- ica (Hymenoptera: Tenthredinidae). Proceedings of the Entomological Society of Washington 69: 95. 1969. Nearctic sawflies I. Selandriinae: Adults (Hymenoptera: Tenthredinidae). United States Department of Agriculture, Technical Bul- letin No. 1398, 48 pp., 10 plates. . 1975. A rose sawfly new to North America (Hymenoptera: Tenthredinidae). Cooperative Eco- nomic Insect Report 25(10): 163-165. . 1979. Symphyta, pp. 3-137. Jn Krombein, K. V., P. D. Hurd, Jr, D. R. Smith, and B. D. Burks, eds. Catalog of Hymenoptera in America North of 248 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mexico, Vol. 1. Smithsonian Institution Press, Sawfly. http://www.nysipm.cornell.edu/factsheets/ Washington, DC. treefruit/pests/eas/eurapsawfly.html. Vikberg, V. and M. Nuorteva. 1997. On the rearing of Nesoselandria morio (Fabricius) and Birka ciner- David R. Smith, Systematic Entomology eipes (Klug) (Hymenoptera, Tenthredinidae), with — 7 gboratory, PSI Agricultural Research Ser- descriptions of their larvae. Entomologica Fennica ; ; Se acsan vice, U.S. Department of Agriculture, “oc Na- Weires, R. W.. Jr. 2001. New York State Integrated tional Museum of Natural History, Smith- Pest Management Program Fact Sheets. Insect SO/ldN Institution, Washington, DC 20560- Identification Sheet No. 20, 1991. European Apple 0/68 (e-mail: dsmith@ sel.barc.usda.gov) ENTOMOLOGICAL SOCIETY OF WASHINGTON REGULAR MEETINGS, 2002—2003 October 3, 2002. Thomas J. Henry, Systematic Entomology Laboratory, USDA, Wash- ington, DC. “‘Ecuadorian Bug-Collecting Adventures: From the Amazon to the Andes” November 7, 2002. Michael Gates, Systematic Entomology Laboratory, USDA, Wash- ington, DC. **Eurytomidae (Hymenoptera: Chalcidoidea): Morphology, Phylogeny, and the New Mexico Bootheel” December 4, 2002. Daniel H. Janzen, University of Pennsylvania, Philadelphia. ““How to Find All the Species of Caterpillars in a Large, Complex Tropical Habitat” January 9, 2003. Gabriela Chavarria, National Wildlife Federation, Reston, VA. “Capitol Hill-Topping in D.C. and Conservation Policy” February 6, 2003. Rob Raguso, University of South Carolina, Columbia. Sphingids and Fragrance (general topic). March 6, 2003. Stuart H. McKamey, Systematic Entomology Laboratory, USDA, Wash- ington, DC. “Hopp(er)ing through Mexico and Venezuela: Adventures in Leathopper- Hunting” April 3, 2003. David A. Nickle, Systematic Entomology Laboratory, USDA, Beltsville, MD. “Life in a Rainforest Canopy: An In-Depth Study of Amazonian Grasshoppers, Katydids, and Termites” May 1, 2003. Barry Knisley, Randolph-Macon College, Ashland, VA. “Tiger Beetle Con- servation: Case Studies with Impacts, Management and Natural History” Meetings at 7:00 PM, Cathy Kerby Seminar Room (CE-340) National Museum of Natural History 10th & Constitution Ave., NW, Washington, DC Meetings Open to the Public New Literature, Specimen Demonstrations, Ento-T-Shirts, etc. Welcome Pre-Meeting Dinner: R. Reagan Building Food Court, 5:30 PM PROC. ENTOMOL. SOC. WASH. 105(1), 2003, p. 249-254 MEMBERSHIP LIST OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The previous list was published in October 1999 with 429 members; the present list contains 418 members from every state in the union except Delaware, Indiana, Maine, Nevada, Oklahoma, Rhode Island, and South Dakota. The largest representation is in Mary- land (48), followed by the District of Columbia (40), California (26), Florida (19), and Virginia (16). 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 25 other countries, on six continents, are represented. The format used in this list follows essentially that of the 1999 list. Names of Honorary Members are capitalized, those of Emeritus Members (25) are italicized, and Life Mem- bers (28) are distinguished by an asterisk following the date they joined the Society. Dates of election to Honorary or Emeritus status are entered in parentheses. In 1988 Dr. Louise M. Russell was elected Honorary Member. In 1993 Dr. Karl V. Krombein was elected Honorary Member. In 1998 Dr. Louise M. Russell was elected Honorary President and Dr. Ronald W. Hodges was elected Honorary Member. In 1999 Drs. Donald M. Anderson and William E. Bickley were elected Honorary Members. I thank Dianne Mathis for her kind assistance in the preparation of this list. Any corrections to the list can be sent to the Corresponding Secretary at the address on the inside 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 Secretary. Abe, Masaki 1988 JAPAN Abrahamson, W. G. 1997 Pennsylvania Adams, J. R. 1963 Maryland 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 Alpert, G. D. 2002 Massachusetts ANDERSON, D. M. 1954 (1999) Mary- land Anderson, L. D. 1944 (1989) California Arce-Perez, R. 2001 MEXICO Archangelsky, M. 1998 ARGENTINA Arduser, M. S. 1999 Missouri Armitage, B. J. 1983 Ohio Arnaud, P. H., Jr. 1955 California Ascher, J. S. 2000 New York Athanas, M. M. 2001 Maryland Baicher, V. V. 2001 Maryland Baixeras, J. 2001 SPAIN Holly B. Williams, Corresponding Secretary Baker, G. T. 1987 Mississippi Ball, G. E. 1948 Alberta Balogh, G. J. 1994 Michigan Barber, K. N. 1985 Ontario Barnes, J. K. 1978 New York Barrows, E. M. 1976 District of Columbia Baumann, R. W. 1972 Utah Baumgardner, D. E. 1992 Texas Becker, V. O. 1987* BRAZIL Bell, R. T. 1955 Vermont Bellinger, R. G. 1972 South Carolina Berry, R. L. 1972 Ohio Bezark, L. G. 1974 California Bicha, W. 1981 Tennessee BICKLEY, W. E. 1949* (1999) Maryland Bilby, P. J. 1993 New Jersey Bilyj, B. 1998 Ontario Blank, S. M. 2002 GERMANY Blom, P. E. 1986 Pennsylvania Bilyj, B. J. 1998 Ontario Bohart, R. M. 1944 California Borkent, A. 1988 British Columbia 250 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bowles, D. E. 1993 Texas Brailovsky, H. 1996 MEXICO Branham, M. A. 1998 Ohio Braswell, W. E. 2000 New Mexico Broda, S. 1991 Maryland Brodel, C. E 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 Burger, J. EF 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 Campbell, D. 2000 North Carolina Carlson, R. W. 1970* Maryland Carroll, J. EF 1977 Maryland Carroll, L. E. 1997 Maryland Cave, R. D. 1977 HONDURAS Chaboo, C. S. 2001 New York Chavarria, G. 2000 Virginia Choi, S.-W. 2002 SOUTH KOREA Clark, W. E. 1975 Alabama Codella, S. G. 2001 New Jersey Cohen, E. A., Jr. 1999 Maryland Connor, E. E 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. EF 1954 Georgia Currie, D. C. 1999 Ontario 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. Illinois Debboun, M. 2002 Maryland Deeming, J. C. 1974* WALES Deitz, L. L. 1982 North Carolina Dennis, S. 1976 Colorado Dewalt, R. E. 1992 Illinois Deyrup, M. A. 1979 Florida Dodson, B. 1998 Virginia Dorchin, N. 2000 ISRAEL Dozier, H. L. 1952* South Carolina Drummond, R. O. 1954 (1987) Texas Duffield, R. M. 1996 District of Columbia Durkin, P. 1999 District of Columbia Eckerlin, R. P. 1990 Virginia Enns, W. R. 1960 Missouri Epstein, M. E. 1994 District of Columbia Erwin, T. L. 1972 District of Columbia Etnier, D. A. 1999 Tennessee Evans, A. V. 2000 Virginia Evans, H. E. 1948 Colorado Evans, W. G. 1957 (1994) Alberta Evenhuis, N. L. 1980 Hawaii Fales, J. H. 1944 Maryland Fee, E D. 1983 Pennsylvania Ferguson, D. C. 1969 District of Columbia Fisher, E. M. 1977 California Fisk F. W. 1968 (1988) Florida Flint, O. S., Jr. 1961 District of Columbia Flowers, R. W. 1994 Florida Floyd, M. A. 1991 Kentucky Fluno, J. A. 1957 Florida Flynn, D. 2002 North Carolina Foote, B. A. 1958 Virginia Foottit, R. G. 2002 Ontario Foster, G. A. 1999 Maryland Frank, J. H. 1994 Florida Freidberg, A. 1979 ISRAEL Freytag, P. H. 1979 Kentucky Fullerton, S. M. 1999 Florida Furth, D. G. 1994 District of Columbia Gagne, R. J. 1966* Maryland Gaimari, S. D. 1995 California Gelhaus, J. K. 1989 Pennsylvania Gimpel, W. F, Jr. 1995 Maryland Glaser, J. D. 1988 Maryland Goeden, R. D. 1982 California Goldarazena, A. 2001 SPAIN Gordon, R. D. 1968 North Dakota Gordon, S.W. 1998 Maryland Gorham, J. R. 1974 (1995) District of Columbia VOLUME 105, NUMBER | Grace, J. K. 1987 Hawaii Grissell, E. E. 1979 District of Columbia Grogan, W. L. 1997 Maryland Guang-Xue, Z 1999 P. R. OF CHINA Guilbert, E. 2000 FRANCE Haines, K. A. 1952 Virginia Halbert, S. E. 1989 Florida Hall, J. P. 2002 District of Columbia Hamilton, S. W. — Tennessee Hanks, L. M. 1993 Illinois Hanson, P. 1985 COSTA RICA Hansson, C. 1985 SWEDEN Harbach, R. E. 1972 UNITED KINGDOM Harlan, H. J. 1988 Maryland Harman, D. M. 1966 Maryland Harris, S. C. 1979 Pennsylvania Harrison, B. A. 1976 North Carolina Harrison, T. L. 1993 Illinois Hastriter, M. W. 1998 Utah Headrick, D. H. 1992 California 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. FE 1970 District of Columbia Heydon, S. L. 1986 California Heyn, M. W. 2002 Florida Hight, S. 1990 Florida Hilton, D. E J. 1990* Quebec HODGES, R. W. 1960* (1998) Oregon Hoebeke, E. R. 1980 New York Hoffman, K. M. 1986 California Houghton, D. C. 2000 Minnesota Howden, H. FE 1948 Ontario Huang, Y.-M. 1968 District of Columbia Husband, R. W. 1972 Michigan Irwin, M. E. 1976 Illinois Ivie, M. A. 1984 Montana Jashenko, R. V. 1997 KAZAKSTAN Jimenez, H. H. 1994 MEXICO Jimenez-Guarda, P. 1999 CHILE Johnson, E. L. 1995 Washington Johnson, J. B. 1987 Idaho Johnson, N. E 1980 Ohio Joseph, S. R. 1957 Maryland Judd, D. 1994 Oregon 251 Kaster, C. H. 1979 Michigan Keffer, S. L. 1993 Virginia Keiper, J. B. 2000 Ohio Keirans, J. E. 1984 Georgia Kennedy, J. H. 1995 Texas Kim, K. C. 1983 Pennsylvania Kimsey, L. S. 1994 California Kingsolver, J. 1963 (1992) Florida Kirchner, R. EF 1981 West Virginia Kitayama, C. 1974 California Kittle, P. D. 1975 Alabama Kjar, D. S. 2001 District of Columbia Knutson, L. V. 1963* FRANCE Kondratieff, B. C. 1992 Colorado Korch, P. P. 1993 Pennsylvania Korytowski, C. A. 2002 PANAMA Kosztarab, M. 1978 (1994) Virginia Kotrba, M. 1997 GERMANY KROMBEIN, K. V. 1941* (1993) District of Columbia Kruse, J. 2000 Alaska Krysan, J. L. 1993 Kentucky Labandeira, C. C. 1993 District of Columbia Lago, P. K. 1984* Mississippi Lakin, K. R. 1993 North Carolina La Vigne, R. 1999 AUSTRALIA Lee, S. SOUTH KOREA Lein, J. C. 1967 TATWAN Levesque, C. 1985 Quebec Lewis, J. A. 1994 District of Columbia Lewis, P. A. 1974 Ohio Lewis, R. E. 1958 Iowa Lingafelter, S. W. 1997 District of Columbia Lisowski, E. A. 1988 Washington Little, R. G. 1993 California Loechelt, H. K. 1988 Washington Loeffler, C. C. 1992 Pennsylvania Lopez-Martinez, V. 2002 MEXICO MacDonald, J. E 1984 Indiana MacKay, W. P. 1982 Texas Magner, J. M. 1953 Missouri Maier, C. T. 1976 Connecticut Main, A. J., Jr. 1965 New York Manley, D. G. 1984 South Carolina Manglitz, G. R. 1956 (1989) Nebraska to Nn i) Mariluis, J. C. 2000 ARGENTINA Marinoni, R. C. 2002 BRAZIL 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 May, E. 1990 Kansas McCabe, T. L. 1977 New York McGovran, E. R. 1937 (1973) Maryland McKamey, S. H. 1989 District of Columbia McPherson, J. E. 1985 Illinois Mead, F. W. 1976 (1995) Florida Menke, A. S. 1969 Arizona Metzler, E. H. 1998 Ohio Micheli, C. 2001 Maryland Michener, C. D. 1994 Kansas Miller, D. R. 1972 Maryland Miller, G. L. 1981 Maryland Miller, R. S. 1981 Montana Miller, S. E. 1980* District of Columbia Miller, T. D. 1988 Idaho Mitchell, R. T. 1949 (1978) Maryland Moore, T. E. 1950 Michigan Morse, J. C. 1976 South Carolina Morse, L. E. 1999 Virginia Moulton, J. K. 1994 Arizona Mudge, A. D. 2001 Oregon Mullens, B. A. 1999 California Munoz-Quesada, E 1996 Minnesota Munro, J. B. 2000 California Munson, S. C. 1938 (1975) District of Columbia Murray, D. L. 2000 Oregon Nakahara, S. 1968 Maryland Neff, S. E. 1969 Pennsylvania Nelson, C. H. 1969 Tennessee Neunzig, H. H. 1956 North Carolina Nguyen, T. C. 2000 New York Normark, B. 2000 Massachusetts Norrbom, A. L. 1989 District of Columbia Novelo-Guiterrez, R. 1999 MEXICO Nuhn, T. P. 1981 Virginia O’Campo, F. C. 2001 Nebraska Ochoa, R. 1999 Maryland O'Neill, K. M. 2000 Montana Opler, P. A. 2001 Colorado PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Orr, R. L. 1990 Maryland Oswald, J. D. 1987 Texas Packauskas, R. J. 1993 Kansas Painter, J. A. 2001 Virginia Pakaluk, J. 1992* District of Columbia Paras, F 1999 Maryland Parker, C. R. 1977 Tennessee Parrish, D. W. 1963 (1987) Maryland Paulo, O. 1998 BRAZIL Pedrosa-Macedo, J.H. 1999 BRAZIL Pennington, W. 2001 Tennessee Perez, O. G. 2001 Florida Perez-Gelabert, D. E. 2000 Maryland Peters, J. G. 1999 Florida Peterson, R. V. 1952 Utah Pham, V. N. 2001 Maryland Philip, M. 2001 North Carolina Pinto, J. D. 1982 California Pogue, M. 1996 District of Columbia Poiner, G. 1999 Oregon Polhemus, D. A. 1993 District of Columbia Polhemus, J. T. 1964 Colorado Polloni, J. 2001 Massachusetts Porter, C. H. 1984 Georgia Pratt, G. EF 2001 California Pratt, H. D. 1943 Georgia Price, R. D. 1963 Arkansas Pujolluz, J. R. 1998 BRAZIL Pulawski, W. J. 1975 California Quio, G.-X. 1999 P. R. OF CHINA Rabaglia, R. J. 2002. Maryland Rainwater, C. F. 1954 (1975) Maryland Rainwater, H. I. 1964 (1983) Maryland Ramsdale, A. S. Wisconsin Raspi, A. ITALY Rawlins, J. E 1974 Pennsylvania Revol, L. M. 1999 FRANCE Ribeiro-Costa, C. S. 1992 Puerto Rico Richards, A. B. 1996 California Richardson, H. H. 1939 (1976) New Jersey Ridge-O’ Connor, G. E. 2000 Connecticut Riley, D. R. 1984 Texas Robbins, R. G. 1979* Maryland Robbins, R. K. 1986 District of Columbia Robbins, T. O. 1989 Texas Robinson, H. E. 1963 District of Columbia VOLUME 105, NUMBER 1 Root, R. B. 1984 New York Rothschild, M. J. 1989 Maryland Roughley, R. E. 2001 Manitoba Rozen, J. G., Jr. 1956 New York Ruiter, D. 1976 Colorado Rumph, J. A. 1996 Washington RUSSELL, L. M. 1930 (1988) Maryland Santana, F J. 1966 Florida Saugstad, E. S. 1979 West Virginia Scarbrough, A. G. 1971 Maryland Schaefer, C. W. 1985 Connecticut Schang, M. M. 2000 ARGENTINA Schauff, M. E. 1980 District of Columbia Schick, K. 1994 California Schiff, N. M. 1991* Mississippi Schmidt, C. H. 1969 North Dakota Schmude, K. L. 1990 Wisconsin Schubert, R. D. 2001 Maryland Schultz, T. R. 2000 District of Columbia Scudder, G. G. E. 1984 British Columbia Sedlacek, J. D. 1988 Kentucky Setlitf, G. P. 2002 Maryland Shaffer, J. C. 1974 Virginia Shands, W. A. 1940 (1991) South Carolina Shaw, S. R. 1991 Wyoming Shepard, W. D. 1992 California Shorthouse, J. D. 1986 Ontario Silva, M. O. 2002 BRAZIL Silva, V.C. 1999 BRAZIL Simmons, R. A. 2002 District of Columbia Sites, R. W. 1989 Missouri Skarlinsky, T. 2001. Florida Skelley, P. E. 1992 Florida Slater, J. A. 1949 Connecticut Sloan, M. J. 1983 (1990) District of Columbia Smiley, R. L. 1964 Maryland Smith, C. F. 1967 (1987) North Carolina Smith, D. R. 1965* District of Columbia Snelling, R. R. 1972 California Solis, M. A. 1985* District of Columbia Spangler, P. J. 1958* District of Columbia Spilman, R. E. 1950 (1977) Maryland Spinelli, G. R. 1983 ARGENTINA Staines, C. L. 1975 Maryland Starr, C. K. 1987 TRINIDAD & TOBAGO Steck, G. J. 1988 Florida Steiner, W. E. 1979 Maryland 253 Steinly, B. A. 1983 Ohio Stewart, R. D. 1985 Maryland Stibick, J. N. L. 1992* Maryland Stoetzel, M. B. 1971 Maryland Strazanac, J. S. 2001 West Virginia Strickman, D. 1988 Virginia Sutherland, C. M. 1974 New Mexico Sutherland, D. W. S. 1973* Maryland Swann, J. 1999 Ontario Tennessen, K. J. 1982 Alabama Thomas, D. B. 1997 Texas Thompson, F.C. 1968* District of Columbia Thompson, J. V. 1953 (1985) New Jersey Togashi, I. 1983* JAPAN Torres-Miller, L. R. 1984 West Virginia Triplehorn, C. A. 1972 Ohio Turner, W. J. 1983 Washington Ulrich, H. 1978* GERMANY Valley, K. 1976 Pennsylvania Vandenberg, N. — District of Columbia Vazquez, A. W. 1957 Virginia Vockeroth, J. R. 1995 Ontario Voegtlin, D. 1981 [llinois Wahl, D. B. 1987 Florida Wallace, M. S. 2002 North Carolina Wallenmaier, T. E. 1979 Michigan 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, T. A. 2001 Quebec White, G. B. 1977* Maryland Whitfield, J. B. 2002 [linois Whitsel, R. H. 1967 California Whitworth, T. 1999 Washington Williams, G. L. 1984 Maryland Williams, H. B. 1977* District of Columbia Wilson, D. 1999 Mississippi Wilson, N. 1957 Iowa Wilterding, J. 1999 Michigan Wojtowicz, J. A. 1981 Tennessee Wood, D. M. 1987 Ontario 254 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Wood, T. K. 1974 Delaware Zack, R. 1982 Washington Woodley, N. E. 1984* District of Zimmerman, E. C. 1965 AUSTRALIA Columbia Zolnerowich, G. 1987 Kansas Woolley, J. B. 1986 Texas Zuccaro, A. E., Jr. 1986 Mississippi Zungolhi, P. 1978 South Carolina Yasunaga, T. 1998 JAPAN Zuparko, R. L. 1993 California Young, D. K. 1981 Wisconsin Zyla, J. D. 2000 Maryland ENTOMOLOGICAL SOCIETY OF WASHINGTON http//entomology.si.edu/ES W/ESEMenus.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 Subscriptions PROC. ENTOMOL. SOC. WASH. 105(1), 2003, pp. 255-256 INSTRUCTIONS FOR AUTHORS General Policy.—Articles for publica- tion are ordinarily accepted only from members of the Society and must be in En- glish. A summary in French, German, Spanish, or Russian is acceptable. Such a summary should be placed immediately af- ter the English abstract and may be a trans- lation of that abstract. Manuscripts should not exceed 15 printed pages (about two and one-half double-spaced typewritten pages equal one page). Manuscripts are peer-re- viewed before acceptance. Final acceptance is the responsibility of the Editor. Articles are published in the order received, not the date of acceptance. Immediate publication is available with payment of full page charges. Notes, book reviews, and obituar- ies are published as space is available. Manuscript preparation.—Submit three printed copies, including copies of illustra- tions, to the Editor. It is not necessary to send original illustrations until requested af- ter the manuscript has been accepted. After acceptance, the final version may be sub- mitted on diskette (DOS or Macintosh, preferably WordPerfect or Microsoft Word; no ASCII files), but a hard copy is also re- quired. Use letter-sized paper. The entire manuscript should be double spaced. Allow adequate (1—1%") margins on all sides. Words to be italicized may be indicated in italic form or underlined. Manuscript con- tents should be in the following sequence and items 1—6 should begin a new page: 1. Title page with title, author(s), and au- thor(s) byline. In upper left, give name, address, phone, fax, and e-mail of the author to whom all correspon- dence and proofs are to be sent. Titles must include order and family place- MEME Css ie) sre ts, © (Hymenoptera: Xyelidae). Taxa of the genus and spe- cies groups levels must have the au- thority name. 2. Abstract; key words 3. Text of paper; acknowledgments at end 4. Literature Cited (see below) 5. Figure legends 6. Appendix; footnotes (if any) 7. Copies of tables; legends for tables are placed at the top of each table. 8. Copies of figures. Illustrations.—There is no extra charge for line drawings or half tones. Authors should plan illustrations for reduction to the dimensions of the printed page and allow room for legends at the bottom. Do not make plates larger than 14 * 18”. Individ- ual figures should be mounted on suitable board, and photographs (glossy only) should be trimmed, grouped together, and abutted when mounted; the printer will in- sert hairlines. Figure numbers are prefera- bly placed at the lower right of each figure. Arrange plates so there is as little wasted space as possible. Include title of paper, au- thor(s), name and address, and illustration numbers on the back of each plate. Figures should be numbered consecutively and usu- ally in the sequence in which they are re- ferred to in the text. Reference in text should be as “Fig. 1,°* “Bigs: 2) 3,7" ““a- ble 1.”’ See recent Proceedings for style of figure legends. Literature Cited.—Give only papers re- ferred to in the text; list alphabetically by author. Spell out references (journal names etc.) completely, including conjunctions and prepositions—no abbreviations. Do not underline; journal names and book titles are set in Roman type. See recent issues for styles for journals, books, book chapters, etc. When referring to references in text, use the following; Henry (1990), (Henry 1990), (Henry 1987, 1990), (Smith 1990, Henry 1992), (Smith 1990; Henry 1990, 1991), (Miller 1990a, 1990b) where two articles are published by the same author in the same year, or Henry et al. 1990 where more 256 than two authors are involved (do not ital- icize ‘‘et al.”’). Citations **in press’’ should be cited as “‘in press,’> not with year of expected publication. Names of persons providing unpublished information should include initials and be referenced in the text as: W. Mathis (per- sonal communication) [information ob- tained orally] or W. Mathis (in litt.) [infor- mation obtained in a writing]. Names of organisms.—The first men- tion of a plant or animal should include the full scientific name including the authority. Use only common names approved in Com- mon Names of Insects and Related Organ- isms published by the Entomological Soci- ety of America. Spell out the entire scien- tific name the first time it is mentioned, thereafter abbreviate generic names: how- ever, do not abbreviate a genus name at the beginning of a paragraph or sentence, or if two or more genera being discussed have the same first letter. Within sentences, al- ways use the genus name or initial preced- ing a species name. Taxonomic papers.—Taxonomic papers must conform to requirements of the Inter- national Code of Zoological Nomenclature. In addition, type specimens must be desig- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nated for new species described, type de- positories must be clearly indicated, and new taxa must be clearly differentiated from existing taxa by means of keys or dif- ferential diagnoses. In short, these papers must conform to good taxonomic practices. Book Reviews.—Send to Book Review Editor (see inside front cover) and see re- cent Proceedings for format. Charges.—Authors will be assessed a charge to cover editing and publication ex- penses. The current charge to members is $40.00 per printed page. These charges are in addition to those for reprints and author’s correction in proof. The charge for pages in excess of 15 printed pages, for non-mem- bers, and papers for immediate publication is $60.00 per printed page. Member authors who are retired or not affiliated with an in- stitution may request to have charges waived and should do so when the manu- script is submitted. A complete or partial waiver may delay publication. Charges for author errors and/or changes in proof, for reprints if ordered, for immediate publica- tion, and for non-members cannot be waived. Acceptance of papers is based only on scientific merit without regard to the au- thor’s financial support. 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. STRATE CRE MSION 0) ayy WROD) ie ese Se Fe ee Sh ca Pe ee Ee ee A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric (GrisselltandiNiichaclS Schaulis S 7; pp aloo) weer ees ees < hse Se ee ae se Revision of the Oriental Species of Aphthona Chevrolat (Coleoptera: Chrysomelidae), by Alexander S. Konstantinov andSteven) W. Lingafelter., 349)pp. 2002 22 — ee Revision of the Genus Anoplophora (Coleoptera: Cerambycidae), by Steven W. Lingafelter and E. Richardeboebeke.23 Opp. 2002 ees eee eee ee ek ieee ee See ee ae 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 TEL SAGAS O31 O ale), TCS Se acta See sree eae Reece a oe A ST ne ee No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- pogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 _____.--__---------- No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. CH ea NSN et AS ae aE a re Rae ca ens Ace ein Ae Re ee epee ee ed No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 (0) op, ULES Sie ce etal sts eT eel nC he ie seers ROM OS Muppet Regt Oi tt 2 NL Pee SN ees eRe aor SSE No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent SUC le Gropani Talo Spps LOS | ye wt an varee sl eee Sh RT a) Tee oe Coe Os Sees) No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 __. No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited by BaBiNordenandvA]SsMenkes216'pp) 1996) Se ee ee ee eee 25.00 No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and Wilbamil: Grocanmir29dipn 1997 sae a ee ee eee ee No. 19. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by honiasrnMennyb49) np OO hr Sok es oe See et PE AE eS SS 2 ey No. 20. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- Pate ere alsa yp) Lo 9 openers tase arate cee ge ee ee ees ek ee ee eee No. 21. New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. [Pretgdars WAM O)fo) oO eS SONORA go Pe ot ee ee eee re eee ee Be eS No. 22. Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: Trichogrammatidae), by John D. Pinto. 287 pp. 1999 ______. No. 23. Revision of the Net-Winged Midges of the Genus Blepharicera Macquart (Diptera: Blepha- riceridae) of Eastern North America, by Gregory W. Courtney. 99 pp. 2000 ____-_-_.------------------ No. 24 Holcocerini of Costa Rica (Lepidoptera: Gelechioidea: Coleophoridae: Blastobasinae), by David Adamskaenlayinip 2200 2 amas ae oe eet RS 18.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume to non-members and $25.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage and handling charges are extra. Dealers are allowed a discount of 10 percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Treasurer, Entomological Society of Washington, “ Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. CONTENTS (Continued from front cover) LARSON, KRISTIN ANN and DAN MYERS HARMAN —Subcortical cavity dimension and inquilines of the larval locust borer (Coleoptera: Cerambycidae) .................-.+-.-.+45- LUIS-MARTINEZ, ARMANDO, JORGE LLORENTE-BOUSQUETS, ISABEL VARGAS- FERNANDEZ, and ANDREW D. WARREN—Biodiversity and biogeography of Mexican - butterflies (Lepidoptera: Papilionoidea and Hesperioidea) ...................2..02.02200200. MARSHALL, S. A. and W. K. REEVES—A new cavernicolous Spelobia Spuler (Diptera: Sphaeroceridae: Limosininae) from the eastern United States ........................0000000- MATHIS, WAYNE N. and MICHELLE D. TRAUTWEIN—-A revision and phylogenetic study of Iipochaera Coquillets (Dipterarse phy dnidae)) ec... cariine ticle eee elses eietert tire NICKLE, DAVID A.—A checklist of commonly intercepted thrips (Thysanoptera) from Europe, the Mediterranean, and Africa at U.S. ports-of-entry (1983-1999). Part 1. Key to genera .... NOYES, JOHN S. and MICHAEL E. SCHAUFF—New Encyrtidae (Hymenoptera) from papaya mealybug (Paracoccus marginatus Williams and Granara de Willink) (Hemiptera: Sternor- datyaalel eCtad PLfei fe fofote(o%e GEVS) tba Bara ssacar cca asAnadosoccsbcriee 4 30st ardor nUng ao lstsean 4 adGueaee Bon POGUE, MICHAEL G.—New synonyms in the genus Obrima Walker (Lepidoptera: Noctuidae), Withiadditionall distribution records): h yee emer eack et eyelets near ise inet tee bies cit era ireee rte PRATHAPAN, K. D. and A. S. KONSTANTINOV—The flea beetle genus Aphthona Chevrolat (Coleoptera: Chrysomelidae) of southern India, with descriptions of seven new species .... RICKSON, FRED R., MELINDA M. RICKSON, KUMAR GHORPADE, BETH B. NORDEN, and KARL V. KROMBEIN—Invertebrate biodiversity (ants, bees, and others) associated with stem domatia of the Indian myrmecophyte Humboldtia brunonis Wallich (Magnolio- Phliy tas Fabaceae) uss den samectatessspreth na aide Oiinia lis At ciaera tists fetes aire me sperseintetee iceel ac op fates eta eee SANCHEZ-PENA, SERGIO R., DONALD R. DAVIS, and ULRICH G. MUELLER—A gregar- ious, mycophagous, myrmecophilous moth, Amydria anceps Walsingham (Lepidoptera: Acrolophidae), living in Atta mexicana (F. Smith) (Hymenoptera: Formicidae) spent fungal GultTe accu MULAL ONS iia.'8 ste. tosis ep ha wae eicletaielatelel cceicrsiel eh tet otto he? AME ects oietercl ieee eats ett te ea SHAFFER, JAY C.—Peoria insularis, a new species of Peoriini (Lepidoptera: Pyralidae: Phyci- tinae’)) from Mississtppivand Wowisianial 2x. tars aletoicinic desielatelsacceeienscs scctes 397 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2003 JONATHAN R. Mawps ey, President MIcHAEL G. PoGuE, Treasurer E. E. GrissELL, President-Elect JoHN W. Brown AND Davip G. Furtu, Program Chairs Stuart H. McKamey, Recording Secretary MicuHaet W. Gates, Membership Chair Ho tis B. WILLIAMS, Corresponding Secretary GABRIELA CHAVARRIA, Past President Jon A. Lewis, Custodian Davin R. Smitn, Editor Publications Committee Raymonpb J. GAGNE THomMas J. HENRY Wayne N. Maruis Honorary President DonALp M. ANDERSON Honorary Members WILLIAM E. BICKLEY Kart V. KROMBEIN RONALD W. HopGeEs Pau 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. 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. 522-523 OF THIS ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 31 March 2003 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. 105(2), 2003, pp. 257-264 PLAGIOGNATHUS REINHARDI JOHNSTON (HEMIPTERA: MIRIDAE): DISTRIBUTION, HABITS, AND SEASONALITY OF A HAWTHORN (CRATAEGUS) SPECIALIST A. G. WHEELER, JR. Department of Entomology, Clemson 29634, U.S.A. (e-mail: awhlr@clemson.edu) University, Clemson, SC Abstract.—The phyline plant bug Plagiognathus reinhardi Johnston, described from Texas in 1935, has been known only from the type locality (College Station). Recent fieldwork produced the first records of this species for the southeastern United States. This mirid developed on glandular hawthorns (Crataegus sp.[p.]; Rosaceae) of series Lachrimatae, mainly in the fall-line sandhills from southern North Carolina through South Carolina and Georgia to eastern Alabama but also in disturbed sandhills in the panhandle and northern peninsula of Florida. Overwintered eggs hatched near vegetative budbreak of host hawthorns: late February to early March in Florida and mid-March in South Carolina. Early instars were found on the expanding leaves, whereas late instars were associated mainly with flower buds and inflorescences. Adults appeared by mid- or late March in Florida and mid-April in South Carolina. Adults of this univoltine mirid were present for about four to six weeks, the females persisting longer than the males. Key Words: Insecta, Miridae, insect distribution, seasonal history, Crataegus, fall-line sandhills Johnston (1935) described the phyline plant bug Plagiognathus reinhardi from College Station, Texas. The type series con- sisted of 80 males and females collected on hawthorn, Crataegus sp. (Rosaceae), from March 29 to April 21, 1933. No further re- cords of this distinctive dark red or reddish- brown mirid have been published since the original description (Henry and Wheeler 1988). Schuh (2001) stated that it is found in the southern United States but did not cite additional records. This plant bug’s sexually dimorphic second antennomere and structure of the male vesica (see Schuh [2001]: fig. 40) do not conform to the cur- rent concept of the genus Plagiognathus. This species, therefore, was given the status of incertae sedis by Schuh (2001). Here I record P. reinhardi from the southeastern United States, cite an additional Texas re- cord, and provide notes on its seasonal his- tory and habits. METHODS AND STUDY SITES I discovered P. reinhardi in South Car- olina in 1988, but surveys to learn more about its southeastern distribution were not initiated until 1999. Crataegus spp. in the fall-line sandhills from southern North Car- olina to eastern Alabama were sampled dur- ing 1999-2001 by tapping branches over a shallow net. Sampling also was conducted in northern Florida in 2000—2002. At each site where adults were found, one or more specimens were collected and deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. During March—May 2000-2001, haw- 258 thorns in the fall-line sandhills of South Carolina were sampled to determine when the overwintered eggs of P. reinhardi begin to hatch, when the adults begin to appear, and the period of time they are present. The main study site was a disturbed sandhill in Richland County, 2.5 km southwest of Pon- tiac. The approximate time of egg hatch and the period of adult occurrence also were de- termined at several sites in the northern peninsula of Florida (see Distribution for specific localities). The number of individ- uals observed on each sample date varied but generally was <20. Early instars were particularly difficult to find because they were not easily dislodged from partially closed vegetative buds. Only lmited num- bers (<10) of nymphs typically were ob- served when overwintered eggs were be- ginning to hatch. First and second instars were collected, placed in 70% ethanol, and sorted to stage in the laboratory using a bin- ocular microscope; later instars were iden- tified in the field and their relative propor- tions estimated. DISTRIBUTION (Fig. 1) In the following list of new records for P. reinhardi, those based only on nymphs are indicated as “*(n).”* All collections were from Crataegus (for comments on identifi- cation of the host, see Habitats and Host Plants) by the author, except the one from Texas and those from Washington County, Florida. ALABAMA: Russell Co., Rt. 165, 5 km N of Fort Mitchell, 32°23.1'N, 85°02.2'W, 9 Apr. 2000 (3 6, 1 2); Rt. 165, 4 km N of Holy Trinity, 32°15.5’N, 85°00.1'W, 9 Apr. 2000 (n); Rt. 169, 1.5 km NW jet. Rt. 431, NNW of Seale, 32°19.6’N, 85°10.5'W, 9 Apr. 2000 (n). FLORIDA: Alachua Co., Rt. 441, 1.7 km NW of High Springs, 29°50.2'N, 82°36.1'W, 3 Mar. 2000 (n) and 22 Mar. 2000 (6 3,5 ¢), 19 Mar. 2002 (n); Columbia Co., Ichetucknee Springs State Park, ca. 6.5 km NW of Fort White, 29°57.5'N, 82°46.0'W, 3 Mar. 2000 (n) and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 22 Mar. 2000 (20 3, 6 @), 2 Mar. 2001 (n) and 17 Mar. 2001 (10 6, 2 2), 19 Mar. 2002 (n); Hamilton Co., Rt. 129, ca. 4 km NW of Jasper, 30°32.6'N, 82°58.4'W, 23 Mar. 2000 (n); Rt. 129, 0.7 km N of Su- wannee River, NE of Suwannee Springs, 30°23.8'N, 82°56.1'W, 23 Mar. 2000 (n); Suwannee Co., Rt. 27, 5 km E of Branford, 29°57.1'N, 82°52.6’W, 3 Mar. 2000 (n); jct. Rts. 75 & 136, 4.6 km SW of White Springs, 30°19.2'N, 82°48.4'W, 21 Mar. 2000 (n); Washington Co., Rt. 279, 7 km N of Greenhead & jct. Rt. 77, 30°34.6'N, 85°40.1'W, 19 Apr. 2001, T. J. Henry & A. Gs Wheeler Jri( oe 1 2) RE793 2:4 kmn S of New Hope, 30°33'17"N, 85°49'14"W, 19 Apr. 2001, T. J. Henry & A. G. Wheeler, Jr. (1 3). GEORGIA: Chattahoochee Co., Rt. 26, 1 km N of Cusseta, 32°18.9'N, 84°46.4'W, 9 Apr. 2000 (3 3, 1 &); Craw- ford Co., Rt. 96, 0.5 km E of Flint River, 8.2 km E of Reynolds, 32°32.8'N; 84°00.7'W, 9 Apr. 2000 (n); Crisp Co., Rt. 257, 1.3 km SW of Dooly Co. line, 0.8 km SW of Lambtown, 32°01.3'N, 83°5.5'W, 21 Apr. 2001 (2 2); Glascock Co., Rt. 123, 0.1 km NNW of Mitchell, 33°13.2'N, 82°42.3'W, 18 May 1999 (1 @); Jefferson Go., Rt. 221, 9:8) kim! IN) ‘of Wrens? 33°18.7'N, 82°22.7'W, 18 May 1999 (4 gd, 8 2); Jones’ Gor, Rt 129)°3/87km) NEtor Gray, 33°02.4'N, 83°30.7’W, 9 Apr. 2000 (n); McDuffie Co., CR-86 nr jet. CR SE 75, 25 km; Wo jof Deanne, 33225-5uNe 82°24.6'W, 18 May 1999 (3 6, 3 @); EI- lington Airline Rd., 3 km SW of Dearing, 33°24.1’N, 82°24.9'W, 1 Apr. 2000 (n); Marion Co., jct. Rts. 41 & 127, 14 km N of Buena Vista, 32°26.4’N, 84°32.1'W, 8 Apr. 1997 (adults, sex unknown) and 9 Apr. 2000 (n); Oconee Co., Rt. 129, 0.5 km SW of Farmington, 33°46.4’N, 83°25.6’W, 9 Apr. (n) & 20 Apr. 2000 (n, 3 d reared); Pulaski Co., Rt. 129A, Hartford community NE of Hawkinsville, 32°17.1'N, 83°26.9'W, 21 Apr. 2001 (3 6, 8 &); Richmond Co., Wheeler Rd., Augusta, 33°28.9'N, 82°05.2'W, 18 May 1999 (2 2); Stewart Co., Rt. 39, 4 km E of Omaha, 32°08.3'N, VOLUME 105, NUMBER 2 Fig. 1. New distribution records of Plagiognathus reinhardi in the southeastern United States (Alabama, Florida, Georgia, North Carolina, South Carolina). Note: See Distribution for more precise locations of collection sites and for a new Texas record. 84°58.3'W, 9 Apr. 2000 (n); Talbot Co., Rt. S05 Se fof. Munction tG@itys7 32°35.14N; 84°26.8'W, 15 Apr. 1990 (n); Rt. 96, Junc- tion City, 32°36.3'N, 84°27.7’'W, 9 Apr. 2000 (3 6, 1 2); Warren Co., Rt. 123, 3.6 km S of jet. Rt. 16, SSW of Warrenton, 33°16.0'N, 82°45.2’W, 18 May 1999 (2 6, 15 2): Wilkinson Co., Rt. 18, 6.1 km S of Gordon, 32°50.4'N, 83°20.0'W, 21 Apr. 2001 (5 6, 4 2). NORTH CAROLINA: Bladen Co., Rt. 242, 14 km N of Eliza- bethtown, 34°45.9'N, 78°36.4’W, 23 May 260 1999 (1 56,3 ¢); Cumberland Co., Rt. 242, 1.9 km S of jet. 210, 4.5 km NNE of Am- mon, 34°22.0'N, 78°33.1’W, 23 May 1999 (1 6,4 @); Richmond Co., Rt. 177, 1.1 km NE of Hamlet, 34°54.4'N, 79°41.1'W, 22 May 1999 6, 17 2); Rt. 177, 0.3 km S$ of Osborne, 34°45.5'N, 79°46.2'W, 23 May 1999 (3 2). SOUTH CAROLINA: Aiken Co., Henderson Heritage Preserve, ca. 6 km NW of Aiken, 33°36.3’N, 81°45.2’W, 24 Apr. 1998 (1 2) & 16 Apr. 2000 (n); North Augusta, 33°30.3'N, 81°57.9'W, 11 Mar. & 7 Apr. 2001 (n); Bamberg Co., Rt. 321, 3.8 km N of Denmark, 33°21.6'N, 81°08.2'W, 31 Mar. 2001 and 7 Apr. 2001 (n); Barnwell Gos Rt 39525: km SSE*of ject RE 787SE of Williston, 33°23.2'’N, 81°26.7W, 31 Mar. 2001 (n); Rt. 78, 6.4 km W of Blackville, 33°22.3'N, 81°20.3’W, 7 Apr. 2001 (n); Cal- houn Co., co. rd. 31 E of jct. Rt. 26, ca. 4 km NW of Sandy Run, 33°48.6'N, 81°00.4'W, 20 May 2001 (1 2); Rt. 176, 5 km) SSEs of "Sandy ) Runs 33°46:37Ne 80°55.5'W, 20 May 2000 (1 ¢); Chester- field Co., jet. Rts. | & SR-13-29, 4.3 km NE of Middendorf, 34°32.6'N, 80°07.4'W, 22 May 1999 (1 6, 8 @&); Edgefield Co., S-19-407 nr. ject. S-19—37, ca. 8.5 km SW of Eureka, 33°39.5'N, 81°51.6'W, 18 May 1999 (1 6, 15 2), 6 June 1999 (1 GY), 12 May 2001 (1 6,5 Q); jet. Rt. 25 & Whit- lock Rd., 8.8 km SSW _ of Trenton, 33°40.0'N, 81°52.4'W, 18 May 1999 (1 gd, 3 2); Fairfield Co., Rt. 34, 0.6 km W of Kershaw Co. line, 12.6 km SSE of Ridge- way, 34°16.4'N, 80°49.9'W, 20 May 2000 (1 6, 2 2); Kershaw Co., jct. S-28-47 & S=28=367, -cayi3.8) km’ “SE of JElein, 34°09.1'N, 80°45.7’W, 22 May 1999 (2 6, 8 92); Rt. 1, ca. 8.5 km NE of Camden, 34°18.8'N, 80°32.4’W, 22 May 1999 (2 @); Lexington) Co; Rt, 302--S of (Cayce; 13 May 1988 (1 @); jct. Rt. 178 and Gantt Mill Rd. 75 km W" off Pelion,233-45.3..N, 81°19.7'W, 6 May 1989 (2 adults, sex un- known); Rt. 321, 0.8 km S of jet. S-32-952, 33°48.4'N, 81°05.8'W, Gaston, 11 & 31 Mar. 2001 (n); Rt. 321, 3 km N of Gaston, 33°50.3'N, 81°05.3'W, 26 Apr. 1998 (1 6, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1 2); Peachtree Rock Preserve, 3.5 km SE of Edmund, 33°49.9'N, 81°12.0'W, 26 May 1995 (2 2), 7 Apr. (n) & 28 Apr. 2001 (4 3,1 2); Marlboro Co., Rt. 177, 0.9 km S of Fulton, 34°46.2'N, 79°47.7'W, 23 May 1999 (2 6, 6 2); Richland Co., Valhalla Dr., 0.1 km S of Rt. 1, 2.5 km SW of Pon- tiac, 34°07.0'N, 80°52.8’W, 15 May (9 6, 6 2), 22 May (1 6, 10 2) & 26 May 1999 (2 2); 16 Apr. (2 2), 6 May (4 6,6 &), & 20 May 2000 (10 2); Sumter Co., Rt. 261, Manchester State Forest, 4.7 km S of Wed- gefield, 33°51.5'N, 80°31.0'W, 26 May 1999 (1 @). TEXAS: Victoria Co., Rt. 59, 9 km NE of Inez, 28°56.5'N, 96°42.5'W, 18 Apr. 19835) i Jp Henny ce A. G? Wheeler wr (adults, sex unknown). HABITATS AND Host PLANTS Plagiognathus reinhardi was found mainly in the fall-line sandhills from south- ern North Carolina through South Carolina and Georgia to eastern Alabama, and in xe- ric sand communities in northern Florida. In South Carolina counties such as Edge- field and Fairfield, which lie mainly in the piedmont and contain a sliver of sandhills, this mirid was found only in the sandhills ecoregion. In Georgia, however, P. rein- hardi was collected in the piedmont as far north as Oconee County (Fig. 1). The host plant of P. reinhardi is a species of Crataegus (or complex of ““microspe- cies” sensu Phipps 1988b) characteristic of xeric sand communities from southern North Carolina to eastern Alabama and northern Florida. This plant often 1s found in disturbed sites such as vacant lots, along highways, and railroad rights-of-way, and it is the most common hawthorn in the sand- hills of South Carolina (Schoenike 1982). Belonging to the section (and series) Lac- rimatae (Phipps et al. 1990), this taxon of- ten is misidentified as C. flava Ait. (e.g., Duke 1961, Radford et al. 1968, Clark 1971, Clewell 1985, Godfrey 1988, Par- tridge et al. 2000; see also Phipps et al. 1990). Crataegus flava, however, is rare or VOLUME 105, NUMBER 2 261 Fig. 2. Host plant of Plagiognathus reinhardi, a hawthorn of series Lachrimatae (probably Crataegus ala- bamensis or C. meridiana), in disturbed sandhills near Pontiac (Richland County), South Carolina. even extinct in the wild, apparently is no longer in cultivation, belongs to another section of Crataegus, and has not been cor- rectly identified subsequent to Beadle’s (1903) treatment (Phipps 1988a, Phipps et al. 1990). In addition to C. flava sensu auctt. Amerr. non Ait., the common Cra- taegus species of southeastern sandhills has been referred to as C. meridiana Beadle (Coker and Totten 1945) and C. michauxii Pers. (e.g., Thorne 1954, Weakley 1997). Specific identifications in Lacrimatae, as in many other sections of Crataegus, are prob- lematic. In many cases the original descrip- tions not only are insufficiently diagnostic to discriminate species, but type specimens, including those for taxa named from the southeastern states by C. D. Beadle and oth- ers, often are in poor condition or lacking. Apomixis and hybridization, as well as polyploidy, contribute to the “Crataegus problem” (e.g., Phipps 1988b, Phipps et al. 1990, Lance 1995). The host of P. reinhardi at the sample site in Richland County, South Carolina, might be C. alabamensis Beadle or C. mer- idiana (Fig. 2). This plant is a shrub or small tree with a dense growth habit, pen- dulous branches at maturity, short and stout thorns, twigs generally zigzag at the nodes, small leaves with their margins conspicu- ously gland dotted and white-tomentose above when young, petioles glandular; in- florescences compact and few flowered (generally 4—7 in cymes) with flowers small and the calyx lobes usually glandular-punc- tate, stamens 20; and fruit oval, yellow to red in clusters of 1—3, nutlets 2—5. Speci- mens from the main study site near Pontiac, South Carolina, labeled Crataegus aff. al- abamensis Beadle by J. B. Phipps, 17 July 2000, and annotated to C. meridiana Beadle by R. Lance, 5 June 2002, are in the Clem- son University Herbarium (CLEMS 55280; Townsend #2200). The morphologically similar hosts of P. reinhardi in Alabama, Florida, Georgia, North Carolina, and at other sites in the South Carolina sandhills also belong to se- ries Lachrimatae but may or may not be conspecific with what is thought to be C. alabamensis or C. meridiana in Richland County, South Carolina. Material from sample sites in Florida will key to C. mu- chauxii in Godfrey (1988) and Wunderlin (1998). Other plants found consistently with C. alabamensis or C. meridiana (or other species of Lachrimatae) in disturbed 262 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sandhills of South Carolina were American joint-weed, Polygonella americana (Fisch. & Mey.) Small (Polygonaceae); eastern prickly-pear, Opuntia humifusa (Raf.) Raf. (Cactaceae); longleaf pine, Pinus palustris Mill. (Pinaceae); silk-grass, Pityopsis gra- minifolia (Michx.) Nutt. (Asteraceae); and turkey oak, Quercus laevis Walter (Faga- ceae). SEASONALITY AND HABITS In the South Carolina sandhills near Pon- tiac (Richland Co.), first instars were ob- served on 12 March 2000. No nymphs had been found on 26 February when vegetative buds were still tightly closed. Collections on | April consisted of one first, two third, two fourth, and two fifth instars. Popula- tions on 16 April contained mostly fifth in- stars, with fourth instars and teneral adults also present. Only adults (4 ¢, 6 2) were found on 6 May, and only adult females (n = 10) on 20 May. The latest record of P. reinhardi in South Carolina, a female from Edgefield County, was 6 June 1999. In 2001, collections in South Carolina on 11 March consisted only of first instars at North Augusta (Aiken Co.) and Gaston (Lexington Co.). Second through fourth in- stars (predominately thirds) were observed at Gaston on 31 March, and fourth instars were collected on that date near Denmark in Bamberg County. Collections on 31 March in Aiken, Bamberg, and Barnwell counties consisted of third through fifth in- stars. On 9 April 2000, adults (3 d, 1 2) were observed at one of three sites (north of Fort Mitchell) that were sampled in Russell County, Alabama, in the East Gulf Coastal Plain. Populations at all three Alabama sites consisted mainly of fourth and fifth instars, although a few second and third instars were present near Seale and a few third in- stars at the site north of Holy Trinity. Mirid populations in the East Gulf Coastal Plain of Georgia also consisted mainly of late in- stars and smaller numbers of teneral adults on 9 April 2000. The populations of P. reinhardi sampled in northern Florida (Alachua, Columbia, and Suwannee counties) consisted of first and second instars on 3 March 2000. At Ichetucknee Springs State Park in Columbia County, adults (20 6, 6 2), many of them teneral, were present on 22 March with about equal numbers of late (fourth and fifth) instars. In 2001, development of this mirid on the single hawthorn that had been sampled the previous March in Ichetucknee Springs State Park was advanced compared to 2000; second through fourth instars were found on 2 March, indicating that egg hatch had begun by late February (no nymphs, however, were found on this same plant on 22 and 26 February 2002). Collections from this same plant on 17 March 2001 indicated that fifth instars were most numerous, with smaller numbers of fourth instars and most- ly teneral adults (10 d, 2 @) also present. The population of P. reinhardi at Ichetuck- nee Springs State Park developed later in 2002 than in either of the two previous sea- sons; second through fourth instars were observed on 19 March. Extensive sampling on 14 April 2001 did not yield adults at the Alachua County site near High Springs, but small numbers of adults were still present in the panhandle (Washington Co.) on 19 April 2001. First and second instars were found with- in unfolding leaves and presumably feed on the young foliage. Third through fifth in- stars were associated mainly with the flower buds and inflorescences of host hawthorns. DISCUSSION The phyline plant bug P. reinhardi, known previously only from the type lo- cality in Texas, can be added to the fauna of the southeastern United States. A char- acteristic insect of the fall-line sandhills of Georgia, North Carolina, South Carolina, and eastern Alabama, it often is found in disturbed sites within pine-scrub oak sand- hill communities. This mirid also can be found in disturbed sandhills in xeric sand communities of northern Florida. Arthro- VOLUME 105, NUMBER 2 pods associated with xeric longleaf pine habitats, which include sandhills, tend to be poorly known (Folkerts et al. 1993). The South Carolina sandhills, important for their biotic richness, provide habitats for numerous plant and animal species that are considered rare or endangered (e.g., Pitt- man 2001). Plagiognathus reinhardi is a hawthorn specialist. The host plant in South Carolina apparently 1s Crataegus alabamensis or C. meridiana, a hawthorn with glandular leaf margins, petioles, and calyx lobes. This plant frequently is misidentified as C. flava Ait. Elsewhere, P. reinhardi also develops on glandular hawthorns of series Lachri- matae. The use of glandular host plants is common in the Miridae (see Wheeler 2001). Plagiognathus reinhardi was not found on nonglandular species of Cratae- gus, including C. spathulata Michx. grow- ing in Oconee County, Georgia, within two meters of a glandular hawthorn that har- bored nymphs of the mirid. Overwintered eggs of this univoltine plant bug begin to hatch at or slightly after vegetative budbreak of their hosts. Egg hatch apparently begins about the second week of March in the South Carolina sand- hills and, in some years, as early as late February in northern Florida. In South Car- olina, adults appear by mid-April, with fifth instars sometimes present until late April. These nymphal “stragglers” usually are parasitized by a euphorine braconid. Adults begin to appear in northern Florida by about mid- to late March. The appearance of adults generally corresponds with the pe- riod of late bloom on host hawthorns. Adults are present only for four to six weeks. Late-season populations, like those of most Miridae (Wheeler 2001), are strongly female biased. The only other phytophagous mirid that co-occurred with P. reinhardi on glandular hawthorns was the orthotyline Heterocor- dylus malinus Slingerland. It was observed at about half of the sites sampled in both Georgia and South Carolina and was pre- 263 sent at one of three sites sampled in Russell County, Alabama (new state record). Her- erocordylus malinus is a univoltine plant bug whose seasonality in Alabama, Geor- gia, and South Carolina is similar to that of P. reinhardi, the overwintered eggs hatch- ing by mid-March and adults appearing about a month later. Nymphs of H. malinus are red to reddish brown, whereas those of P. reinhardi are yellow to yellow green. ACKNOWLEDGMENTS I am grateful to J. B. Phipps (University of Western Ontario, London) for identifying the hawthorn from Richland County, South Carolina; J. F Townsend (formerly with Clemson University) for identifying several plant species from disturbed sandhills, pre- paring herbarium specimens of Crataegus, and identifying C. spathulata from Georgia; T. J. Henry (Systematic Entomology Lab- oratory, USDA, % National Museum of Natural History, Washington, D.C.) for ver- ifying the identification of P. reinhardi and for his companionship in the field in Florida and Texas; S. H. Bennett (South Carolina Department of Natural Resources, Colum- bia) for issuing a permit that allowed me to collect insects in the Henderson Heritage Preserve; D. C. Soblo (formerly with the South Carolina Nature Conservancy, Co- lumbia) for allowing me to collect in the Peachtree Rock Preserve; G. J. Steck (Flor- ida Department of Agriculture and Con- sumer Services, Gainesville) and the Divi- sion of Recreation and Parks, Florida De- partment of Environmental Protection, Tal- lahassee, for the permit authorizing the collection of insects in Ichetucknee Springs State Park: G. R. Carner (Clemson Univer- sity) for photographing the host plant, and P. H. Adler (Clemson University) for offer- ing suggestions that improved the manu- script. LITERATURE CITED Beadle, C. D. 1903. Crataegus, pp. 532-569. In Small, J. K. Flora of the Southeastern United States. The author, New York. 2604 Clark, R. C. 1971. The woody plants of Alabama. An- nals of the Missouri Botanical Garden 58: 99— 242. Clewell, A. E 1985. Guide to the Vascular Plants of the Florida Panhandle. University Presses of Flor- ida, Tallahassee, 605 pp. Coker, W. C. and H. R. Totten. 1945. Trees of the Southeastern United States, Including Virginia, North Carolina, South Carolina, Tennessee, Geor- gia, and Northern Florida. University of North Carolina Press, Chapel Hill, 419 pp. Duke, J. A. 1961. The psammophytes of the Carolina fall-line sandhills. Journal of the Elisha Mitchell Scientific Society 77: 3-25. Folkerts, G. W., M. A. Deyrup, and D. C. Sisson. 1993. Arthropods associated with xeric longleaf pine habitats in the southeastern United States: A brief overview, pp. 159-192. /n Proceedings of the Tall Timbers Fire Ecology Conference No. 18, The Longleaf Pine Ecosystem: Ecology, Restoration and Management, May 30—June 2, 1991. Tall Timbers Research Station, Tallahassee, Florida. Godfrey, R. K. 1988. Trees, Shrubs, and Woody Vines of Northern Florida and Adjacent Georgia and Al- abama. University of Georgia Press, Athens, 734 PP- Henry, T. J. and A. G. Wheeler, Jr. 1988. Family Mir- idae Hahn, 1833 (= Capsidae Burmeister, 1835): The plant bugs, pp. 251-507. Jn Henry, T. J. and R. C. Froeschner, eds. Catalog of the Heteroptera, or True Bugs, of Canada and the Continental Unit- ed States. E.J. Brill, Leiden. Johnston, H. G. 1935. Five new species of Miridae (Hemiptera). Bulletin of the Brooklyn Entomolog- ical Society 30: 15—19. Lance, R. 1995. The Hawthorns of the Southeastern United States. The author, Fletcher, North Caro- lina, 136 pp. Partridge, E. C., W. A. Baker, and G. Morgan-Jones. 2000. Notes on Hyphomycetes. LXXVII. A new species of Ramichloridium, R. bacillisporum, oc- curring on leaf glands of Crataegus flava in Ala- bama. Mycotaxon 75: 147-152. Phipps, J. B. 1988a. Re-assessment of Crataegus flava PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Aiton and its nomenclatural implications for the Crataegus serial name flavae (Loud.) Rehd. and its sectional equivalent. Taxon 37: 108—113. . 1988b. Crataegus (Maloideae, Rosaceae) of the southeastern United States, I. Introduction and series Aestivales. Journal of the Arnold Arboretum 69: 401-431. Phipps J. B., K. R. Robertson, P. G. Smith, and J. R. Rohrer. 1990. A checklist of the subfamily Ma- loideae (Rosaceae). Canadian Journal of Botany 68: 2209-2269. Pittman, B. 2001. Sandhills fall line, pp. 21—25. In Carswell, L. and P. Robinson, eds. Conserving South Carolina: Sustaining and Protecting our Natural Resources. The Nature Conservancy, Co- lumbia, South Carolina. Radford, A. E., H. E. Ahles, and C. R. Bell. 1968. Manual of the Vascular Flora of the Carolinas. University of North Carolina Press, Chapel Hill, 1183 pp. Schoenike, R. E. 1982. A Dendrology of South Car- olina Trees, Native and Introduced, 3rd printing. Department of Forestry, Clemson University, Clemson, South Carolina, 288 pp. [unpublished manuscript, Cooper Library, Clemson University. ] Schuh, R. T. 2001. Revision of New World Plagiog- nathus Fieber, with comments on the Palearctic fauna and the description of a new genus (Heter- optera: Miridae: Phylinae). Bulletin of the Amer- ican Museum of Natural History 266: 1—267. Thorne, R. E 1954. The vascular plants of southwest- ern Georgia. American Midland Naturalist 52: 257-327. Weakley, A. S. 1997. Flora of the Carolinas and Vir- ginia: Working Draft of 21 July 1997. The Nature Conservancy, Southeast Regional Office, Chapel Hill, North Carolina. Wheeler, A. G., Jr. 2001. Biology of the Plant Bugs (Hemiptera: Miridae): Pests, Predators, Opportun- ists. Cornell University Press, Ithaca, New York, 507 pp. Wunderlin, R. P. 1998. Guide to the Vascular Plants of Florida. University Press of Florida, Gainesville, 806 pp. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 265-282 REVISION OF DESMOMETOPA LOEW (LITOMETOPA SABROSKY) (DIPTERA: MILICHITDAE), WITH DESCRIPTIONS OF SIX NEW SPECIES I. BRAKE AND A. FREIDBERG (IB) Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany (e-mail: 1.brake.zfmk @ uni-bonn.de); (AF) Department of Zo- ology, The George S. Wise faculty of Life Sciences, Tel Aviv University, Tel Aviv 69978, Israel (e-mail: afdipter@ post.tau.ac.il) Abstract.—The Afrotropical genus Litometopa Sabrosky is relegated to subgeneric sta- tus within Desmometopa Loew and revised. Desmometopa (L.) glabrifrons (Sabrosky), the type species, is redescribed, and six new species, D. (L.) brachycephala, D. (L.) dolichocephala, D. (L.) flavicornis, D. (L.) glandulifera, D. (L.) nigrifemorata, and D. (L.) sabroskyi, are described. The subgenus is redescribed, including a description of sclerotized structures within the male abdomen possibly representing a gland, and a spe- cialization of the female ovipositor, known as the secondary ovipositor. A key to all species is given. The relationship between Litometopa and other taxa within Desmometopa and among species of Litometopa is discussed. Key Words: In 1965 Sabrosky established the genus Litometopa for his new species, L. glabri- frons Sabrosky, based on four specimens collected in Tanzania. The genus was not recorded again until Brake (2000) discussed it in her study of the Milichiids. Although museum specimens are relatively scarce, about 130 recently have become available to us, including six new species, which are described herein. In his description of Litometopa, Sabros- ky stated that Litometopa is closely related to Desmometopa Loew and wrote: “In a sense, it [Litometopa] is an extreme form of the latter [Desmometopa], with interfrontal hairs and stripes absent, and orbital bristles and mesonotal hairs greatly reduced.” In Sabrosky (1983) recorded several polished black spe- cies, which were intermediate between Des- mometopa and Litometopa in that they shared most (plesiomorphic) characters of his revision of Desmometopa, Diptera, Milichiidae, Desmometopa (Litometopa), new status, Afrotropical Desmometopa but lacked the (apomorphic) interfrontal stripes. He mentioned the pos- sibility that these were degrees of reduction from the interfrontal stripes of Desmome- topa but prefered to retain the distinctness of the stripes as uniquely characteristic of Desmometopa. According to the phyloge- netic study of Litometopa and Desmome- topa present in this paper, Litometopa is closely related to some Desmometopa spe- cies and is part of this genus. Therefore, we include Litometopa as a subgenus of Des- Based on new characters, the subgenus Litometopa and its type species, D. (Litometopa) glabrifrons (Sabrosky), are redescribed. In her key to genera of Mili- chiidae Brake (2000) pairs Litometopa with the branch leading to Desmometopa, Lep- tometopa Becker, and Madiza Fallén, dif- fering by the presence of only one orbital seta and by the absence of interfrontal se- tulae, except for an anterior pair. The iden- mometopa. 266 tification of species of Litometopa is in some cases quite difficult, because, while they do not differ in male or female ter- minalia, leg coloration and the shape of the head do differ but these characters are somewhat variable. As an added compli- cation, the shape of the head is sexually di- morphic. Future research on Litometopa should in- clude histological sections of the male ab- domen, which contains sclerotized reser- voirs of possible glandular function. The terminology essentially follows McAlpine (1981) with a few exceptions. We follow White et al. (2000) in using “*mi- crotrichia’”” instead of McAlpine’s “‘prui- nescence,’ because the body surface is cov- ered with microscopic outgrowth of the cu- ticle and not with dust or a waxy substance (pruinose). For the vertical setae we use the terms “medial”? and “‘lateral’’ (White et al. 2000) instead of the traditional “inner” and “‘outer,”” respectively. The terminology of the prothorax follows Speight (1969), and that of the male genitalia Cumming et al. (1995). The definition of “dark” is brown and black. Descriptions are composite. For the most part information given in the de- scription of the genus is not repeated in the species descriptions. We thank the following curators and in- stitutions for lending specimens. HU Museum fiir Naturkunde der Humboldt Universitat, Berlin, Germany (Dr. M. Kotrba) Staatliches Museum fiir Natur- kunde, Stuttgart, Germany (Dr. H.-P. Tschorsnig) Tel Aviv University, Tel Aviv, Israel Fakultat fiir Biologie, Universi- tit Bielefeld, Bielefeld, Germa- ny (Dr. M. von Tschirnhaus) National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA (Dr. W. N. Mathis) SMNS TAU UBI USNM Some paratypes will be deposited in the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON following institutions (as well as the USNM): BM The Natural History Museum, London, U.K. NMW.Z_ National Museum and Galleries of Wales, Cardiff, U.K. ZFMK Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany ZSM Zoologische Staatssammlung Miinchen, Germany Desmometopa subg. Litometopa Sabrosky, new status Litometopa Sabrosky 1965: 4. Type species: Litometopa glabrifrons Sa- brosky. Original designation. Description.—Head (Figs. 4—5, 7-8, 10— 11): About as long or longer than high, black, only base of arista and often base of palpus yellowish, polished except for mi- crotrichose antenna, fovea, and ventral fa- cial margin. Frons glabrous, polished, 1.3— 2.0X as long as broad (length measured from ptilinal fissure to base of postocellar setae; breadth measured at narrowest point between eye margins). Frontal width nearly equal in male and female (Fig. 10); lacking interfrontal stripes or setulae, except pair of setulae anteriorly. Frontal triangle small, not extended beyond ocelli. One reclinate and slightly lateroclinate orbital seta and 2 medioclinate frontal setae. Two proclinate setulae between orbital seta and posterior frontal seta, anterior seta slightly larger than posterior one, possibly representing reduced anterior orbital seta. One medioclinate se- tula present close and anterior to posterior frontal seta, and one medioclinate setula present anterior to anterior frontal seta. La- teroproclinate ocellar and medial and lateral vertical setae present. Postocellar setae par- allel to slightly convergent. Lunule visible, extended to basal part of first flagellomere, triangular, pointed, with pair of setulae. Face concave, antenna short, first flagello- mere round; arista long, slender, microscop- ically pubescent. Vibrissal angle produced, VOLUME 105, NUMBER 2 267 Fig. 1. usually half as broad as first flagellomere, but sometimes equibroad or broader (vi- brissal angle measured from anteriormost point of eye margin to anterior tip of vi- brissal angle), vibrissa well developed. Gena about 0.1—0.2* as high as eye. Post- orbital region (i.e., space between eye and postocular setulae) in male 2—3 longer than in female, up to about a third as long as eye (Fig. 4) (length of eye measured as horizontal line between anteriormost and posteriormost points of eye; length of post- orbital region measured at posteriormost point of eye). Palpus flat, elongate spatulate to rather broad (Figs. 6, 9), with small se- tulae and two longer setae at anterolateral Desmometopa dolichocephala, male, habitus (after Brake 2000). edge; in male entirely microtrichose, in fe- male ventrally glabrous and shiny. Clypeo- labral membrane at base of palpus slightly sclerotized. Prementum broader and thicker than combined labella, which have four pseudotracheae each. In female, pseudotra- cheal teeth on medial pseudotrachea ex- tremely long and thin (Fig. 20, arrow), bent to about 90°, and with small subsidiary tooth at angle. Small subsidiary teeth also present on teeth of other pseudotracheae. In male, teeth on all pseudotracheae about equally long and neither bent nor toothed (Fig. 19). Thorax: Black, narrow and elongate (Fig. 1). Mesonotum 1.5—2.0 as long as 268 broad, finely shagreened, subshiny, almost bare of setulae, with only dorsocentral row of setulae and median (unpaired) acrosti- chal irregular rows posteriorly, and few scattered setulae later- ally. Anepisternum and furcasternum elon- gate and fused. Basisternum with precoxal bridge, similar to variant ““Q/T” of Speight (1969) (Fig. 12). Pleura strongly shiny, though in part somewhat rugose, anepime- ron and Katepisternum posteriorly and mer- on, katatergite and anatergite entirely dull, gray microtrichose. Scutellum shiny, gla- brous. Chaetotaxy: | long postpronotal seta and 2 postpronotal setulae, of which one may be longer and directed dorsad, 2 no- topleural setae, | presutural represented by setula, 1 short supraalar, | long postalar, | short intraalar, 2 dorsocentral setae, anterior one about half length of posterior one, and row of dorsocentral setulae progressively shortened anteriorly, 1 apical and 1 basal scutellar setae, basal seta /, as long as api- cal seta, anepisternum bare, | katepisternal and no anepimeral seta. Legs: Foreleg large, appearing raptorial, with elongate coxa and often incrassate fe- mur with anteroventral and posteroventral rows of 6-8 strong spines each (Fig. 1). Fo- recoxa bright yellow, and foretibia and for- etarsus black in all studied species, whereas forefemur varies between species, from pre- dominantly black to almost entirely yellow. row, that becomes 2 Foreleg appears thicker due to brush of black setulae ventrally at distal end of tibia and on tarsus. Mid- and hindlegs more or less normal. Mid- and hindtibiae and mid- and hindtarsi except distal tarsomeres yel- low. Basal two tarsomeres of hindleg with posteroventral brush of either pale (yellow) or dark (brown or black) setulae. Pale se- tulae more coarsly and more spirally fluted than normal black setulae in D. brachyce- phala (Fig. 21). Structure of setulae in brush not studied in other species. Wing: Hyaline. R,,; and M,,, very slightly convergent (Fig. 13). Last section of M,,, about 2.75 as long as penultimate PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON section. Cell cup closed, but only weakly differentiated. Abdomen: Black, slender and elongate. Male usually with paired sclerotized inter- nal saclike structures (see next section). Male terminalia similar to those of other Desmometopa species. Cercus as large as epandrium in lateral view (Fig. 17); 2—4 long setae laterally on epandrium, number sometimes varying between sides. Surstylus partly fused to epandrium, covered by se- tulae medially (Fig. 18), similar to flame of candle in ventral view. Pregonite with 2 se- tulae distally. Distiphallus forming long membranous tube, which is widened in middle, dorsobasally with sclerotized strip. Female with tergite and sternite 6 and 7 not reduced in size (reduced in many other Milichiidae). Lateral margins of sternite 7 and tergite 7 overlapping (Fig. 22). Sternite 6 anteromedially with small pointed process (Fig. 16). Segment 7 internally with sec- ondary oOvipositor (Brake 2000; and see next section). Female internal reproductive system without any sclerotized structures. Spermathecal ducts elongated and loosely (i.e., the ducts are not lying directly next to each other) rolled together distally into one small coil. Spermathecal duct basally wider and possibly surrounded by muscles. Duct and surrounding epithelium in coil narrow- er. No sclerotized spermathecal capsule. Distal end of spermathecal duct projecting out of coil and surrounded by epithelial gland cells. Tip of duct slightly enlarged (see fig. 24H, D. sabroskyi, in Brake 2000). Peculiarities of the male and female gen- italia—Males of Litometopa usually have peculiar paired structures within the fifth abdominal segment (Fig. 14). Each struc- ture consists of two sclerotized reservoirs, which are surrounded by tissue. The reser- voirs are elongate sacs connected by ducts, which are sclerotized in the same manner. The duct leading to the anterior reservoir is longer than the one leading to the posterior reservoir. The ducts lie dorsally, but the sacs are directed ventrad. Near the termin- alia, the ducts of both reservoirs meet and VOLUME 105, NUMBER 2 fuse to form a common duct. The common ducts of either side open close to each other dorsally between the fifth tergite and the terminalia. On each common duct there is a spiracle, probably spiracle 7. A second spiracle lies in the membrane between the duct opening and the fifth tergite. This is probably the spiracle 6. There does not seem to be a valve for the reservoirs. The sclerotized wall of the reservoirs 1s covered by slight, longish indentations, similar to the surface of a golfball. A few short se- tulalike structures are on these indentations towards the lumen of the reservoir. It is pos- sible that each of these structures represents a connection between the surrounding tis- sue and the reservoir. However, in SEM dis- sections no holes in the tip of the setulalike structures could be discerned. The reser- voirs in all dissected specimens (both dry and alcohol-preserved material) seemed to be empty. We suppose that this structure represents a gland, based on the tissue sur- rounding the reservoirs and on the setula- like structures in the reservoirs. We will use the term ‘“‘abdominal reservoirs” for this structure. The abdominal reservoirs possi- bly evolved from synsternite 7/8, which is absent in Litometopa, because the 6th and 7th spiracle are often in synsternite 7/8 in Milichiidae and especially in Desmometo- pa. In females there is a specialized structure inside segment 7, which is called a second- ary Ovipositor and is used for oviposition (Fig. 15) (Brake 2000). Presumably, while ovipositing, the secondary ovipositor is everted telescopelike between sternite 7 and the subanal plate. Basally the structure con- sists of a broad membranous ring with many anteriorly directed barbed spines. Distally there are two weakly sclerotized strips each with a row of yellow setulae. These strips originate ventrolaterally and meet dorsally at the functional tip of the ovipositor. Between these strips, which rep- resent sternite 8, les the genital opening. When the secondary ovipositor is retracted, only the basal part is turned over, with the 269 barbed spines now directed posteriorly. In addition to the secondary ovipositor, Lito- metopa 1s characterised by a very small ter- gite 8, a bare supra-anal plate and short, round cerci. This combination of characters is not restricted to the subgenus Litometopa, but is found in Desmometopa species and in Pholeomyia and is therefore probably plesiomorphic. Distribution.—Afrotropical (Fig. 2): Ethiopia, Kenya, Tanzania, Uganda, Rwan- da, Congo, Nigeria, and South Africa. This apparently disjunct pattern indicates a pos- sibly wider and more homogenous distri- bution in the Afrotropical Region. Biology.—Nothing is known about the biology of Litometopa species. Most spec- imens were swept with a net from flowers, savanna trees and bushes, and some speci- mens were sampled by canopy fogging. Since sclerotized reservoirs are found in the abdomen of males only, it is possible that they represent glands that produce sex pheromones. Abdominal glands are quite rare in Diptera. Hennig (1973) gives an overview of the known cases. If only pre- sent in one sex, glands were found mostly in females, for example in Phoridae, Laux- aniidae, and some Tephritidae. However in other Tephritidae, Chloropidae (Thaumato- myia notata Meigen), and Milichiidae (Madiza glabra Fallén) glands were found only in males. These glands have no simi- larity to the abdominal reservoirs in Lito- metopa, because they are not sclerotized. The only record of sclerotized glandlike structures is from Empididae (Smith and Davies 1964). However, in contrast to Li- tometopa, the two pairs of abdominal or- gans of males of Austrodrapetis lie beneath the third and fourth tergite and have no ap- parent opening to the surface. Rectal glands are known from Sepsidae, Tephritidae, and Coelopidae and were shown to have a de- fensive function in Sepsidae (Meier and Dettner 1998). Cladistic analysis.—The data set for the computer-based cladistic analysis comprises 19 characters and 13 taxa, including 12 spe- 270 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON @ flavicornis | | @ glandulifera | | & brachycephala | Oglabrifrons | Osabroskyi \ \ \ | | ~\ Adolichoceph 31a \ & nigrifemorata, : =} Fig. 2. cies of Desmometopa and the stem-species pattern of Milichiidae as an outgroup. Des- mometopa m-nigrum Zetterstedt is a repre- sentative of the subgenus Desmometopa Loew; D. tarsalis Loew, D. floridensis Sa- brosky, D. meridionalis Sabrosky, and D. melanderi Sabrosky are representatives of the subgenus Platophrymia Williston of Desmometopa, and the other species belong to the subgenus Litometopa Sabrosky. All characters were treated as unordered. The cladistic analysis was carried out using the exhaustive search option “‘ie’’ of Hennig86, which finds the most parsimonious clado- gram(s). The precise character distribution can be seen in the data matrix (Table 1). Distribution records of Desmometopa (Litometopa) in Africa. HEAD l. i) Interfrontal stripes: (O) not present; (1) present. In Desmometopa and some Pholeomyia and Phyllomyza_ species, the row of interfrontal setulae is em- phasized by sclerotizations at the base of the setulae and the development of microtrichia. In such cases the rows are called interfrontal stripes. Interfrontal stripes are the only known apomorphic character for the genus Desmometopa (Sabrosky 1983). . Posterior orbital seta: (0) present; (1) absent. According to Brake (2000) the small medio-reclinate seta posterior to VOLUME 105, NUMBER 2 Table 1. Character matrix for species of Desmo- mentopa (Litometopa) and outgroup. Stem-species pattern of Milichiidae 00000 02000 00 . (D.) m-nigrum . (P.) tarsalis . (P.) floridensis . (P.) meridionalis . (P.) melandert . (L.) dolichocephala 01 . (L.) glabrifrons . (L.) glandulifera . (L.) sabrosky . (L.) flavicornis . (L.) nigrifemorata OLTA0! 14 . (L.) brachycephala 10000 00000 ( 10000 11000 00 10000 11000 00110 0010 10000 11100 00110 0010 10000 11100 0010 1 Je Ey a Ot Ue et UE Ls Spa a a et it gt ie Ef) EE OAT LO ae | )0 0010 00010 5 the large lateroclinate orbital setae in Desmometopa is homologous to the posterior (“upper”) orbital seta in the stem-species pattern of Milichiidae. This seta is apomorphically absent in Litometopa. Middle orbital seta: (QO) lateroclinate;: (1) latero-reclinate. As discussed above, the posterior orbital seta in Li- tometopa 1s absent. Therefore the pos- teriormost orbital seta in this subgenus is probably homologous to the middle orbital seta in the stem-species pattern of Milichtidae (Brake 2000). A change in the inclination of the middle orbital seta from lateroclinate and only slightly reclinate to mainly reclinate and only slightly lateroclinate occurred several times within the Milichiidae and is apo- morphic for Litometopa. The anterior orbital seta in Litometopa is possibly represented by the anterior of two se- tulae between middle orbital seta and frontal setae. Anterior frontal seta: (0) 0.66—1.00% as long as posterior frontal seta; (1) about half as long as posterior frontal seta. In Milichidae the frontal setae usually have the same length. . Setulae between frontal setae: (0) two; 271 (1) one. The number of setulae between the frontal setae in Milichiidae is often two. In Desmometopa the number is two in all studied species of the sub- genera Desmometopa and Platophry- mia. Therefore the presence of only one setula is considered to be apo- morphic. Face: (O) not concave; (1) concave. The comparatively protuberant ventral mar- gin of the head and deeply concave face as seen in profile, is one of the characters Sabrosky (1983) used to separate the subgenus Platophrymia from the subgenus Desmometopa, in which the face is only slightly concave. Since the face is only slightly concave in most Milichiidae, this character state is supposed to be plesiomorphic. Palpus coloration in male: (QO) basally yellow; (1) entirely black. The colora- tion of the palpus varies within Mili- chiidae and within Therefore the polarity of this character is unknown. Palpus shape in male: (O) not enlarged: (1) enlarged. In the stem-species pat- tern of Milichiidae the palpus is short and clavate. The shape of the palpus in several Desmometopa species differs Desmometopa. from the stem-species pattern in that it is slightly longer with the width a third of the vertical eye diameter or more. Within Desmometopa an enlarged pal- pus occurs in two species of the sub- genus Desmometopa and in less than half of the species of Platophrymia. Palpus microtrichia in female: (0) en- tirely microtrichose; (1) polished ven- trally. In all Milichiidae we studied (for list of studied species see Brake 2000), including members of Desmometopa, the palpus is entirely microtrichose. Therefore a ventrally polished palpus is considered apomorphic. THORAX 10. Presutural seta: (0) as long as or longer than supraalar seta; (1) shorter than su- 272 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON praalar seta. The size reduction of the presutural seta is apomorphic for Lito- metopa. 11. Mesonotum setulation: (0) with many setulae; (1) almost bare of setulae ex- cept for dorsocentral row of setulae and median (unpaired) acrostichal row, that becomes two irregular rows posteriorly, and few scattered setulae laterally. In all Milichiidae we studied, the meson- otum is covered by many acrostichal setulae. The reduction of the number of acrostichal setulae is therefore consid- ered apomorphic for Litometopa. 12. Suture between anepisternum and fur- casternum (Q) present; (1) absent. In all Milichiidae we studied, the suture be- tween anepisternum and furcasternum is present. The absence of this suture is therefore considered apomorphic for Litometopa. LEGS 13. Forecoxa coloration: (0) black; (1) yel- low. In most Milichiidae all coxae are black. However, the forecoxa is yellow- ish in several species of Neophyllomyza and Leptometopa. 14. Forecoxa: (0) not elongate; (1) elon- gate. In Litometopa and Platophrymia the thorax and forecoxa are elongate, especially in relation to the height of the thorax at the base of the coxa. 15. Forefemur coloration: (0) black; (1) slightly yellow at base; (2) basal 0.25— 0.60 yellow; (3) almost entirely yellow except for anterodorsal brown. stripe distally. In most Milichiidae, including members of Desmometopa, the forete- mur is black. Therefore a yellow fore- femur is considered apomorphic. 16. Midfemur and hindfemur coloration, except for apices: (O) black; (1) yellow. In most Milichiidae, including mem- bers of Desmometopa, the midfemur and hindfemur are black. Therefore yellow mid- and hindfemora are con- sidered apomorphic. 17. Apices of midfemur and hindfemur: (0) black; (1) yellow. Dark legs with yel- low apices of the midfemur and hind- femur are known in several species of Milichiidae, but seem to have evolved several times independently. As_ the apices are black in all studied species of the subgenera Desmometopa and Platophrymia, yellow apices are con- sidered to be apomorphic. 18. Posteroventral brush on hindtarsus: (0) black or brown; (1) yellow. The col- oration of this brush varies within Mil- ichiidae, and the polarity of this char- acter is unknown. ABDOMEN 19. Sclerotized reservoires in male abdo- men: (Q) absent; (1) present. Sclero- tized reservoirs are unique to Litome- topa and have neither been found in other Milichiidae, nor in other acalyp- trate flies. The cladistic analysis found 48 most par- simonious trees of 32 steps (consistency in- dex: 0.65; retention index: 0.86). The Nel- son tree of these 48 trees is depicted in Fig. 3. Discussion (numbers in square brackets refer to the characters used in the cladistic analysis).—Litometopa is closely related to Desmometopa (Platophrymia) (Sabrosky 1965, 1983), sharing the shiny black later- oventral corner of the facial plate, imme- diately mesad of the vibrissal angle and the concave face [6] (Brake 2000). However, there are two rows of strong spines on the forefemur in Litometopa, but only one an- teroventral row of weak spines in these Pla- tophrymia species. Within Platophrymia, Litometopa is most closely related to the group around D. floridensis, D. melanderi, and D. meridionalis, sharing the enlarged palpus [8] and the elongated yellow [13, 14] forecoxa and often incrassate forefemur. However, Litometopa is Afrotropical, whereas the related Platophrymia spp. are Nearctic or Neotropical. The close relation- ship between Litometopa and some Plato- VOLUME 105, NUMBER 2 273 Desmometopa Platophrymia Litometopa I & O oO © o = fo wy St S o ce OU me 2 8 Say suan Ses 8 Se 8 a 2 2 = 6 So Sass or. Sores = O eS oO iS fe) =. .o) 3 — Oo oS S) 6. Sep ee SS ie ah OM eter S OF £ = ¢ G&S 6 oe s- 2] > g fe) oO {e) pit de ie) Oo oO peg 2 See Sf Sf = sa So oS: Ss © 4a GiGi GQ) 2 AVA ee pe) 2 [14-0 [116-0 7-1 18-1 is LJ18- 7-1 6-1 Ie 17-0 ats [119-0 [116-1 5a), 15-2 17-1 [] 7-0, 8-0 2-1, 3-1,9-1, 10-1, he} 13-0 AMEN AES [11-0, 4-1, 18-0, 19-1 14-1 [1 8-1, 13-1 6-1 [i7-1, 18-1 r1-1 Fig. 3. Phylogenetic tree of Desmometopa subgenera; shaded hatchmarks indicate character changes that either reverse or evolved at least twice on the tree; black hatchmarks indicate changes that do not reverse and evolved only once; reductions are marked with a dark shading. phrymia species results in a paraphyly of Platophrymia. However, we have decided to keep both subgenera separate, as they are easily recognisable and differ from each other in several characters, and a more rig- orous phylogenetic study of Platophrymia and of the whole genus Desmometopa, which would clarify the presence of mono- phyletic groups within the genus, is still needed. The subgenus Litometopa, as diagnosed above, is clearly a monophyletic group. Its apomorphies are: [2] posterior orbital seta lost (plesiomorph: present): [3] middle or- bital seta (in Litometopa the actual posterior seta) reclinate (plesiomorph: lateroclinate): anterior orbital seta reduced in size (ple- siomorph: as large as the other orbital se- tae); [9] female palpus polished ventrally (plesiomorph: palpus entirely microtricho- se); [10] presutural seta reduced in size (plesiomorph: not reduced); [11] mesono- tum almost bare of setulae (plesiomorph: covered with setulae); [12] suture between anepisternum and furcasternum lost (ple- siomorph: present); and probably also [19] the presence of abdominal reservoirs in males (plesiomorph: not present), though the state of this character is unknown for D. nigrifemorata. Within Litometopa, the yellow forefemur [15-3] is the synapomorphic character of a clade comprising D. brachycephala, D. dol- ichocephala, D. flavicornis, and D. sabros- 274 kyi. These four species together with D. glabrifrons and D. glandulifera form a clade that is supported by the following apomorphic characters: only one setula pre- sent between frontal setae [5—1l]. basal 0.25—0.60 of forefemur yellow [15-2], and apices of mid- and hindfemora yellow [17— 1]. KEY TO SPECIES OF DESMOMETOPA (LITOMETOPA) 1. Posteroventral brush of hindtarsus yellow ... 2 — Posteroventral brush of hindtarsus black or brown 2. Mid- and hindfemora black. Male with first fla- gellomere black icra eee D. (L.) brachycephala, new species — Mid- and hindfemora yellow. Male with first fla- gellomere yellowish medially and basally .... 3 . Male with postorbital region about ¥, as long as eye — Male with postorbital region about /,, as long Ww D. (L.) flavicornis, new species ASHOY Cover vesycke eiowuaney cn eemateakes D. (L.) species A 4p Remora yellow. ceccon.. ore at ian caer 5) — At least distal % of forefemur black; mid- and hindfemora mainly or entirely black ...... . 6 an . Head of male (Fig. 7) distinctly longer than high, vibrissal angle distinctly produced and antenna in deep fovea. Head of female (Fig. 8) not distinctly longer than high, vibrissal angle less produced than in male. Palpus black and enlarged in both sexes (Fig. 9). Anterior frontal seta 0.66—1.00* as long as posterior frontal Setamperuey- ter. D. (L.) dolichocephala, new species — Head of both sexes as long as high, vibrissal angle only slightly produced in both sexes (Fig. 11). Palpus black or yellow with black tip, not enlarged. Anterior frontal seta less than half as long as posterior frontal seta .......... Be ene Pee en D. (L.) sabroskyi, new species 6. Forefemur almost entirely black, only slightly yellow at base. Mid- and hindfemora entirely black. Both sexes with black enlarged palpus (as in Fig. 9) OER t Eines D. (L.) nigrifemorata, new species — Forefemur yellow on basal 0.25—0.60, rest black. Mid- and hindfemora entirely black, or apices yellow. Palpus not enlarged, base often WANE on Ste Ces gure oes saga be aoe 7 7. Forefemur yellow on basal 0.60. Apex of mid- and hindfemora yellow PES eee D. (L.) glabrifrons (Sabrosky 1965) — Forefemur yellow on basal 0.25—0.40. Mid- and hindfemora entirely black ra bad Rl Sees D. (L.) glandulifera, new species PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Desmometopa (Litometopa) brachycephala Brake and Freidberg, new species (Figs. 4-6, 19-21) Litometopa sp. 2: Brake 2000: 11. Diagnosis.—This species is distinguished from its congeners by the following com- bination of characters: Forefemur almost entirely yellow, with anterodorsal brown stripe distally; mid- and hindfemora pre- dominantly black, apices yellow; postero- ventral brush on hindtarsus yellow. Male.—Predominantly black; forecoxa, forefemur except anterodorsal brown stripe distally, apices of mid- and hindfemora, mid- and hindtibiae, and mid- and hindtarsi except distal tarsomeres yellow. Palpus largely yellow with black tip. Posteroven- tral brush on hindtarsus yellow. Head: An- terior frontal seta about half as long as pos- terior frontal seta (Fig. 4). Postorbital re- gion very long, about 4% as long as eye. Pal- pus as in Fig. 6. Wing: Length: 1.8—2.6 mm (holotype: 2.5 mm). Abdomen: Tergites 1— 4 microtrichose and subshiny medially, lat- eral and posterior margins polished; micro- trichose spots progressively decreasing pos- teriorly, so that tergite 4 microtrichose only anteromedially or entirely polished. Tergite 5 and all sternites polished. Abdominal res- ervoirs present. Male genitalia as in Fig. 16. Female (Fig. 5).—Differs from male in palpus more extensively black and slightly broadened. Type material.—Holotype male: “UGANDA. S.W.: Fort Portal, 5 km NW, 2,000 m, 10.1.1996, I. Yarom & A. Freid- berg.” The holotype is double mounted, is in excellent condition, and is deposited in TAU. Paratypes: 10 d and 3 9, same col- lection data as holotype (TAU, USNM). UGANDA. S.W. Kabale, 7 km NE, 1,950 m, 23.XII.1995, I. Yarom & A. Freidberg (S d; TAU); Mpigi, 1.400 m, 40 km SW Kampala, 22.X1I.1995, I. Yarom & A. Freidberg (2 3; TAU); Kisoro, 2,000 m, 26.X11.1995, I. Yarom & A. Freidberg (1 6; TAU). KENYA. Bungoma, 12.— VOLUME 105, NUMBER 2 205 Figs. 4-11. 4-6, Desmometopa brachycephala. 4, Head, lateral view, male. 5, Head, lateral view, female. 6, Palpus, male. 7-10, D. dolichocephala. 7, Head, lateral view, male. 8, Head, lateral view, female. 9. Palpus, male. 10, Head, frontal view. 11, D. sabroskyi, head, lateral view, male. Scales: 0.1 mm. 276 13.1.1996, I. Yarom & A. Freidberg (1 6, forelegs missing; TAU); 10.X.1998, F Kap- lan & A. Freidberg (1 6; TAU). ETHIO- PIA. Shewa, Wendo Genet, 2,100 m, 29.1.2000, A. Freidberg & I. Yarom (6 d, 5) 2s) TAUL INMW:Z, BM) 1-900 im; 28.1.2000, A. Freidberg & I. Yarom (9 d, 3 2; TAU, SMNS). Non-type material KENYA. West Po- kot, Chepareria, 4—5.X1.1983, A. Freidberg (1 2; TAU). UGANDA. S.W. Ishaka, 25 km N, 1,900 m, 5.1.1996, I. Yarom & A. Freidberg (1 2; TAU). Etymology.—The species name is de- rived from the Greek brachys = short and cephalae = head, denoting the short head. Remarks.—The non-type female from Kenya differs from the type specimens in the black apex of the forefemur. The non- type female from Uganda was not added to the type series, because the head is shriv- eled. Desmometopa (Litometopa) dolichocephala Brake and Freidberg, new species (Figs. 1, 7-10, 17-18) Litometopa sp. 1: Brake 2000: 11. Diagnosis.—This species is distinguished from its congeners by the following com- bination of characters: Male with head lon- ger than high, distinctly produced vibrissal angle, and strongly warped upward facial margin. Both sexes with anterior frontal seta 0.66—1.00* as long as posterior seta. Palpus black and enlarged. Forefemur al- most entirely yellow with anterodorsal brown stripe distally; mid- and hindfemora yellow; posteroventral brush on hindtarsus dark. Male.—Predominantly black; coxae and femora, mid- and hindtibiae and mid- and hindtarsi except distal tarsomeres yellow. Head: Anterior frontal seta 0.66—1.00%* as long as posterior frontal seta (Figs. 7-8). Postorbital region /—Y,* as long as eye. Vi- brissal angle distinctly produced anteriorly, as broad as first flagellomere, the angle em- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON phazised by shiny black lateroventral corner of facial plate immediately mesad of vibris- sal angle and usually warped forward and upward beyond it. Ventral margin of head comparatively long and face deeply con- cave as seen in profile, antenna in deep fo- vea. Palpus enlarged (Fig. 9). Wing: Length: 1.9—2.8 mm (holotype: 2.6 mm). Abdomen: Tergites 1—4 microtrichose and subshiny medially, lateral and posterior margins polished, microtrichose spots pro- gressively decreasing posteriorly, so that tergite 4 is microtrichose only anteromedi- ally. Tergite 5 and all sternites polished. Abdominal reservoirs present. Male geni- talia as in Figs. 17-18. Female.—Differs from male in head not elongated (Fig. 8) and vibrissal angle less than half as long as first flagellomere. Type material—Holotype male: ““TAN- ZANIA, Same, Rt. Bl, 8—16.1X.1992, A. Freidberg.”’ The holotype is double mount- ed, is in excellent condition, and is depos- ited in TAU. Paratypes: 9 d and 1 2, same collection data as holotype (TAU, USNM). TANZANIA. Same, Rt. B1, 22. VIII.1996, A. Freidberg (2 6; TAU); 8.IX.1996, A. Freidberg (7 6, 1 2; TAU, HU). Etymology.—The species name is de- rived from the Greek dolichos = long and cephalae = head, denoting the particularly long head of the male. Desmometopa (Litometopa) flavicornis Brake and Freidberg, new species Diagnosis.—This species is distinguished from its congeners by the following com- bination of characters: First flagellomere in male yellowish medially and at extreme base, dark only around base of arista; for- efemur in both sexes almost entirely yel- low, with anterodorsal brown stripe distally: mid- and hindfemora yellow; posteroventral brush on hindtarsus yellow. Male.—Predominantly black; forecoxa, forefemur except anterodorsal brown stripe distally, mid- and hindfemora, mid- and hindtibiae, and mid- and hindtarsi except distal tarsomeres yellow. Palpus yellow VOLUME 105, NUMBER 2 277 i 7 cell cup’... CUA phap phap scl Figs. 12-18. 12, Desmometopa sabroskyi, prothorax, ventral view (from Brake 2000), bs—basisternum, cx— base of coxa, pb—precoxal bridge. 13-14, Desmometopa sp. A, male. 13, Wing (from Brake 2000), CuA— anterior cubitus, cell cup—posterior cubital cell, dm-cu—discal medial-cubital crossvein, M—media, R—radius, r-m—radial-medial crossvein. 14, Abdominal reservoirs, ventral view, h—setulae on indentations, r—reservoir, t—tissue, o—opening, s—spiracle, T—tergite. 15—16, D. sabroskyi, female. 15, Tip of ovipositor with secondary Ovipositor inverted, lateral view (from Brake 2000), ce—cerci, S—sternite, sba—subanal plate, spa—supra-anal plate, T—tergite. 16, Sternite 6 (from Brake 2000), p—process at anterior margin. Scales: 0.1 mm. 17-18, D. dolichocephala, male terminalia. 17, Lateral view. 18, Viewed in direction of black arrow in Fig. 17. Scales: 0.1 mm. 278 with light brown to brown tip. Postero- ventral brush on hindtarsus yellow. Head: Anterior frontal seta half as long as poste- rior frontal seta. Postorbital region about ¥.X as long as eye. Palpus as in Fig. 6. Wing: Length: 2.1—-2.5 mm (holotype: 2.2 mm). Abdomen: Tergites 1—5 microtrichose and subshiny medially, polished laterally. Abdominal reservoirs present. Male geni- talia as in Fig. 17. Female.—Differs from male in entirely black first flagellomere. Palpus black except mesobasally. Type material—Holotype male: “‘NI- GERIA: Ife, 2 Aug. 1969/J. T. Meddler Coll.” The holotype is double mounted, is in excellent condition, and is deposited in the USNM. Paratypes: 2 d and 2 @, same collection data as holotype (USNM). NI- GERIA. Niger State: Mariga River, 80 km NW Minna, 11.XII.1987, Fini Kaplan (1 2: TAU). Etymology.—The species name is de- rived from the Latin flavus = light and cor- nus = horn, denoting the yellowish first fla- gellomere of the males. Desmometopa (Litometopa) glabrifrons (Sabrosky), new combination Litometopa glabrifrons Sabrosky 1965: 4. Diagnosis.—This species is distinguished from its congeners by the following com- bination of characters: Basal ¥; of forefemur yellow, remainder black; mid- and hindfem- ora black, except for apices; posteroventral brush on hindtarsus dark; male without ab- dominal reservoirs. Male.—Predominantly black; basal *% of palpus, forecoxa, basal ¥, of forefemur, api- ces of mid- and hindfemora, mid- and hind- tibiae and mid- and hindtarsi except distal tarsomeres yellow. Head: Anterior frontal setae half as long as posterior frontal seta. Postorbital region ¥.—¥, as long as eye. Pal- pus as in Fig. 6. Wing: Length: 2.2-2.6 mm (holotype: 2.6 mm). Abdomen: Tergites 14 microtrichose and subshiny medially, polished laterally, microtrichose spots pro- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gressively decreasing posteriorly; tergite 5 and all sternites polished. Abdominal res- ervoirs absent, but small base of ducts pre- sent (in dissections of paratype and non- type material). Male genitalia as in Fig. 17. Female.—Differs from male in basal half of palpus yellow and palpus slightly broad- er. Type material.—The holotype male bears several labels: ““O. Afrika, T. T. Marangu, 1.20. Maerz 1959, Lindner leg.”’ On the lower side of the label the date is specified as “17.11.” The second label reads ““Ho- lotype, Adesmometopa glabrifrons, male, C. W. Sabrosky.” The third label says “Adesmometopa glabrifrons, det. Sabros- ky.’ The fourth label reads “‘Litometopa glabrifrons Sabrosky.”” The second and third labels were probably written by Sa- brosky himself and indicate that he first considered naming the genus Adesmome- topa. However, in his description he named the genus Litometopa and therefore the cor- rect name is stated on the fourth label, which was probably not written by Sabros- ky himself. The holotype is double mount- ed, is in good condition except for the slightly shriveled head, and is deposited in the SMNS. One male paratype bears the first label with the same data as the holo- type but with “10.III.” on the lower side (USNM). The second label says “Paratype, Litometopa glabrifrons, male, C.W. Sabros- ky,’ and the third label reads ““Litometopa glabrifrons, det. Sabrosky.”” For the other two original paratypes see below. Additional specimens examined.—TAN- ZANIA. Marangu, E. Lindner, 10.11.1959 (1 2; SMNS). SOUTH AFRICA. Natal, Uvongo, South Coast, 11.X.1983, A. Freid- berg (2 6, 1 2; TAU); North Transvaal, Tzaneen, 1977, Van Eeden, at Euphorbia flowers (1 2; USNM). KENYA (West). Ka- penguria—Tartar Road, 24.X1.1989, A. Freidberg & E Kaplan (1 2; TAU). ETHI- OPIA. Shewa, Wendo Genet, 1,900 m, 28.1.2000, A. Freidberg & I. Yarom (1 6; TAU); 2,100 m, 29.1.2000, A. Freidberg & I. Yarom (2 6; TAU). VOLUME 105, NUMBER 2 Remarks.—Two of the original para- types, one male and one female (allotype), were found not to be conspecific with the holotype and are redescribed below as D. sabroskyi. These two specimens differ from the holotype of D. glabrifrons by the yel- low mid- and hindfemora and the almost entirely yellow forefemur with an antero- dorsal brown stripe distally. One male from South Africa, Natal, Uvongo, differs from the holotype in the forefemur being black on the distal half posteriorly and distal 7; anteriorly and in the shape of the head, which is similar to D. brachycephala. The female from South Af- rica, North Transvaal, differs from the ho- lotype in the black apices of the mid- and hindfemora. These specimens nevertheless are considered variants of D. glabrifrons. Desmometopa (Litometopa) glandulifera Brake and Freidberg, new species Diagnosis.—This species is distinguished from other congeners by the following combination of characters: Basal ,—7, of forefemur yellow, rest black; mid- and hindfemora black; posteroventral brush on hindtarsus dark. Male.—Predominantly black; forecoxa, basal Y,—7, of forefemur, mid- and hindtib- iae and mid- and hindtarsi except distal tar- someres yellow. Mid- and hindtibiae in some specimens quite dark. Palpus brown with darker tip, but in some specimens pal- pus mesobasally yellow. Head: Anterior frontal seta half as long as posterior frontal seta. Postorbital region ¥%—/;* as long as eye. Palpus as in Fig. 6, in some specimens slightly narrower. Wing: Length: 1.9—2.4 mm (holotype: 2.4 mm). Abdomen: Ter- gites 1—4 microtrichose and subshiny me- dially, polished laterally, microtrichose spots progressively decreasing posteriorly; tergite 5 and all sternites polished. In some specimens anterior margin of tergite 5 sub- shiny. Abdominal reservoirs present. Male genitalia as in Fig. 17. Female.—Does not differ from male ex- 279 cept for the usual differences between male and female. Type material.—Holotype male: ““ETHI- OPIA: Shewa, Wendo Genet, 1,900 m, 28.1.2000, A. Freidberg & I. Yarom.”’ The holotype is double mounted, is in excellent condition, and is deposited in TAU. Para- types: 3 ¢ and 1 2, same collection data as holotype (TAU, USNM). ETHIOPIA. Shewa, Wendo Genet, 2,100 m, 29.1.2000, A. Freidberg & I. Yarom (1 6; TAU); Gamo, Gofa, 3 km NE Arba Minch, 1,300 m, ‘Forest’, 5.11.2000, A. Freidberg & I. Yarom (1 2; TAU). KENYA. Bungoma, 12.-13.1.1996, I. Yarom & A. Freidberg (1 2; TAU). KENYA. Kabarnet, 11.- 12.X.1998, F Kaplan & A. Freidberg (1 ¢; TAU); Tsavo West, Ngulia Lodge, 16— 17.VIII.1983, A. Freidberg (1 2; TAU). RWANDA. Rusumu, Ibanda Makera, 2°09'S/30°55'E, 1,350 m, canopy fogging on Teclea nobilis, gallery forest; 7.X1.1993, Th. Wagner, stored in ethanol (X 1006- Tn.5,, 1 9s XxX. 1006-En.6,,.1 do: X 1006- in.8,, 1-6; X 1006-T.n.10;.1 3). (ZSM, ZFMK, UBI). TANZANIA. Lugoba, Rt B 1218, 25.VIII.1996, A. Freidberg (3 ¢, 1 ?; TAU); Manga, Rt. B1218, 6.1X.1996, A. Freidberg, on mango, (1 d; TAU). Etymology.—The species name is de- rived from the Latin glands = gland and ferre = carry, bear, denoting the abdominal reservoirs. Desmometopa (Litometopa) nigrifemorata Brake and Freidberg, new species Diagnosis.—This species is distinguished from its congeners by the following com- bination of characters: Both sexes with en- tirely black and enlarged palpus; forefemur black except for base; mid- and hindfemora black; posteroventral brush on hindtarsus dark. Male.—Predominantly black; forecoxa, base of forefemur, mid- and hindtibiae and mid- and hindtarsi except distal tarsomeres yellow. Palpus black. Head: Anterior fron- tal seta half as long as posterior frontal seta. Two setulae between frontal setae. Length 280 Figs. 19-22. lateral view, female. 21, First tarsomere of hindleg, female, y s—yellow setae, d s—dark setae. 22, D. sabroskyi; ovipositor, lateral view (from Brake 2000), S—sternite, T—tergite. Scales: 0.1 mm. of postorbital region not recognizable be- cause of shriveled head. The same holds true for the vibrissal angle. Palpus as in Fig. 9. Wing: Length: 2.5—2.6 mm (holotype: 2.6 mm). Abdomen: Tergites 1—4 microtri- chose and subshiny medially, polished lat- erally, microtrichose spots progressively decreasing posteriorly; tergite 5 and all ster- nites polished. Female.—Differs from male in slightly narrower palpus. In female from Uvongo, anterior frontal seta about 7, as long as posterior frontal seta. Type material.—Holotype male: “SOUTH AFRICA: Natal, Uvongo, South Coast, 11.X.1983, A. Freidberg.”’ The ho- lotype is double mounted, is slightly shriv- eled, and is deposited in TAU. Paratypes: | 2, same collection data as holotype (TAU). SOUTH AFRICA. Itala Game Reserve, 27°30'S/31°20'E, 27.—29.1.1994, U. Géllner Geo AEIw): PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SEM, 19-21, Desmometopa brachycephala. 19, Labella, lateral view, male. 20, Tip of labella, Etymology.—The species name is de- rived from the Latin words niger = black and femur = thigh, femur, denoting the black femora. Desmometopa (Litometopa) sabroskyi Brake and Freidberg, new species (Figs. 11-12, 15-16, 22) Litometopa ?glabrifrons: Brake 2000: 11 (in part). Diagnosis.—This species is distinguished from its congeners by the following com- bination of characters: Forefemur almost entirely yellow with anterodorsal brown stripe distally; mid- and hindfemora yellow; posteroventral brush on hindtarsus dark. Male.—Predominantly black; fore- and hindcoxae, forefemur except anterodorsal brown stripe distally, mid- and hindfemora, mid- and hindtibiae, and mid- and hindtars1 except distal tarsomeres yellow. Palpus yel- VOLUME 105, NUMBER 2 low with dark tip or entirely dark. Head: Anterior frontal seta less than half as long as posterior frontal seta. Postorbital region moderately long, /%—’;* as long as eye (Fig. 11). Wing: Length: 1.8—2.0 mm (holotype: 2.0 mm). Abdomen: Tergites 1—4 microtri- chose and subshiny medially, polished lat- erally; tergite 5 and all sternites polished. Female.—Does not differ from male ex- cept for the usual differences between male and female. Type material.—Holotype ¢d: ““UGAN- DA: S.W., Fort Portal, 5 km NW, 2,000 m, 10.1.1996, I. Yarom & A. Freidberg.’’ The holotype is double mounted, is in excellent condition, and is deposited in TAU. Para- types: | d, same collection data as holotype (TAU); 1 2, same collection data but **7.1.1996”" (TAU). NIGERIA. Ife, 2. Aug. 1969/J. T. Meddler Coll. (3. 2; USNM). UGANDA. S.W. Kabale, 7 km NE, 1,950 m, 23.XII.1995, I. Yarom & A. Freidberg (1 2; TAU); District Masindi, Budongo Forest, N’River Sonso, 1°45’N/31°25’E, 1,200 m, canopy fogging on Cynometra al- exandri (C.a.) and Rinorea beniensis (= ar- disiifolia) (R.a.), seasonal rain forest, 19.VI.—31.VII.1995, Th. Wagner, stored in ethanol (X 1259-C.a.3, 2 9; X 1259-C.a.16, Oe Xe 1259-Reall, 2. 25 X% 1259=R a3: 3 OX 1259-Rea.7, 1-2; X 1259-Real2: lS; X1259-R als; 1 23 XK 1259-Rals, 1 Ss X 1259-R.a.16, 1 2) (ZSM, ZFMK, UBD). CONGO. Kivu-Sud, Irangi, 1°54’S/ 28°27'E, 950 m, canopy fogging on Carapa grandifolia, rain forest, 7.X1.1993, Th. Wagner, stored in ethanol (X 1006-C.g.5, 6, 15, 18, 1 2) (ZSM). TANZANIA. Maran- gu, 3.II1 1959, E. Lindner (1 6, 1 &, para- and allotype of D. glabrifrons, Sabrosky, SMNS). Etymology.—This species is named in memory of Curtis W. Sabrosky, a friend and dipterist, who dominated the taxonomic study of Milichiidae in the twentieth cen- tury and described the genus Litometopa. Remarks.—The female from Uganda, Kabale differs from the holotype in the slightly broader palpus. The male and fe- 281 male from Tanzania, Marangu, and one fe- male from Nigeria, Ife, differ from the ho- lotype in the dark apex of the forefemur. Desmometopa (Litometopa), species A (Figs. 13-14) Litometopa ?glabrifrons: Brake 2000: 11 (in part). These specimens differ from D. flavicor- nis in the shape of the head in male. Post- orbital region in male short. Palpus nearly yellow in male, black except for medial base in female, and slightly broader than in male. Mid- and hindfemora yellow to light brown. The females cannot be differentiat- ed from the females of D. flavicornis. Material examined.—UGANDA. District Masindi, Budongo Forest, N’River Sonso, 1°45'N/31°25'E, 1,200 m, canopy fogging on Rinorea beniensis (R.a.) and Teclea no- bilis (T.n.), seasonal rain forest; 19.VI.— 31.VH.1995, Th. Wagner, stored in ethanol (X 1259-R.a.3, 4 2; X 1259-R.a.17, 1 @; XM 1259-Tn2) 2965. X 1259-En. 13, 1 2) (ZSM). Remarks.—We refrain from naming this species, because there are just two males, which are very similar to D. flavicornis, so that species A might just be a variant or subspecies of the latter species. The above listed females cannot be differentiated from the females of D. flavicornis, and therefore cannot give support for naming species A. ACKNOWLEDGMENTS We are grateful to M. Balke, Dr. K. Kiontke, Prof. Dr. W. Sudhaus, and Dr. W. N. Mathis for reviewing a draft of this pa- per. We thank Dr. L. Cederholm of the En- tomologica Scandinavica Supplements for the permission to use the figures 1, 7, 10, 12-13, 16, and 22 from Brake (2000). LITERATURE CITED Brake, I. 2000. Phylogenetic systematics of the Mili- chiidae (Diptera, Schizophora). Entomologica Scandinavica Supplement 57: 1-120. Cumming, J. M., B. J. Sinclair, and D. M. Wood. 1995. Homology and phylogenetic implications of male 282 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON genitalia in Diptera—Eremoneura. Entomologica Scandinavica 26: 120-151. Hennig, W. 1973. 31. Diptera (Zweifliigler). Handbuch der Zoologie 4(2), 2/31 (Lfg. 20): 1-337. McAlpine, J. EF 1981. Chapter 2. Morphology and ter- minology: adults, pp. 9-63. In McAlpine, J. F et al., eds, Manual of Nearctic Diptera, Vol. 1. Re- search Branch, Agriculture Canada, Monograph 27. Ottawa, vi + 1-674 pp. Meier, R. and K. Dettner. 1998. The ultrastructure of a rectal gland in the Sepsidae and the composition of its secretion. 4th International Congress of Dip- terology, 6.-13. Sept. 1998, Oxford, Abstract Vol.: 143-144. Sabrosky, C. W. 1965. East African Milichiidae and Chloropidae (Diptera). Stuttgarter Beitraége zur Naturkunde 138: 1-8. . 1983. A synopsis of the world species of Des- mometopa Loew (Diptera, Milichiidae). Contri- butions of the American Entomological Institute 19(8): 1-69. Smith, K. G. V. and R. G. Davies. 1965. Glandlike abdominal structures of possible epigamic func- tion in the Diptera. Nature 4994; 321-322. Speight, M. C. D. 1969. The prothoracic morphology of acalyptrates and its use in systematics. Trans- actions of the Royal entomological Society of London 121: 325-421. White, I. M., D. H. Headrick, A. L. Norrbom and L. E. Carroll. 2000. 33 Glossary, pp. 881-924. In Aluja, M. and A. L. Norrbom, eds. Fruit Flies (Te- phritidae): Phylogeny and Evolution of Behaviour. CRC Press, Boca Raton. PROC, ENTOMOL. SOC. WASH. 105(2), 2003, pp. 283-291 NEW PEORIINI (LEPIDOPTERA: PYRALIDAE: PHYCITINAE) FROM BRAZIL JAY C. SHAFFER Department of Biology 3E1, George Mason University, Fairfax, WA 22030-4444. U.S.A. (e-mail: jshaffe 1 @gmu.edu) Abstract.—Metacommotria beckeri, n. gen., n. sp., Cipopeoria camura, n. gen., n. sp., and Harnochina digitata, n. sp. are described from Brazil. A description is also provided for the female of Zapalla dentata Shaffer, the first known female representative of the genus. The latter species, previously known only from the type locality of south- eastern Brazil, is herein recorded from Planaltina (Distrito Federal), Alto Parafso (Goias), and Chapada Guimaraes (Mato Grosso). Photographs of adult moths are provided for all four species, male head profiles for the three new species, and male antenna photomicro- graphs and wing venation drawings for M. beckeri and C. camura. Male genitalia pho- tographs are given for M. beckeri, H. digitata, Z. dentata, and female genitalia photo- graphs for C. camura and Z. dentata. Key Words: I recently had the pleasure of examining a fine assemblage of approximately 100 specimens of Brazilian Peoriini collected by Vitor Becker. Among these were three un- described species and the first known fe- male of the genus Zapalla Shaffer, that of Z. dentata Shaffer. MATERIALS AND METHODS Prepared microscope slides were mount- ed in Euparal. Moths and slides were ex- amined using a Wild M5 stereo microscope, and slides were also examined at high pow- er and photographed with an Olympus Van- ox AHBT3 microscope. Color names fol- low the ISCC-NBS Color-Name Charts (Kelly 1965) except for small structures where only general designations could be given. Moths and color samples were viewed together using the Wild microscope and a fluorescent ring light. Holotypes and most paratypes are deposited in the Museu Nacional, Rio de Janeiro [NMRJ], a single Peoriini, Phycitinae, moths, Neotropical, taxonomy paratype each for M. beckeri and C. camura is deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC [USNM]. All other speci- mens are in the collection of Vitor Becker [VOB], Brasilia, Brazil. Metacommotria Shaffer, new genus Type species.—Metacommotria beckeri Shaffer, new species. Diagnosis.—The male genitalia are sim- ilar to those of Commotria Berg, particu- larly in possessing an uncus with a pair of lateral spicate processes which are long, curve cephalad, stout at the base and taper to a sharp point. The medial process of the uncus is somewhat bandlike, not matching that of any particular species of Commotria, but falling within the range of variation (most narrowly bandlike) exhibited by that genus. Indeed, this species would fit well within Commotria but for the male anten- nae, which are subserrate rather than pec- tinate as in Commotria. Figs. 1-3. = | mm (1,2), 0.5 mm (3). Description.—Male: Frons conical. La- bial palpus (Fig. 8) porrect to somewhat obliquely ascending with third segment porrect. Maxillary palpus cylindrical, not reaching end of frons. Proboscis much re- duced, not usually visible between labial palpi. Ocellus normal, black with clear center, separated from eye by about its own width. Antenna (Fig. 11) shaft sub- serrate, basal segments fused and en- closed by parallel rows of scales, but oth- erwise unmodified. Forewing (Fig. 1) with ten veins; R, stalked with R,,; about 7; length of latter; R,,, stalked about ¥, that length; from well before upper outer angle of cell; M, from the angle; M,., fused, from lower outer angle of cell, usu- ally from same point as CuA, (M,,, well separated from CuA, in holotype); CuA, from well before the angle. Hind wing with seven veins (1A, 2A, and 3A counted as one vein): M,., fused and stalked with CuA, from about /, to over ¥; length of latter, from lower outer angle of cell: CuA, from very near the angle. Genitalia (Figs. 13—16) with medial pro- cess of uncus an irregular somewhat semi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Wing venation and male palpus. 1, Metacommotria beckeri. 2-3, Cipopeoria camura. Scale bar circular band; lateral process a long, curved, sharp-pointed spine. Gnathos arms flat, incomplete, medially with a small sclerite or a few small teeth. Juxta conical, lacking scale tufts. Valva with costa termi- nating in a small tooth; sacculus hirsute, at base with a cluster of long stiff slightly curved setae, about half as long as valva and parallel to valva axis. Aedoeagus cylin- drical, about 4 times as long as wide; vesica with two cornuti (Figs. 15—16), one approx- imately twice as large as the other, each bearing small pointed scales on one side; vesica devoid of microspines. Etymology.—The generic name _ is formed by adding the Greek meta (in its meaning of near) to Commotria, in refer- ence to the similarity in male genitalia. The name is not necessarily intended to imply close phylogenetic relationship. Gender: feminine. Metacommotria beckeri Shaffer, new species (Figs. 1, 4, 8, 11, 13-16) Diagnosis.—This is the only known spe- cies in the genus. VOLUME 105, NUMBER 2 285 Figs. 4-7. type. 6, Harnochina digitata, holotype. 7, Zapalla dentata, 3d, Alto Paraiso, Brazil; to show 3 spots on vein 1A+2A. Scale bar = 2 mm (6 & 7 to same scale). Description.—Male: Frons brownish pink dorsally, light brownish pink to yel- lowish white laterally. Labial palpus with basal segment white, second segment white to light brownish pink on ventral half of outer side, gradually becoming brownish pink on dorsal half of outer side: third seg- ment brownish pink. Vertex, occiput, pata- gium, and tegula light reddish brown, te- gula darker near wing base. Forewing (Fig. 4) length 8.0—9.5 mm (n = 5). Costa brown on basal third of wing, narrow but promi- nent band of yellowish white between costa and subcosta extends from wing base to apex. Broader band of nearly solid brown extending from wing base to apex, bounded anteriorly by subcosta, posteriorly by 1A+2A near wing base and by M, near apex; a second less prominent brown band between cell and 1|A+2A, this band diluted with yellowish pink scales. Band of yellow- Adults showing right wings. 4, Metacommotria beckeri, holotype. 5, Cipopeoria camura, holo- ish pink extending from distal half of cell to outer wing margin, bounded approxi- mately by M, and CuA,. Ground predomi- nantly yellowish pink posterior to L[A+2A. Band of yellowish white on cubitus, espe- cially prominent at lower outer angle of cell, extending onto M,,, and CuA,, poorly defined to absent on CuA,, prominent on 1A+2A. Dark brown spots between veins on outer wing margin. Genitalia as described for the genus. Holotype d.—Brazil, Espirito Santo, Linhares, 40 m, 05—09 April 1992, V. O. Becker; 82680; d genitalia on slide 2671, J. C. Shaffer [NMRJ]. Paratypes.—3 d, same data as holotype, one specimen J. C. Shaffer genitalia slide 2678 [USNM]; another, wing slide 2757; third undissected [NMRJ]. Other specimen examined.—d, Brazil, Rondonia, Vilhena, 600 m, 9 December 286 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 8-10. Harnochina digitata, holotype. Scale bar 1997, V-©: Becker] 111786; 9. €. Shatter genitalia slide 2679; [VOB]. Remarks—A female (J. Shaffer genitalia slide no. 2701) from Acailandia, Maranhao is essentially identical in wing maculation and venation and may be conspecific. Etymology.—The specific epithet honors the collector, Dr. Vitor Becker. Cipopeoria Shaffer, new genus Type species.—Cipopeoria camura Shaf- fer, new species. Diagnosis.—The male genitalia some- what resemble those of Homosassa Hulst, notably in that the lateral arms of the uncus are slender, straight, unbranched, tapering, and ventrally directed in a plane perpendic- ular to the body axis. The gnathos of Ci- popeoria \acks the stout lateral arms seen in Homosassa. Female genitalia of Cipo- peoria have a large signum, those of Hom- osassa lack a signum. The male labial palpi of Cipopeoria are slender and upturned (Fig. 9), those of Homosassa are broader and obliquely ascending (see Fig. 2, Shaf- fer, 1976b). Description.—Labial palpus of (Figs=3; male 9) with second segment slender, curved basally, upturned; of female porrect, about three times as long as eye diameter. Maxillary palpus moderately well devel- oped, extending just beyond eye in male, nearly to end of frons in female. Antenna Heads, left profile. 8, Metacommotria beckeri, holotype. 9, Cipopeoria camura, holotype. 10, 0.5 mm (8-10). shaft sublaminate in male, basal segments fused, posterior surface with sinus, sinus surface somewhat concave and bordered by parallel pair of short scale tufts; shaft in fe- male filiform, finely ciliate, lacking basal modification. Ocellus well developed, sep- arated from eye by about its own width. Forewing (Fig. 2) with eleven veins; R, well separated from base of R,,; latter from well before upper outer angle of cell; R, from about ! , and R,,, from just over 4 length of R,,;; M, from upper outer angle; M,,, stalked about Y,; its length, from lower outer angle; CuA,, from very near the an- gle; CuA, from well before the angle. Hind wing with seven veins (1A, 2A, and 3A counted as one vein); M,., fused, stalked with CuA, about 7, its length. Male genitalia (Figs. 17—20) with medial lateral process basally broad, subtriangular, distal- process of uncus weak, bandlike; ly slender, straight, unbranched, tapering, and ventrally directed in a plane perpendic- ular to the body axis. Gnathos a broad bi- lobed plate with a pair of anteriorly directed medial spines and along lateral margin of each lateral lobe a pair of ventromedially directed spines. Juxta horseshoe shaped, somewhat flared laterally. Valva with costa terminating in blunt tooth. Aedoeagus sub- cylindrical, 2.5 times as long as maximum width, tapering somewhat posteriorly; ve- sica with a pair of prominent subequal ser- VOLUME 105, NUMBER 2 Figs. 11-12. 287 Male right antennae, basal region, partially denuded. 11, Metacommotria beckeri, holotype. 12, Cipopeoria camura, holotype. Scale bar = 0.2 mm (11, 12). rate cornuti (Figs. 19—20) and a few minute teeth. Female genitalia (Figs. 23—24) with ovi- positor lobes finely setose, dorsal and pos- terior elements meeting at 160° angle, weakly sclerotized. Apophyses slender; posterior apophysis weakly curved, about ¥, length of anterior apophysis; anterior apophysis straight. Ovipositor separated lit- tle, if any, from eighth abdominal segment, the latter heavily sclerotized, its posterior 7; strongly folded. Ostium bursae membra- nous. Posterior half of ductus bursae well sclerotized, considerably flattened, smooth; anterior half membranous, with scattered minute triangular teeth. Corpus bursae membranous, with large irregular platelike signum bearing numerous inwardly directed spines. Ductus seminalis from tapering pos- terior end of corpus bursae (Fig. 23, arrow). Etymology.—The generic name is de- rived from the name of the type locality of the type species combined with a reference to the Tribe Peoriini. Gender: feminine. Cipopeoria camura Shaffer, new species (Figs. 2, 3, 5, 9, 12, 17—20, 23-24) Diagnosis.—This is the only known spe- cies in the genus. Description.—Frons grayish reddish brown dorsally, yellowish white ventrally. Labial palpus with first and second seg- ments grayish reddish brown dorsally, yel- lowish white ventrally and laterally except grayish reddish brown near dorsal margin and with a few scattered scales of that color elsewhere; third segment pale orange yel- low throughout. Vertex grayish reddish brown; yellowish white posterior and pos- teriolaterally from antenna base. Occiput grayish reddish brown dorsally, similar be- hind eye, but interrupted with narrow bars of yellowish white. Patagium and tegula light grayish reddish brown, darker at wing 288 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 13-22. Aedoeagus. 15, removed. 18, Aedoeagus. 19, Small cornutus. 20, Large cornutus. 21—22, Harnochina digitata, holotype. 21, Genitalia, aedoeagus removed. 22, Aedoeagus. Scale bar = 0.5 mm (13-14, 17-18, 21-22), 0.1 mm (15-16, 19-20). base. Forewing (Fig. 5) length 6.0—11.0 mm (n = 7). Costa grayish reddish brown at base, light orange yellow distally. Ground white anterior to cubitus, heavily sewn with a mixture of grayish reddish brown and dark brown scales, lightest on distal half of wing. Ground light brown posterior to cell and near outer wing mar- Male genitalia. 13-16, Metacommotria beckeri, holotype; 13, Genitalia, aedoeagus removed. 14, Large cornutus. 16, Small cornutus. 17-20, Cipopeoria camura, holotype. 17, Genitalia, aedoeagus gin, a large irregular dark brown patch be- tween 1A+2A and cell, extending posterior to |A+2A. Terminal line of dark brown spots between veins. All wing markings dif- fuse, ull defined; no veins with white trac- ing. Genitalia as described for the genus. Holotype o6.—Brasil, Minas’ Gerais, VOLUME 105, NUMBER 2 1.400 m, Serra do Cipo, 17-19 April 1991, V. O. Becker; no. 78056; 3 genitalia on slide 2672 J. C. Shaffer [NMRJ]. Paratypes.—2 d, same data as holotype; no. 78056, genitalia slide 2680 [USNM]: no. 78052, genitalia slide 2694. 2 2, same data as holotype: no. 78053, genitalia slides 2690, 2704 [NMRJ]. Other specimens examined (1 d, 1 2 ).— 3, Nova Lima, Minas Gerais, 25-27 De- cember 1982, V. O. Becker; no. 50267; ¢ genitalia on slide 2670. 2, Bonito, Bahia, 1,000 m, 25 April 1991, V. O. Becker; no. 78464; not dissected [VOB]. Etymology.—The specific epithet is de- rived from the Latin camur (turned inward) in reference to the inward curving hooks of the gnathos. Harnochina digitata Shaffer, new species (Figs. 6, 10, 21-22) Diagnosis.—The division of the distal portion of the valva of the male genitalia into two slender digitate processes, one membranous, the other a heavily sclerotized extension of the costa is unique within the genus, as is the V-shaped medial process of the gnathos. In the only other known spe- cies in the genus, H. rectilinea Dyar, the distal half of the valva (see Shaffer 1976a, fig. Ile) is angled relative to the basal half, broadly triangular, the apex rounded and simple; the medial process of the gnathos unbranched, forming a tapering spine. Description.—Male: Labial palpus (Fig. 10) short, about 1.9 times as long as eye diameter; upturned, third segment porrect. Maxillary palpus short (not clearly seen in holotype). Antenna shaft subserrate, cilia about half as long as segment width near base of shaft; basal segments unmodified, sinus absent. Forewing length 8.5 mm (n = 1); with eleven veins; R, well before upper outer angle of cell; R, free from cell, near to origin and closely parallel to R;.;; R3., stalked with R; about 7% its length, from be- fore the angle; M, from the angle; M,,; stalked about half its length, from lower outer angle; CuA, from well before the an- 289 gle; CuA, separated from CuA, by about same distance. Ground overall light grayish brown, produced from a mixture of brown- ish pink and dark brown scales on a field of yellowish white scales. Markings darker than ground, diffuse, somewhat indistinct: small discal spot between base of M, and base of M,,,; antemedial, postmedial, ter- minal bands; fringe with two light and two dark bands. Hind wing with seven veins (1A, 2A, and 3A counted as one vein); M3, ; fused, stalked with CuA, about 7% its length: from lower outer angle of cell. Genitalia (Figs. 21-22) with medial pro- cess of uncus bearing a pair of smooth tri- angular posterior projections, more distally and dorsally a pair of irregular setose pro- jections; lateral processes a pair of knobbed arms, these setose on posterior margin, smooth elsewhere. Gnathos with medial process a weakly sclerotized plate with a pair of divergent anterior arms and a well sclerotized V-shaped process, its apex pos- teriorly directed; lateral arms of gnathos broad, flat, weakly sclerotized. Juxta a semilunar plate, posterior margin emargin- ate. Valva subquadrate, ventral half covered with still anteriorly directed setae; costa heavily sclerotized, smooth, the free distal portion about 7 length of entire costa; membranous digitate process closely par- allel to free portion of costa. Anterior mar- gin of vinculum truncated. Aedoeagus about three times as long as maximum width, somewhat constricted on posterior half; vesica unarmed, lacking microspines. Holotype ¢.—Brazil, Goids, Alto Parat- so, 1,400 m, 1-6 November 1996, V. O. Becker; 98791; d genitalia on slide 2683, J. C. Shaffer [NMRJ]. Other specimens examined.—None. Etymology.—The specific epitaph is de- rived from the Latin digitus in reference to the distal digitate extension on the valva of the male genitalia. Zapalla dentata Shaffer (Figs. 7, 25) Zapalla dentata Shaffer 1976a: 322, 3h; 61, 9f. 290 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 23-25. origin. 24, Enlarged to show signum. 25, Zapalla dentata, genitalia (corpus bursae bearing spermatophore), arrow shows ductus seminalis origin. Scale bar = 0.5 mm (23, 25), 0.2 mm (24). Previously known only from the male holotype (type locality: ““S.E. Brazil”), the Becker material includes specimens from Planaltina (Distrito Federal), Alto Paraiso (Goias), and Chapada Guimaraes (Mato Grosso). Specimens from the latter locality comprise three males and one female (J. Shaffer slide 2700) which forms the basis for the following description. Description.—Male (see Shaffer, 1976). Female: Labial palpus porrect, about 3.5 times as long as eye diameter. Maxillary palpus very short, extending little if at all beyond pilifer, yellowish white. Eye diameter 0.55 mm. Ocellus well de- veloped, black with clear center, separated from eye by about its own diameter. Antenna Female genitalia. 23-24, Cipopeoria camura. 23, Genitalia, arrow shows ductus seminalis filiform, very finely ciliate. Genitalia (Fig. 25) with ovipositor lobes finely setose; upper element a slender well sclerotized rod, ver- tical element a diffuse lightly sclerotized plate about 3.75 times as high as wide. Apophyses straight or nearly so, anterior apophysis about 1.3 times as long as poste- rior apophysis. Ostium bursae membranous. Ductus bursae moderately well sclerotized, extensively folded longitudinally, unarmed; joining corpus bursae near its posterior end. Corpus bursae about twice as long as wide; unarmed, lacking signum and microspines [Fig. 25 shows remnants of a single sper- matophore nearly filling the corpus bursae]. Ductus seminalis from tapering posterior end of corpus bursae (Fig. 25, arrow). VOLUME 105, NUMBER 2 Remarks.—This is the first female known for the genus. The genitalia are generalized and show no apparent apomorphies. Exter- nally dentata and deliquella (Zeller) are readily separated by the number of dark spots on vein 1A+2A, two spots in deli- quella, three spots in dentata. An adult male of dentata is illustrated herein (Fig. 7) as the photograph (fig. 3h) in Shaffer 1976a does not show the third (middle) spot clearly. ACKNOWLEDGMENTS I am most grateful to Vitor Becker, Bra- silia, Brazil, for the loan of specimens from his personal collection, and thank Alma So- lis (USDA, Systematic Entomology Labo- 291 ratory) and an anonymous reviewer for helpful comments on the manuscript. LITERATURE CITED Kelly, K. L. 1965. ISCC-NBS color-name charts illus- trated with centroid colors. Standard Sample No. 2106. Supplement to National Bureau of Stan- dards Circular 553. 4 pp. (unnumbered), 18 sheets color samples. U.S. Government Printing Office. Washington, DC. Shaffer, J. C. 1976a. A Revision of the Neotropical Peoriinae (Lepidoptera: Pyralidae). Systematic Entomology |: 281-331, Figs. 1-13, Maps 1-5, Plates 1-16. . 1976b. Two new species of Peoriinae (Lepi- doptera: Pyralidae) from Texas. Proceedings of the Entomological Society of Washington 78(4): 431— 434. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 292-299 REVIEW OF THE NEW WORLD LEAFHOPPER GENUS EXCULTANUS OMAN (HEMIPTERA: CICADELLIDAE: DELTOCEPHALINAE) STUART H. MCKAMEY Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, “ National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: smckamey @sel.barc.usda.gov) Abstract.—The taxonomy of the New World genus Excultanus is reviewed. Excultanus argillaceus (Van Duzee) is placed under EF. excultus (Uhler), n. syn. Excultanus para- conus, n. sp. is described from Cost Rica. The known southward limit of the genus is extended from Guatemala to Misiones, Argentina. A key to the 12 species is provided, as well as a checklist with critical synonymies and distribution summaries. Key Words: About 150 of the 350 genera of U.S. leaf- hoppers have species in Mexico, yet only about 200 species occur in both countries. Furthermore, the number of leafhoppers known from the U.S. (approximately 3,000) is more than double the number known from Mexico, despite its relatively large size and topographical complexity. The sit- uation for Central America is similar: 80 genera but only 65 leafhopper species are shared with the U.S., and fewer than 900 Central American species have been de- scribed. These figures suggest two impor- tant generalizations: (1) the Mexican and Central American faunas must be taken into account to provide comprehensive identifi- cation tools for U.S. leafhopper genera; and (2) the Mexican and Central American fau- nas are poorly known and likely contain several thousand leafhopper species new to science, including many belonging to gen- era that also occur in the United States. The genus Excultanus is one such genus. Excultanus the United States, Mexico, Central America, from which one new species is here described, and South America. While the genus includes a known occurs in taxonomy, vector, Athysanini, Excultanus vector of potential importance to California celery crops [1.e., E. incurvatus (Osborn & Lathrop); Nielson 1968b], there have been no synopses of the whole genus. Literature citations designated with key letters are consistent with the bibliographies by Metcalf (1964a) and Oman et al. (1990). REVIEW OF LITERATURE Oman (1949a) erected the new subgenus Texananus (Excultanus) for eight species having a broadly excavated sternum VII in the female and lacking a posterior extension of the connective in the male genitalia. Crowder (1952a) provided a key to the four species that occur in the United States. Linnavuori (1959b) elevated Excultanus to generic status, the genus being distinguish- able from other Philepsius-like genera by the absence of a posterior projection of the connective below the aedeagus. When Lin- navuori (1959b) elevated Excultanus to ge- neric status, he treated only the Central American species but clearly intended it to include the Nearctic species also treated un- der the genus-group name, by citing Oman (1949a). Likewise, Nielson (1968b) explic- VOLUME 105, NUMBER 2 itly accepted the generic rank of Excultanus but mentioned only E. incurvatus, the sin- gle species known to vector phytopatho- gens. McKamey (2000) affirmed the placement of the three species already included in Ex- cultanus and referred nine other species to the genus: six previously in Texananus (Ex- cultanus) and three [E. parrai (DeLong), E. plummeri (DeLong) and E. horridus (DeLong)] from Texananus but previously unplaced to subgenus, bringing the total number of species to 12. One new species is described and one older species is placed in synonymy in this paper. MATERIALS AND METHODS Specimens for the study are from the Na- tional Museum of Natural History, Smith- sonian Institution, Washington, DC (NMNH); the Instituto Nacional de Biodiv- ersidad, Santo Domingo de Heredia, Costa Rica (INBIO); the DeLong collection of Ohio State University, Department of En- tomology, Columbus, Ohio, USA (OSUC): and the California Academy of Sciences, San Francisco, California (CASC). The pygofers and genitalia were prepared by briefly boiling the abdomen in a weak potassium hydroxide (KOH) solution. KOH-treated parts were preserved in glyc- erin in polypropylene microvials beneath the specimens. Digital images were captured at 400 dpi resolution by mounting a Sony DKCS5000° digital camera on a Leica MZ-APO™ ste- reoscopic microscope (for the pygofer) or Leitz Diaplan™ compound microscope (for the aedeagus). The ventral and dorsal margins of the male pygofer of some species [e.g., E. co- nus (DeLong)] are softer than surrounding areas of the pygofer and curl inwards if the abdomen remains in KOH too long. Hence, DeLong’s (1939c, 1944h) description of the pygofer of E. conus being ‘“‘curled inwardly on ventral side, appearing tapered to a pointed apex,” while consistent with his il- lustration (1944h, fig. 3), is an artifact of 293 preparation. Brief treatment in KOH, such as the method used here, does not curl the pygofer margins. RESULTS Genus Excultanus Oman Texananus (Excultanus) Oman 1949a: 142. Type species by original designation: Jassus excultus Uhler 1877. Excultanus; Linnavuori 1959b: 197 [elev. status |. Diagnosis.—Differing from other Delto- cephalinae in having head narrower than pronotum (Fig. 1); forewing with appendix well developed; pronotum and usually fore- wing with vermiculate pigment lines (pl. 39, fig. | of Oman 1949a); subgenital plates with setae; male with genital connective lacking posterior projection below aedea- gus; female sternum VII deeply excavated [except E. hebraeus (Ball)] and often with portion of underlying membranes visible. Resembles Texananus and other Phlepsius- like genera in having forewing covered with ramose or vermiculate pigment lines (Fig. 1). Notes.—Illustrations of the female ster- num VII and male pygofer and genitalia for most species were provided by DeLong (1944h) and Crowder (1952a). KEY TO SPECIES OF EXCULTANUS 1. Male pygofer less than or equal to length of subgenital plates; female sternum VII with portion of posterior underlying membranes visible, if excavated almost to base and with minute median projection (£. conus), then sternum VII length approximately *%4 its max- LUAU Wi Le een Pettey. te eco teiiees! ca: sonar 2 — Male pygofer extending beyond subgenital plates; female sternum VII with portion of posterior underlying membranes not visible or, if visible (E. dorothyae and E. eugeneus), then triangular and attaining posterior margin of sternum VII or excavated almost to base, with minute median projection and sternum VII length approximately % its maximum Wid th peeps sacs ch Aki ctcs Yee 2s 12 Pa Bre Ns. fee to % 6 2. Male with anal segment X strongly produced ventrally, hanging within pygofer; female sternum VII deeply excavated, margins of ex- cavation notched at mid length... . . plummert PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Male with anal segment X normal, ringlike, not produced ventrally; female sternum VII deeply excavated without notch at mid length along margins, or not excavated and with me- AianMOtch’ yo \oe plenanesopewagens) oceans 3 Male pygofer with conspicuous inner brushes of darkened bristles distally on dorsal and ventral margins; male pygofer length in lat- eral view about 1.5 < height; female sternum VII deeply excavated and underlying mem- branesinot; visible’ -ecereeneen teens. horridus Male pygofer without inner brushes of bristles on margins; male pygofer in lateral view about as long as tall; female sternum VII deeply excavated and underlying membranes visible or if membranes not visible (E. he- braeus), then not excavated and with median NOCHE cet mee neg Rete piatigchays (od-toge ee eee 4 . Male pygofer apex bluntly rounded; female sternum VII with visible portion of posterior underlying membranes rectangular, extending from median excavation to lateral margin Lee AOL) halt kee dorothyae Male pygofer apex truncate; female sternum VIL underlying membranes either not visible or with visible portion oval and lying entirely within deep excavation (as in Fig. 5) .... . Male pygofer apex bilobed dorsoposteriorly; aedeagal shaft extending nearly to dorsal mar- gin of pygofer; female sternum VII not ex- cavated, with median notch, underlying mem- an branes nOteviIStDlem cco < ae arieye ete ore hebraeus Male pygofer apex single-lobed; aedeagal shaft extending dorsally to about mid height of pygofer; female sternum VII deeply exca- vated and with small median projection, un- derlying membranes visible ........ eugeneus Male pygofer without distal brush of bristles on ventral margin or apex; male pygofer in lateral view with distal half forming an equi- lateral triangle; female sternum VII sinuously excavated and smoothly concave medially with underlying pair of membranes triangular FOO RO OrO Sa eto o rere See Ar 6 excultus Male pygofer with conspicuous inner brush of darkened bristles distally (Fig. 2), though not always at apex; male pygofer in lateral view with distal half length greater than pygofer height, not forming an equilateral triangle; fe- male sternum VII excavation either not sinu- ous or bearing acute notch medially, with un- derlying pair of membranes oval (Fig. 5) .. 7 Male pygofer with inner brushes of bristles on dorsal and ventral margins; female sternum VII excavated approximately halfway to base with acute, V-shaped median notch ......... parrat Male pygofer with inner brushes of bristles on ventral margins or at apex, or both (Fig. 2), but not on dorsal margins; female sternum VII excavated at least % to base, excavation medially either smoothly concave or with smalliimedian projection’ 2-2 2. «-eise sce 8 Male pygofer with inner brushes of bristles on ventral margins, sometimes continuing to apex; female sternum VII length at least % its maximum width, posterior angles acute (Fig. 5) or excavation with small median projec- tion, or with both conditions ........... 9 — Male pygofer with inner brushes of bristles at apex only; female sternum VII length approx- imately 4 its maximum width, posterior an- gles truncate or rounded, excavation without Medians projechlon are aie eee 11 9. Male pygofer apex weakly sclerotized, with- out bristled fringe; female sternum VII with posterior angles rounded, excavated almost to oo base and with distinct median projection . . . Monsbeusheleneesewehor beta rcke ewan e theeae paralus — Male pygofer with strong sclerotization and bristle-fringed ventral margins continuing to apex; female sternum VII with posterior an- gles acute, excavation depth variable, some- times with minute median projection .... 10 10. Male pygofer with bristle-fringed ventral mar- gins linear (whether curled inwardly in KOH- prepared specimens or not curled inwardly); female sternum VII excavated almost to base Pac ye PONCE CO fe. -Obrett O80 san OMCLAT oo Gecko OHO conus — Male pygofer with bristle-fringed ventral mar- gins sinuous (Fig. 2); female sternum VII ex- cavated about % its length (Fig. 5) ..... Serio Cee oon Dat deetto iy Chon ac paraconus, 1.Sp. 11. Male subgenital plates triangular, connective anterior arms V-shaped, and style with preap- ical concavity U-shaped; length including wings in repose 5.7—6.2 mm; female sternum VII with visible portions of underlying mem- branes about as long as wide ...... incurvatus — Male subgenital plates subtriangular but weakly attenuate on lateral margins, connec- tive anterior arms U-shaped, and style with preapical concavity V-shaped; length includ- ing wings in repose 7.0—8.0 mm; female ster- num VII with visible portions of underlying membranes about twice as long as wide Beta ouchcye MMe Reet N ie cl eres aie neomexicanus Excultanus paraconus McKamey, new species (Figs. 1-5) Diagnosis.—Male pygofer with ventral margins sinuous and bearing a brush of dense bristles; female sternum VII with posterior angles acute, excavated posteri- orly for about % of its length. VOLUME 105, NUMBER 2 Figs. 1-5. ventral view, Excultanus paraconus. 1, Male, dorsal view. 2, Male pygofer, right portion of ventral margin, showing bristle-fringed, sinuous ventral margin. 3, Aedeagus, connective, and styles, dorsal view. 4, Aedeagus, lateral view. 5, Female sterna, ventral view, showing excavated sternum VII with visible ovoid portions of underlying membranes. Description.—Measurements (mm). Length: male 6.6—7.0; female 7.0—7.6. Width across eyes: male 1.9, female 1.9— 210) Male and female similar externally. Fron- toclypeus and pronotum darkly mottled fus- cous, vertex and scutellum paler, forewing covered throughout with vermiculate infus- cate marks (Fig. 1). Abdomen: Male. Py- gofer heavily sclerotized throughout, trian- gular in lateral view, covered with macro- setae, ventral margins (Fig. 2) sinuous in ventral view, with dense fringe of dark bris- tles, which are longest at convexities, al- most absent at concavity, pygofer length exceeding subgenital plates by half length of their mesal margins; subgenital plates tri- angular, lateral margins straight. Connective (Fig. 3) Y-shaped, anterior arms subparallel and separated from each other by approxi- mately their length. Aedeagus (Fig. 4) ar- ticulated to connective, U-shaped, gradually tapering from base to apex; style (Fig. 3) with broad base quickly tapered, at mid length abruptly narrowed to slender distal ¥3. Female sternum VII (Fig. 5) with pos- terior angles acute, excavated posteriorly for about *% its length. Types.—Holotype ¢ [USNM] COSTA RICA: Puntarenas Prov., Puntarenas, 12- 296 VII-1972, J. Maldonado C[apriles]. Para- types [USNM]: COSTA RICA: 9 mi. NW Esparta, 22-VII-1965, 1 3d, 1 @. Paratypes [INBIO]: COSTA RICA: Guanacaste Prov., Finca Jenny, 30 km N. Liberia, Guanacaste National Park, [X-1989, R. Espinoza, 8 d 3 2, INBIO Bar Code No. CRI000025028, CRI000025546, CRIO00152442, CRI- 000476339, CRI000476749, CRI00047674 1, CRI000025219, CRI000476734, CRI- 000476733, CRIO00025606, CRI000476725; same locality, X-1989, E. Araya & R. Espi- noza, 3 ¢ 4 2, INBIO Bar Code No. CRI- 00008 1664, CRIO00108821, CRIO000109428, CRIO00130815, CRI000131397, CRI- 000130888, CRIO00131051; same locality, XI-1989, 1 ¢, E. Araya & R. Espinoza, IN- BIO Bar Code No. CRI000138115; Estacion Palo Verde, elev. 10 m, 12-23-XI-1991, R. U. Chavarria & D. Acevedo, Malaise 1 °, INBIO Bar Code No. CRIO002800906, same locality 12 VII-12 IX-1991, 1 2, INBIO Bar Code No. CRIO02800983; Estacion Experi- mental E. Jimenez Nunez, 30-VI-1993, G. Allen, Malaise 6:00 pm—8:00 am, 1 @, IN- BIO Bar Code No. CRI002088831; same lo- cality and nocturnal Malaise, 12-VHI-1993, | 2, INBIO Bar Code No. CRI002072231; same locality and nocturnal Malaise, 15-VUI- 1993, 2 ¢6, 2 2, INBIO Bar Code No. CRI- 002088173, CRIO01951879, CRI002072694 and CRI002072719; Parque Nacional Barra Honda, elev. 100 m, VIII 1994, M. Reyes, 1 2, CRIOO2005809. Note.—One difference between E. par- aconus and E. conus may be the strong cur- vature of the aedeagus in the latter. The var- iation of aedeagal curvature observed else- where in the genus when there are more specimens, for example in E. excultus, sug- gests that this is not as reliable a feature to distinguish the species. The distinctive ven- tral margin of the male pygofer in E. par- aconus, on the other hand, appears consis- tent and can usually be observed in undis- sected specimens. Etymology.—The specific epithet (adjec- tive) is formed by adding the Greek prefix PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON “para-,” for near, to the name of the spe- cies it resembles most, Excultanus conus. CHECKLIST AND DISTRIBUTION SUMMARY FOR OTHER SPECIES OF EXCULTANUS E. conus (DeLong) Texananus conus DeLong 1939c: 386 [n.sp.]. Texananus (Excultanus) conus; Oman 1949a: 142 [subgen. comb.]. Excultanus conus; McKamey 2000: 214 [n.comb.]. Summary of distribution.—MEXICO: Chiapas. NICARAGUA. Note.—Maes et al. (1999) reported E. co- nus from Nicaragua. E. dorothyae (DeLong) Texananus dorothyi [sic] DeLong 1939c: 387 [n.sp.]. Texananus (Texananus) dorothyae; DeLong & Hershberger 1949a: 173 [subgen. comb.]. Texananus (Excultanus) dorothyae, Oman 1949a: 142 [subgen. comb.]. Excultanus dorothyae; McKamey 2000: 214 [n.comb.]. Summary of distribution —MEXICO: Guerrero, Nuevo Leon, Puebla, Morelos, San Luis Potosi. UNITED STATES: Ari- zona, New Mexico, Texas. Note.—In the original description, the species was explicitly dedicated to Dorothy Johnson Knull, so the correct spelling of the name is as used by DeLong & Hershberger (1949a) and others, including this work. E. eugeneus (Ball) Phlepsius (Texananus) eugeneus Ball 1918b: 384 [n.sp.]. Lectotype 6 USNM. Texananus eugeneus; DeLong & Cald- well 1937c: 43 [n.comb.]. Texananus (Excultanus) eugeneus: Oman 1949a: 142 [subgen. comb.]. Excultanus eugeneus; McKamey 2000: 214 [n.comb.]. VOLUME 105, NUMBER 2 Summary of distribution.—COSTA RICA: Guanacaste, Finca Jenny, 30 km N. Liberia, Guanacaste National Park, IX- 1989, R. Espinoza, INBIO Bar Code No. CRION00024958 and CRIO00025105. MEX- ICO: Guerrero, Jalisco, Morelos. Notes.—This species was originally de- scribed from 12 syntypes. Previously re- corded only from Mexico, the known range is here extended to Costa Rica. A syntype from the USNM is here designated as the lectotype to stabilize the nomenclature of the species. The lectotype has the following label data: “*Amula/Guerrero,/6000 ft./Sept. He He Smith, = DYPE ~Cotype INo:/ [blank]/U.S.N.M.,” “‘Phlepsius/eugeneus/ Ball,” and a lectotype label. E. excultus (Uhler) Jassus excultus Uhler 1877a: 467 [n.sp.]. Phlepsius excultus; Southwick 1892a: 288 [n.comb. ]. Phlepsius (Texananus) excultus; Ball 1918b: 386 [subgen. comb.]. Phlepsius argillaceus Van Duzee 1923a: 185 [n.sp.; holotype ¢ CASC], new synonymy. Texananus agrillaceus {sic}; DeLong & Caldwell 1937c: 43 [n.comb. ]. Texananus excultus; DeLong & Cald- well 1937c: 43 [n.comb.]. Texananus (Texananus) excultus: DeLong & Hershberger 1949a: 176. Texananus (Excultanus) agrillaceus [sic]; Oman 1949a: 142 [subgen. comb. |. Texananus (Excultanus) excultus; Oman 1949a: 142 [subgen. comb. ]. Excultanus excultus; Linnavuori 1959b: 198 [n.comb.]. Excultanus agrillaceus [sic]; Mc- Kamey 2000: 214 [n.comb.]. Summary of distribution —ARGENTI- NA: Misiones. BAHAMAS. BELIZE. EL SALVADOR. GUATEMALA: San_ José. MEXICO: Baja California, Chiapas, Jalis- co, Sonora, Veracruz. UNITED STATES: 297 Alabama, Arizona, California, Florida, Georgia, Louisiana, New Mexico, North Carolina, South Carolina, Texas. Notes.—The specimens from Argentina are identical with respect to all features ex- cept the aedeagal apex is almost in contact with the base, i.e., they are the closest to the O-shape in the U- to O-shape variation in this species. Also, although the species was already the most widespread, it is even more dispersed than previously thought, from the United States and Bahamas to Ar- gentina (USNM specimen). The holotype of argillaceus was examined and dissected; the aedeagus is U-shaped with a weakly de- flexed apex, also falling within the variation of E. excultus. The type locality of argil- laceus, Baja California, is consistent with the other records of excu/tus in northern Mexico. E. hebraeus (Ball) Phlepsius (lowanus) hebraeus Ball 1918b: 383 [n.sp.]. Texananus hebraeus; DeLong & Cald- well 1937c: 44 [n.comb.]. Towanus hebraeus,; DeLong & Hersh- berger 1948d: 115 [n.comb.]. Excultanus hebraeus; Linnavuori 1959b: 198 [n.comb.]. Summary of distribution —GUATEMA- LA: Guatemala City. MEXICO: Chiapas, Guerrero. Note.—This species was described from eight females, including one from the Vi- enna Museum that was not examined in this study. A syntype from the USNM is here designated as the lectotype to stabilize the nomenclature of the species. The lectotype has the following label data: ‘*Amula/Guer- rero,/6000 ft./Sept. H. H. Smith,” “TYPE,” “Cotype No./[blank]/U.S.N.M.,” ‘‘Phlep- sius/hebraeus/Ball,”’ and a lectotype label. E. horridus (DeLong) Texananus horridus DeLong 1944h: 234 [n.sp.]. Excultanus horridus,; McKamey 2000: 214 [n.comb.]. 298 Summary of distribution.—MEXICO: Guerrero, Oaxaca. E. incurvatus (Osborn & Lathrop) Phlepsius (lowanus) incurvatus Os- born & Lathrop 1923a: 346 [n.sp.]. Texananus incurvatus; DeLong & Caldwell 1937c: 44 [n.comb.]. Texananus (Texananus) incurvatus: DeLong & Hershberger 1949a: 178 [subgen. comb.]. Texananus (Excultanus) incurvatus; Oman 1949a: 142 [subgen. comb.]. Excultanus incurvatus; Nielson 1968b: 285 [n.comb.]. Summary of distribution —MEXICO: Jalisco, Sonora. UNITED STATES: Arizo- na, California, Georgia, New Mexico. Note.—This species, a vector of western strain of North American aster yellows vi- rus, is potentially important in the spread of this disease in celery in California. E. neomexicanus (Baker) Phlepsius neomexicanus Baker 1895b: 13 [n.sp.; holotype ¢ USNM]. Phlepsius (lowanus) neomexicanus; Osborn & Lathrop 1923a: 346 [sub- gen. comb.]. Phlepsius (Texananus) neomexicanus; Ball 1931g: 85 [subgen. comb.]. Texananus neomexicanus; DeLong & Caldwell 1937c: 44 [n.comb.]. Texananus (Texananus) neomexican- us; DeLong & Hershberger 1949a: 176. Texananus (Excultanus) neomexican- Oman 1949a: 142 [subgen. comb. |. Excultanus neomexicanus; McKamey 2000: 214 [n.comb.]. us; Summary of distribution.—MEXICO: Sinaloa. UNITED STATES: Arizona, New Mexico, Texas. E. paralus (DeLong) Texananus paralus DeLong 1939c: 385 [n.sp.]. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Texananus (Excultanus) paralus; Oman 1949a: 142 [subgen. comb.]. Excultanus paralus; McKamey 2000: 214 [n.comb.]. Summary of distribution. —MEXICO: Chiapas, Guerrero, Michoacan, Morelos. E. parrai (DeLong) Phlepsius parrai DeLong 1939c: 382 [n.sp.; holotype d OSUC]. Texananus parrai; DeLong 1944h: 234 [n.comb.]. Excultanus parrai; McKamey 2000: 214 [n.comb.]. Summary of distribution. —MEXICO: Guerrero. E. plummeri (DeLong) Texananus plummeri DeLong 1939c: 385 [n.sp.; holotype ¢ OSUC]. Texananus cassus DeLong 1939c: 385 [n.sp.; holotype ¢ OSUC]. Synon- ymy by DeLong 1944h: 233. Excultanus plummeri; McKamey 2000: 214 [n.comb.]. Summary of distribution —MEXICO: Chiapas, Guerrero, Michoacan, Morelos. Note.—Excultanus plummeri and its syn- onym Texananus cassus represent opposite sexes of the same species, of which a mixed series (males of females) was later collect- ed. ACKNOWLEDGMENTS For lending specimens I am grateful to Carolina Godoy (INBIO), Andrey Sharkov (OSUC), and Keve Ribardo (CASC). The manuscript was improved by suggestions from P. Freytag (University of Kentucky), T. Henry, D. Smith (both USDA/ARS Sys- tematic Entomology Laboratory) and two anonymous reviewers. LITERATURE CITED Baker, C. EF 1895b. New Homoptera received from the New Mexico Agricultural Experiment Station. I. Psyche 7(suppl. 1): 13-14. Ball, E. D. 1918b. The phlepsids of Mexico and Cen- VOLUME 105, NUMBER 2 tral America (Homoptera Cicadellinae). Annals of the Entomological Society of America 11: 381— SE . 1931g. New species of Phlepsius with notes on others (Homoptera, Cicadellidae). Pan-Pacific Entomologist 8: 85-89. Crowder, H. W. 1952a. A revision of some Phlepsius- like genera of the tribe Deltocephalini (Homop- tera, Cicadellidae) in America north of Mexico. University of Kansas Science Bulletin 35: 309— 541. DeLong, D. M. 1939c. Fauna Mexicana.—Il. (Insecta Homoptera, familia Cicadellidae, genus Phlep- sius). Los phlepsidos (Phlepsius y Texananus) de Mexico (Homoptera-Cicadellidae). Anales de la Escuela Nacional de Ciencias Biologicas 1: 379— 405. . 1944h. The Mexican species of leafhoppers of the genus Texananus (Homoptera: Cicadelli- dae). Journal of the Washington Academy of Sci- ences 34: 228-239. DeLong, D. M. and J. S. Caldwell. 1937c. Check List of the Cicadellidae (Homoptera) of America, North of Mexico. Ohio State University Press, Co- lumbus, iv + 93 pp. DeLong, D. M. and R. V. Hershberger. 1948d. A syn- opsis of the genus /owanus in North America in- cluding Mexico. Ohio Journal of Science 48: 107— LAS: . 1949a. A review of the North American spe- cies of Texananus (Homoptera—Cicadellidae) north of Mexico. Ohio Journal of Science 49: 173-187. Linnavuori, R. 1959b. Revision of the Neotropical Deltocephalinae and some related subfamilies (Homoptera). Annales Zoologici Societatis Zool- ogicae Botanicae Fennicae *Vanamo’ 20(1): [i-ii], 1-370. Maes, J. M., C. Godoy, and P. H. Freytag. 1999. Nue- vos reportes de Cicadellidae (Homoptera) de Nic- aragua. Revista Nicaraguense de Entomologia 49: 9-46. 299 McKamey, S. H. 2000. A review of the New World leafhopper subgenus Texananus (Jowanus Ball) (Hemiptera: Cicadellidae: Deltocephalinae) with a checklist and distribution summary for species in the genus. Proceedings of the Entomological So- ciety of Washington 102(1): 212-224. Metcalf, Z. P. 1964a. General Catalogue of the Ho- moptera. Fascicle VI. Cicadelloidea. Bibliography of the Cicadelloidea (Homoptera: Auchenorrhyn- cha). United States Department of Agriculture, Agriculture Research Service, 349 pp. Nielson, M. W. 1968b. The Leafhopper Vectors of Phytopathogenic Viruses (Homoptera, Cicadelli- dae). Taxonomy, Biology, and Virus Transmis- sion. United States Department of Agriculture Technical Bulletin No.1382, 386 pp. Oman, P. W. 1949a. The Nearctic Leafhoppers (Ho- moptera: Cicadellidae): A Generic Classification and Check List. Memoirs of the Entomological Society of Washington No. 3, [v] + 253 pp. Oman, P. W., W. J. Knight, and M. W. Nielson. 1990. Leathoppers (Cicadellidae): A Bibliography, Ge- neric Check-list and Index to the World Literature 1956-1985. CAB International Institute of Ento- mology, Wallingford, UK. [iv] + 368 pp. Osborn, H. and F. H. Lathrop. 1923a. The genus Phlep- sius in North America (Homoptera). Annals of the Entomological Society of America 16: 310-350. Southwick, J. EF 1892a. Notes on local Jassidae. Sci- ence 19: 287-288. Uhler, P. R. 1877a. Report upon the insects collected by P. R. Uhler during the explorations of 1875, including monographs of the families Cydnidae and Saldae and the Hemiptera collected by A. S. Packard, Jr. Bulletin of the United States Geolog- ical and Geographic Survey of the Territories 3(2): 355-475. Van Duzee, E. P. 1923a. Expedition of the California Academy of Sciences to the Gulf of California in 1921. The Hemiptera (true bugs, etc.). Proceed- ings of the California Academy of Sciences 12(11): 123-200. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 300-303 A NEW SPECIES OF CIRROSPILUS WESTWOOD (HYMENOPTERA: EULOPHIDAE) FROM SOUTHEASTERN ARIZONA MICHAEL W. GATES AND MICHAEL E. SCHAUFF Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: mgates @sel.barc.usda.gov; mschauff @sel. barc.usda.gov) Abstract.—Cirrospilus infuscatus, n. sp. (Hymenoptera: Eulophidae), is described from southeastern Arizona. This species was reared from Tischeria bifurcata Braun (Lep- idoptera: Tischeriidae) on Ceanothus fendleri A. Gray (Rhamnaceae). Key Words: Species of the cosmopolitan genus Cir- rospilus Westwood. parasitize cryptically- feeding insects, most often leaf-rollers, leaf- miners and galling insects in the orders Lepidoptera, Diptera, Hymenoptera and Coleoptera (Schauff et al. 1997, Boucek 1988). Species of Cirrospilus are predomi- nantly larval ectoparasitoids, but also have been documented both as hyperparasitoids on primary ichneumonoid parasitoids and as primary parasitoids on several life stages of the primary host (Boucek and Askew 1968). Over 300 nominal species of Cir- rospilus occur worldwide, primarily in the Holarctic (Noyes 1998). Species of Cirrospilus possess a two- segmented funicle, complete notauli that extend to the transscutal articulation (TSS) (Fig. 2), and the postmarginal vein is shorter than or equal in length to the stig- mal vein (Schauff et al. 1997). However, the location of the axilla can vary within Cirrospilus from being almost entirely posteriad to slightly anteriad of the TSS (LaSalle, personal communication). Fur- ther, as a member of Cirrospilini (Gauthier et al. 2000), Cirrospilus are characterized by a transverse sulcus on the face (Fig. 3) Hymenoptera, Eulophidae, Cirrospilus, taxonomy, Chalcidoidea and the posterior separation of the pro- pleura. A closely related genus, Zagram- mosoma Ashmead, has been separated from Cirrospilus on the basis of characters that intergrade between the two genera (Gordh 1978, LaSalle 1989, Gates 2000). For example, the placement of the axilla relative to the TSS cited by Yefremova (1996) appears useful in differentiating the two genera in some instances, but by no means all. For a discussion of generic character intergradation, see Gates (2000) and a summary in Table 1. Acronyms used are: USNM = National Museum of Natural History, Smithsonian Institution, Washington, DC; DLWC = Da- vid Wagner collection, University of Con- necticut, Storrs, CT. Cirrospilus infuscatus Gates and Schauff, new species (Figs. 1-5) Type material.—Holotype, 2. “AZ: Co- chise Co., Huachuca Mtns., Hunter Cyn 5100’, 14-IV-1986 Wagner; DLW Lot: 86D70, em. 26-IV/10-V1986, Host: Tis- cheria bifurcata” (USNM). Paratypes, 3 ° 1 d, same data as holotype (1 2 1 ¢ point- VOLUME 105, NUMBER 2 301 Table 1. Character distribution in Cirrospilus and Zagrammosoma. Character Cirrospilus Zagrammosoma Vertex vaulted Rarely Typical Propodeum with strong median carina Often Rarely Notaulus orientation Extends straight to TSS Curves to intercept axilla Axilla shape Triangular Elongate Axilla placement Mostly posterior to TSS Mostly anterior to TSS mounted, | 2 slide/SEM stub mounted convex in lateral view. Mesopleuron gla- (USNM): 1 @ card mounted (DLWC). Etymology.—This species is named for its infuscate wing pattern. Diagnosis.—This species has the follow- ing unique features within Cirrospilus: pet- iolate gaster in both sexes with the petiole 1.4—1.7* as long as broad; an infuscate fore wing; strong, wide submedian lines on the scutellum; and the brachypterous male with the apex of the fore wing not extend- ing beyond the basal % of the gaster. Description.—Female: Length 1.6—1.7 mm. Head and body deep golden, gaster dark brown laterally and with transverse bands dorsally, antenna dark brown. Legs golden, tarsi paler except apical tarsomeres brown. Wings hyaline except forewing in- fuscate in medial half (Fig. 5), venation brown. Head: Finely reticulate with even cov- erage of fine brown setae. Roughly quad- rate in frontal view, 1.0—1.1* as high as broad (Fig. 3). Malar space 0.4—0.5* eye height. Eye glabrous (minute setae appar- ent at high magnification) (Fig. 3). Scrobal depression shallow, unmargined. Scape 4.6-5.5* as long as broad, two anelli transverse and short; pedicel 2.3—2.8% as long as broad; Fl 1.3—1.6* as long as broad; F2 1.0—1.8 as long as broad; clava 2.0—2.4* as long as broad, 3-segmented, tapering apically (Fig. 4). Mesosoma: Transversely reticulate ante- riorly, becoming reticulate medially to gla- brate posteriorly (Fig. 2). Midlobe of me- soscutum |.1—1.2* as long as broad. Scu- tellum 1.0—1.1 as long as broad, with dis- tinct submedial grooves most closely approaching each other anteriorly (Fig. 2); brous to faintly reticulate. Propodeum gla- brous with fine median carina; callus with row of at least 8 setae. Fore wing 2.5—3.1 x as long as broad, setose in apical half (Fig. 5); marginal vein 8.0—-10.0* as long as postmarginal vein; stigmal vein 3.3—3.5 as long as postmarginal vein. Metasoma: Petiole 1.4—1.7X as long as broad, finely rugulose (Fig. 1). Gaster |.1— 1.2 as long as mesosoma. Tips of Ovipos- itor sheaths visible in dorsal view (Fig. 2). Male: Length 1.2 mm. Identical to fe- male except as follows: malar space 0.6 eye height; scape 4.2 as long as broad; pedicel 1.8 as long as broad; Fl 2.3 as long as broad; F2 1.2 as long as broad; clava 2.6 as long as broad; midlobe of mesoscutum 1.3% as long as broad; fore- wing brachypterous, 3.8% as long as broad; marginal vein 20 as long as postmarginal vein; stigmal vein 6.0* as long as post- marginal vein; petiole 1.4 as long as broad; gaster 0.9 as long as broad. Variation.—The type series is essentially invariant. Biology.—These specimens were reared from Tischeria bifurcata Braun (Lepidop- tera: Tischeriidae) mining the leaves of Ce- anothus fendleri A. Gray (Rhamnaceae) in the Huachuca Mountains of southeastern Arizona. A small proportion of total leaf mines collected were formed by a species of Recurvaria (Lepidoptera: Gelechiidae), but it is unlikely that these parasitoids emerged from those mines (Wagner, per- sonal communication). Like other species of Cirrospilus (Gates 2000), C. infuscatus may be a gregarious parasitoid, as all spec- imens emerged on the same day. Unfortu- 302 PROCEEDINGS OF THE ENTOMOLOGICAI transverse sulcus > Figs. 1-5. Cirrospilus infuscatus, female scutal articulation nately, it is unknown if all specimens emerged from a single host insect. Tis- cheria bifurcata forms an irregular track on a single leaf with the mesophyll not completely consumed between epidermal layers. The frass is forced out of the lower thrust through the lower epidermis (Wagner, field epidermis and the pupa is also notes ). SOCIETY OF WASHINGTON 1, Dorsal petiole. 2, Dorsal mesosoma and gaster; TSS trans- 3, Frontal head. 4, Antenna. 5, Fore wing ACKNOWLEDGMENTS We are grateful to David Wagner (Uni- versity of Connecticut) for allowing access to his reared leaf miner materials where this species was discovered. Thanks also to John LaSalle (CSIRO, Canberra, Australia), John Heraty (University of California, Riv- erside, CA), Norm Woodley and Dave Smith (both USDA, Systematic Entomolo- VOLUME 105, NUMBER 2 gy Laboratory, Washington, DC) for critical comments on earlier drafts of this manu- script. LITERATURE CITED Bouéek, Z. 1988. Australasian Chalcidoidea (Hyme- noptera): A Biosystematic Revision of Genera of Fourteen Families, with Reclassification of Spe- cies. CABI, Wallingford, United Kingdom. 832 Pp- Boucek, Z. and R. Askew. 1968. Palaearctic Eulophi- dae (excl. Tetrastichinae) (Hym. Chalcidoidea). Index of Entomophagous Insects. Le Francois, Paris. 260 pp. Gates, M. 2000. A new species of Cirrospilus West- wood (Hymenoptera: Eulophidae) from the south- western United States and Mexico. Proceedings of the Entomological Society of Washington 102(1): 58-61. Gauthier, N., J. LaSalle, D. Quicke, and H. Godfray. 2000. Phylogeny of Eulophidae (Hymenoptera: Chalcidoidea), with a reclassification of Eulophi- 303 nae and the recognition that Elasmidae are derived eulophids. Systematic Entomology 25: 521-539. Gordh, G. 1978. Taxonomic notes on Zagrammosoma, a key to the Nearctic species and descriptions of new species from California (Hymenoptera: Eu- lophidae). Proceedings of the Entomological So- ciety of Washington 80: 344-359. LaSalle, J. 1989. Notes on the genus Zagrammosoma (Hymenoptera: Eulophidae) with description of a new species. Proceedings of the Entomological Society of Washington 91: 230-236. Noyes, J. 1998. Catalogue of the Chalcidoidea of the World. CD-ROM Expert Center for Taxonomic Information, Amsterdam, The Netherlands. Schautf, M., J. LaSalle, and L. Coote. 1997, Chapter 10. Eulophidae, pp. 327-429. In Gibson, G., J. Huber and J. Woolley, eds. Annotated Keys to the Genera of Nearctic Chalcidoidea (Hymenoptera). NRC Research Press, Ottawa, Ontario, Canada. Yefremova, Z. 1996. Notes on some Palaearctic and Afrotropical species of the genus Zagrammosoma (Hymenoptera, Eulophidae). Zoologicheskiy Zhurnal 74(10): 46—54. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 304-311 REDESCRIPTION OF THE LARVA OF GOMPHUS MILITARIS HAGEN (ODONATA: GOMPHIDAE), WITH DISTRIBUTIONAL AND LIFE HISTORY NOTES Brett H. P. LANDWER AND ROBERT W. SITES Enns Entomology Museum, Department of Entomology, and Center for Agroforestry, University of Missouri, Columbia, MO 65211, U.S.A. (e-mail: bhl1065@mizzou.edu and SitesR @ missouri.edu) Abstract.—tThe final larval instar of Gomphus militaris Hagen is described, diagnosed, and figured from exuviae and larval specimens from Missouri, Texas, and Kansas. A previous description was erroneously attributed to G. militaris, but actually pertained to Arigomphus lentulus. Thus, the use of many previously published characteristics to iden- tify this species will result in misidentification. Larvae of G. militaris were common and widespread in ponds in the prairie region of Missouri. We report data on life history based on larval and adult collections. Key Words: Gomphus militaris Hagen is primarily a species of the southern Great Plains of North America and has been reported from Iowa and North Dakota, south to New Mex- ico and Nuevo Leon, Mexico (Dunkle 2000, Needham et al. 2000). It was the “most frequent species of the genus” in Oklahoma (Bick and Bick 1957), the most widely distributed and commonly encoun- tered species of Gomphidae in Texas (Ab- bott 2001), and common throughout Kansas (Huggins et al. 1976, Beckemeyer and Hug- gins 1997). Recently, it was reported to be common in western Missouri (L. Trial and J. F Belshe, in litt.). Due in part to its limited range, infor- mation regarding the ecology and life his- tory of G. militaris 1s scant. It is a species of ponds or sluggish streams (Bird 1934; Beckemeyer and Huggins 1997; Dunkle 2000; L. Trial and J. E Belshe, in litt.) where the larvae burrow in the sediment and detritus along the margins (personal ob- servation). The adult flight season is 19 Odonata, Gomphidae, Gomphus, Arigomphus, larva, Missouri March to 15 August in Texas (Abbott 2001), and June and July in Kansas (Be- ckemeyer and Huggins 1997). In Texas, most specimens collected between 18 May and | June were reported to be teneral (Bick and Bick 1957). In Oklahoma, the peak of larval emergence occurred during mid-May (Bird 1934). Specimens reared in Kansas by D. G. Huggins emerged during early June (unpublished data). We conducted a survey of the larval Odonata of ponds in the prairie (sensu Stey- ermark 1963, Pfleiger 1989, Bailey 1998) or plains (sensu Branson 1944, Thornbury 1965, Thom and Wilson 1980, Omernick 1987) regions of Missouri in 1998 and 1999. When determining specimens col- lected during this survey, we repeatedly en- countered a single morphospecies of Gom- phidae that could not be determined using existing keys that include the Midwestern fauna (i.e., Needham and Westfall 1955, Needham et al. 2000). These larvae super- ficially resembled those of the genus Ari- VOLUME 105, NUMBER 2 gomphus Needham in general form and most often keyed to that genus. However, these larvae lacked the large, jagged teeth on the labial palps and the mid-dorsal ridge characteristic of larvae of Arigomphus. Moreover, adults of G. militaris frequently had been collected at ponds in the region, but no larvae had been encountered that could be attributed to this species using the published characters. However, our larvae possessed several characteristics which were contradictory to those attributed to G. militaris in the existing keys (1.e., Needham and Westfall 1955, Young and Bayer 1979, Needham et al. 2000) and in the larval de- scription given by Bird (1934). Specimens were sent to K. J. Tennessen who determined that they were indeed G. militaris and informed us that larvae of this species may not agree with published char- acters (personal communication). We also reared a male specimen and determined it as G. militaris. We present a redescription of the larva of G. militaris, with distributional data and notes on its life history in Missouri. MATERIALS AND METHODS We measured 10 specimens: exuviae of one partially emerged male specimen, ex- uviae of one unreared final instar, and eight final instars. Three of the larvae were from Texas, one from Kansas, and the remaining Six specimens from Missouri. All measurements were performed in dorsal view and represent maximum values, unless otherwise stated. Due to non-uni- form telescoping of the abdomen, exact measures of total length and length of ab- domen are impossible to obtain (see Calvert 1934, Huggins and Harp 1985). Therefore, these measurements were performed to the nearest mm using calipers. All other mea- surements were performed using an ocular micrometer. The head was measured from the anterior margin of the clypeus to the posterior margin of the postocciput, and the abdomen to the tip of the caudal append- ages. The outer surface was considered the dorsal aspect of the leg. Wingpads were 305 measured from the dorsal juncture with the thorax to the tip. Mid-dorsal and mid-ven- tral abdominal segment lengths were mea- sured from anterior to posterior margins of a sclerite, excluding dorsal hook, if present. Lateral margins of abdominal segments were measured from the anterolateral cor- ner to the tip of the lateral spine. Lateral spines were measured from the posterior margin of the tergite immediately adjacent to the spine, to the tip of the spine. Basal width of abdominal segments were maxi- mum values, measured just beyond the slight basal constriction. To compensate for lateral distortion of the abdomen in exuviae and preserved larvae (see Tennessen and Louten 1984, Huggins and Harp 1985), the abdomen was depressed for abdominal seg- ment width measurements (except segment X) so that the ventral surface was flat in cross section. Caudal appendages were measured from the posterodorsal margin of segment X to the tip of the appendage. SYSTEMATICS Gomphus militaris Hagen Description.—Total length 28-30 mm (mode = 30 mm). Overall body shape elon- gate fusiform, slightly depressed (Fig. 1). Body color light brown with darker brown markings. Dorsal surface of body finely granulose with glabrous patches on muscle scars. Minute, scalelike setae emanating from granules. Ventral surface generally not granulose, but with very fine setae. Larvae collected in situ with fine dorsal setation generally obscured by layer of fine silt and detritus. Head: Width 5.25—5.67 mm (¥ = 5.50), length 3.08—3.33 mm (y = 3.21). Postero- lateral margins abruptly rounded. Clypeus extending anteriorly to apex of second an- tennal segment, labrum to about middle of third segment. Compound eyes with mesal margins converging posteriorly. Antenna with segments III and IV darker than light brown general body color, segment III ex- panding slightly to apex and bowed up- 306 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON “or syst ‘on sn KiNUDRD Figs. 1—4. Gomphus militaris larva. 1, Dorsal view of larva. 2, Ventral view of labium. 3, Blade of labial palp (detail of inset of Fig. 2). 4, Dorsal view of abdominal segments VII-X and terminal appendages of female. VOLUME 105, NUMBER 2 ward. Ratio of antennal segment lengths 5: 3:13:1. Prementum, in ventral aspect, con- stricted in proximal ca. %, lateral margins subparallel in apical % (Fig. 2). Mid-ventral length of prementum, excluding hinge, 3.20—3.32 mm (¥ = 3.25), width 2.40—2.64 mm (y = 2.51). Anterior margin of pre- mentum between bases of labial palps (lig- ula) evenly convex with dense fringe of long, stout, scalelike setae, and without dis- tinct median tooth (Fig. 2). Blade (first seg- ment) of labial palp rounded apically, with 7-12 (mode = 9) very short, nearly obso- lete, squarely truncate teeth along the slightly arcuate mesal margin, distal-most tooth triangular but not longer than remain- ing teeth (Fig. 3). Long, conspicuous hairs on antennal segments, distal margin of la- brum, and lateral margin of head beneath the compound eye from base of antennae to postocciput. Glabrous areas on dorsum of head comprising proximal portion of clyp- eus; V-shaped spot in center of frons; oval area lateral to V-shaped spot and adjacent to mesal margin of compound eye: spots immediately anterior to postocciput at mid- line and ca. 7 length from mid-line to pos- terolateral corner; thin, elongate spot above posterolateral corner. Thorax: Areas of darker pigmentation variously developed, comprising margins of mesothoracic spiracle; dorsal portion of meso- and metapleural sutures; thin, longi- tudinal line dorsally between mesepisterna; immediately lateral to anal angle of meta- thoracic wingpads. Prothorax with ring shaped glabrous area lateral to mid-line, confluent anterolaterally with glabrous area on pleuron above base of coxa. Glabrous areas also present on and immediately ad- jacent to meso- and metapleural sutures. Fringe of long, conspicuous hairs along lat- eral margins of thorax. Pigmentation of legs slightly darker at apex of tibia. Legs hairy, with distinct fringes on dorsal and ventral surfaces. Pro- and mesotibial borrowing hooks well developed. Length of hind fe- mur 5.42-5.83 mm (¥ = 5.53 mm), apex attaining ca. mid-length of abdominal seg- 307 ment IV. Glabrous areas of legs comprising triangular areas on both anterior and pos- terior surfaces of coxae; triangular area on the ventral surface of trochanters; longitu- dinal lines extending from base to near apex of femora on anterior, posterior, and dorsal surfaces; similar, less conspicuous lines on tibiae. Length of metathoracic wingpad 6.42—6.83 mm (y = 6.61 mm), attaining ca. mid-length of abdominal segment IV. Me- sothoracic wingpad slightly overhanging anterior margin of abdominal segment IV. Wingpads densely granulose and pubescent on margins, less so on major veins of meta- thoracic wingpads, remainder glabrous. Dark coloration at nodi and tips of meso- and metathoracic wingpads, and proximal anal region of metathoracic wingpad. Abdomen: Slender and gradually taper- ing to tip (Fig. 1), moderately depressed. Lateral margins with fringe of long setae. Posterior margin of tergites with sparse fringe of shorter setae, densest near mid- line. Length of abdomen 18—21 mm (mode = 21 mm), maximum width 7.0—7.4 mm (y = 7.2 mm). Integument glabrous at muscle scars, external surfaces of terminal append- ages, and entirety of segment X. Segments I-VHI dorsally with paired dark spots near mid-length of each segment just lateral to mid-line (Fig. 4). Mesal margin of each muscle scar also with dark spot. Segment IX with dark spot at posteromedial corner of muscle scar (Fig. 4). Spots absent from segment X, and fading on segments anterior to IV. Lateral margins diverging moderately to segment IV, converging gradually be- yond segment V to parallel-sided segment X (Fig. 1). Dorsal mid-line without ridge or impressed line. Segments I-VII with low, posteromedial protuberance occupying Ca. % length of segment. Posteromedial protu- berance of segment VIII very low. Female with rudimentary lobes of vulvar lamina projecting slightly from posterior margin of abdominal sternite VIII. Dorsum of seg- ment IX, in cross section, nearly tectate an- teriorly, circular posteriorly. Mid-dorsal length of segment IX, excluding hook, 83— 308 99% (¥ = 92%) basal width, and 84—94% (vy = 87%) mid-ventral length. Segment 10 cylindrical. Basal width of segment X 83— 97% (y¥ = 93%) mid-dorsal length. Lateral spines present on abdominal segments VII—IX (rarely a minute spine on VI). Lat- eral spines of VII and VIII small, compris- ing 8-14% (¥ = 10%) and 11-20% (¥y = 13%), respectively, of lateral margin of seg- ment. Lateral spine of segment IX long, slender, and somewhat appressed to side of segment X, 36-50% (¥ = 43%) as long as mid-dorsal length of segment X, usually ex- tending slightly short of mid-length of seg- ment X. Length of lateral spine of segment IX 0.56—0.72 mm (y = 0.64 mm). Mid-dor- sal posterior hook present on segment IX, usually present on segment VIII, and oc- casionally present on segment VII. Hook on segment VIII and VII, if present, minute and formed by gradual recurvature of pos- terior margin of tergite on either side im- mediately lateral to mid-line, to form an acuminate process. Hook of segment IX flat, slightly larger than that of segment VIII, more abruptly protruding from pos- terior margin of tergite, and slightly over- hanging anterior margin of segment X. Length of epiproct 80-88% (y = 84%) mid- dorsal length of abdominal segment X, and 93-100% (¥ = 97%) length of paraprocts. Male epiproct, in dorsal view, abruptly nar- rowed immediately beyond mid-length, forming a pair of rounded, lateral humps. Female epiproct tapering more regularly to tip (Fig. 4). Length of cercus 83-96% (y = 92%) that of epiproct. Diagnosis.—The larva of Gomphus mil- itaris can be distinguished from North American congeners by the following com- bination of characters: length of abdominal segment X subequal to or decidedly greater than basal width; mid-dorsal length of ab- dominal segment IX subequal to or less than basal width; lateral spines of abdomi- nal segment VI absent or minute; distal margin of labial palp rounded and without end hook; teeth of labial palp truncate and nearly obsolete. Gomphus militaris can be PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON distinguished from species of Arigomphus by the absence of a mid-dorsal ridge on the anterior portion of the middle abdominal tergites, and the teeth of the labial palp small, squarely truncate, and much shorter than basal width. Examination of earlier in- star specimens indicates that these charac- teristics are quite stable in specimens as small as ca. 15 mm in total length. Abdom- inal segment X becomes slightly more elon- gate and the lateral spines of IX extend a proportionately shorter distance in progres- sively earlier instars. Material examined.—TEXAS: Kimble Co.: N. Liano R. 1.7 mi. W. on Rt. 377, V-24-1987, RWS, 1 larva; Mason Co.: James R. near London, IV-11-1992, RWS, 2 larvae; KANSAS: Elk Co.: Big Caney R.—1 mi. W. Grenoda at U.S. 160 hwy. br., V-12-1982, B. Coler & D. G. Huggins, 1 larva; MISSOURI: Audrain Co.: Robert M. White II Conservation Area, UTM (Uni- versal Transverse Mercator) zone 15 S59TOS0E, 4353000N, IV-24-1999, BHPL & N. Whiteman, | larva: Boone Co.: Ashland Lake, UTM zone 15 568980E 4290420N, V-6-2001, BHPL, | male (partially emerged V-26 2001); Caldwell Co.: Bonanza Con- servation Area, UTM zone 15 418010E 4385720N, III-10-1999, BHPL & N. Whiteman, | larva; Monroe Co.: Otter Cr., UTM zone 15 569650E 4384400N, VI-21- 2001, BHPL, exuviae (1 specimen); Platte Co.: Guy B. Park Conservation Area, UTM zone 15 345150E 4365040N, IIl-21-1999, BHPL & N. Whiteman, Co.: Douglas Branch Conservation Area, UTM zone 15 379030E 4198180N, If-22- 1999, BHPL & N. Whiteman, | larva. Additional Missouri material exam- ined.—Atchison Co.: Tarkio Prairie Con- servation Area, BHPL & N. Whiteman, UTM zone 15 313200E 4485890N, [X-12— 1998: same data, UTM zone 15 312230E 4486100N, BHPL & N. Whiteman; Au- drain Co.: Robert M. White I Conserva- tion Area, UTM zone 15 597100E 4353 150N, [X-9-1998, BHPL & N. White- man; Barton Co.: Shawnee Trail Conser- 1 larva; Vernon VOLUME 105, NUMBER 2 vation Area, UTM zone 15 359735E 4143150N, LX-26-1998; Dorris Creek Prai- rie Conservation Area, UTM zone 15 390765E 4137060N, I[X-27-1998, BHPL & N. Whiteman; same data, VI-9-1999, BHPL & N. Whiteman; Clear Creek Conservation Area, UTM zone 15 390900E 4166300N, VI-9-1999, BHPL & N. Whiteman; Carroll Co.: Bunch Hollow Conservation Area, UTM zone 15 449350E 3380150N, [X-13- 1998, BHPL & N. Whiteman: Harrison Co.: Wayne Helton Memorial Conservation Area, UTM zone 15 432000E 4454000N, VI-23-1999, BHPL & N. Whiteman; Henry Co.: Connor O. Fewel Conservation Area, UTM zone 15 439600E 4259650N, LX-20- 1998, BHPL & N. Whiteman; Holt Co.: Riverbreaks Conservation Area, UTM zone 15 321210E 4420700N, VI-15-1999, BHPL & N. Whiteman; Knox Co.: Henry Sever Lake Conservation Area, UTM zone 15 587200E 4429840N, I[X-6-1998, BHPL & N. Whiteman; Lafayette Co.: Maple Leaf Lake Conservation Area, UTM zone 15 431410E 4316450N, [X-18-1998, BHPL & N. Whiteman; Livingston Co.: Poosey Con- servation Area, UTM zone 15 442000E 4420220N, IX-12-1998, BHPL & N. Whiteman; St. Clair Co.: Taberville Prairie Conservation Area, UTM zone 15 414800E 4211890N, IX-26-1998, BHPL & N. Whiteman; Vernon Co.: Little Osage Prai- rie Natural Area, UTM zone 15 381800E 4180440N, IX-26-1998, BHPL & N. Whiteman: Worth Co.: Emmett and Leah Seat Memorial Conservation Area, UTM zone 15 396450E 4471620N, I[X-12-1998, BHPL & N. Whiteman. DISCUSSION The results demonstrate that previously published characters will not correctly dis- tinguish larvae of Gomphus militaris from those of congeners. In their key to species of the subgenus Gomphus, Needham and Westfall (1955) characterized G. militaris as the only species of the subgenus not pos- sessing lateral spines on abdominal seg- ments VII and VIII. As such, the larva of 309 G. militaris will key to G. spicatus Hagen with Needham and Westfall (1955). This shortcoming is corrected only partially by Needham et al. (2000), who characterized G. militaris as possessing lateral spines on abdominal segments VIII and IX, or IX only. In that key, the larva can not be keyed to any species. As lateral spines are present on abdominal segments VIL-IX. The results of this analysis also reveal the inadequacy of existing keys to reliably dis- tinguish G. militaris from species of Ari- gomphus. The abdomen of species of Ari- gomphus has been characterized as taper- ing, whereas that of species of Gomphus as ending more abruptly (Needham and West- fall 1955, Young and Bayer 1979, Needham et al. 2000). However, the abdomen of G. militaris 1s tapering, as in species of Ari- gomphus. Also, abdominal segment X of species of Arigomphus has been character- ized as ‘‘distinctly’> longer than wide (Young and Bayer 1979, Needham et al. 2000) and segment IX as wider than long (Needham et al. 2000), whereas abdominal segment X in species of Gomphus has been characterized as not longer than wide (Young and Bayer 1979), or if longer, then with segment IX longer than wide (Need- ham et al. 2000). As shown here, segment X of G. militaris is usually longer than wide, but abdominal segment IX is not. Fi- nally, species of Gomphus with the abdo- men acuminate and abdominal segment X longer than wide have been characterized as possessing large end hooks on the labial palps (Needham et al. 2000); however, G. militaris possesses no such end hooks. The inability of published characters to diagnose the larva of G. militaris is not due to intraspecific variation. Larvae from throughout the range of this species were consistent in the manifestation of key char- acteristics. In addition to the specimens measured in the description, six final instar exuviae, reared by D. G. Huggins, and nu- merous larvae from Kansas (Kansas State Biological Survey collection, housed at the shown _ here, 310 University of Kansas Natural History Mu- seum and the Snow Museum of Entomol- ogy) were examined but not measured, and diagnostic characters were consistent throughout. Further, although Young and Bayer (1979) accurately figured the labium and abdomen of G. militaris, the characters provided in their key were inadequate to di- agnose the species. Much of the confusion can be traced to the published larval description by Bird (1934), upon which existing keys seem, at least partially, to be based. Bird’s descrip- tion was based on the exuviae from the emergence of a single female from a pond in Oklahoma on Y-10-1932. The teneral imago escaped ‘one hour and seventeen minutes after the nymph had left the wa- ter.’ The association was based upon pho- tographs and notes of the teneral imago. The senior author has examined the exuvial specimen described by Bird and determined it to be that of Arigomphus lentulus (Need- ham). Determination of female specimens of Gomphidae is difficult, even when working with mature material in the laboratory, and field identification of teneral specimens is highly suspect. Additionally, adult speci- mens of Arigomphus are less darkly pig- mented than those of related genera, and G. militaris displays reduced dark markings relative to other species of the genus Gom- phus. Further, the nymph of A. /entulus was unknown at the time of Bird’s description. Thus, Bird’s field identification was a rea- sonable error. The description by Bird was the first for- mal description of the larva of A. lentulus, and should be cited as such. There can be little doubt that the exuviae examined by BHPL are those described by Bird. The vial bears his determination label as Gomphus militaris, and the female specimen was col- lected by Bird in McClain Co., Oklahoma on V-10-1932, which matches the data giv- en in his description. Only one specimen bearing this information was found in Bird’s collection, housed at the Sam Noble PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Oklahoma Museum of Natural History, University of Oklahoma (Ken Hobson, per- sonal communication). No information re- garding an associated imago is recorded on the label. The specimen exactly matches Bird’s figures and agrees with the descrip- tion, except on two points: Bird stated that lateral spines are present only on abdominal segment IX, and that no dorsal “teeth” are present on any abdominal segment. How- ever, the figures which accompany Bird’s description clearly depict a small dorsal hook on abdominal segment LX, and indi- cate lateral spines on at least segments VIII and IX. The specimen does, in fact, bear a small dorsal hook on segment IX, and small lateral spines on segments VII and VIII in addition to the large lateral spines of seg- ment IX. Additionally, it should be noted that other larval specimens of A. lentulus often bear a small dorsal hook on segment VIII. Interestingly, as evidenced by Bird’s description, the length of abdominal seg- ment X of specimens of A. lentulus often is no greater than the basal width. Gomphus militaris was the most fre- quently encountered species of the family Gomphidae in ponds in the prairie region of Missouri. Final instars were collected only during the spring, but much smaller instars were also collected at this time. Samples from early summer contained only intermediate instars, and samples from late summer contained both early and late in- stars. A final instar collected on 6 May died while emerging on 26 May, and exuviae were collected on 21 June. This emergence, coupled with adult collection data, estab- lishes the flight season as 26 May to 5 Au- gust in Missouri. The instar distribution and flight season data indicate that G. militaris emerges somewhat synchronously during the spring and at least some individuals may require more than one year to complete development in Missouri. The larval distribution records indicate that G. militaris is more widespread in Mis- souri than previously shown. The collection records from Shelby, Audrain, and Knox VOLUME 105, NUMBER 2 counties extend the known eastern limit of the species range ca. 150 km beyond that given in the most recently published, de- tailed account (Dunkle 2000). Also, the col- lection of exuviae from Shelby County demonstrates adult recruitment at this limit, and the Knox County record places the known limit within ca. 40 km of the Mis- sissippi River. ACKNOWLEDGMENTS We thank Ken J. Tennessen for determi- nation of specimens and a critical review of the manuscript. We are grateful to Jordan Yochim, University of Kansas Natural His- tory Museum, and Dr. James S. Ashe, Snow Entomological Museum, University of Kansas, for the opportunity to examine ma- terial housed in their collections. Dr. Ken Hobson, Sam Noble Oklahoma Museum of Natural History, University of Oklahoma, was instrumental in obtaining Dr. Bird’s specimen. Linden Trial, Missouri Depart- ment of Conservation, provided adult ma- terial. Funding for RWS was provided in part by project #PSSLO232 of the Missouri Agricultural Experiment Station. LITERATURE CITED Abbott, J. C. 2001. Distribution of dragonflies and damselflies (Odonata) in Texas. Transactions of the American Entomological Society 127: 189— 228. Bailey, R. G. 1998. Ecoregions—The Ecosystem Ge- ography of the Oceans and Continents. Springer- Verlag, New York, 176 pp. Beckemeyer, R. J. and D. G. Huggins. 1997. Checklist of Kansas dragonflies. The Kansas School Natu- ralist 43: 1-16. Bick, G. H. and J. C. Bick. 1957. The Odonata of Oklahoma. The Southwestern Naturalist 2: 1-18. Bird, R. D. 1934. The emergence and nymph of Gom- phus militaris (Odonata, Gomphinae ). Entomolog- ical News 45: 44—46. 311 Branson, E. B. 1944. The geology of Missouri. The University of Missouri Studies 19: 1-535. Calvert, P. P. 1934. The rates of growth, larval devel- opment and seasonal distribution of dragonflies of the genus Anax (Odonata: Aeshnidae). Proceed- ings of the American Philosophical Society 73: 1— 70. Dunkle, S. W. 2000. Dragonflies Through Binoculars: A Field Guide to Dragonflies of North America. Oxford University Press, Oxford, 266 pp. Huggins, D. G. and G. L. Harp. 1985. The nymph of Gomphus (Gomphurus) Westfall (Odonata: Gomphidae). Journal of the Kansas En- tomological Society 58: 656-661. Huggins, D. G., P. M. Liechti, and D. W. Roubik. 1976. Species accounts for certain aquatic macroinver- tebrates from Kansas (Odonata, Hemiptera, Co- leoptera, and Sphaeriidae), pp. 13-77. In New Re- cords of the Fauna and Flora of Kansas for 1975, Technical Publications of the State Biological Sur- vey of Kansas, no. 1. Needham, J. G. and M. J. Westfall, Jr. 1955. A Manual of the Dragonflies of North America (Anisoptera). University of California Press, Berkeley, 615 pp. Needham, J. G., M. J. Westfall, Jr, and M. L. May. 2000. Dragonflies of North America. Scientific Publishers, Gainesville, 939 pp. Omernik, J. M. 1987. Ecoregions of the conterminous United States. Annals of the Association of Amer- ican Geographers 77: 118-125. Pflieger, W. L. 1989. Aquatic Community Classifica- tion System for Missouri. Aquatic series no. 19. Missouri Department of Conservation, Jefferson City, Missouri, 69 pp. Steyermark, J. A. 1963. Flora of Missouri. lowa State University Press, Ames, 1,725 pp. Tennessen, K. J. and J. A. Louten. 1984. The true nymph of Gomphus (Gomphurus) crassus Hagen (Odonata: Gomphidae), with notes on adults. Pro- ceedings of the Entomological Society of Wash- ington 86: 223-227. Thom, R. H. and J. H. Wilson. 1980. The natural di- visions of Missouri. Transactions of the Missouri Academy of Sciences 14: 9-23. Thornbury, W. D. 1965. Regional Geomorphology of the United States. John Wiley and Sons, New York, 609 pp. Young, W. C. and C. W. Bayer. 1979. The dragonfly nymphs (Odonata: Anisoptera) of the Guadalupe River Basin, Texas. The Texas Journal of Science 31: 85-97. ozarkensis PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 312-319 ANNOTATED LIST OF THE BARK AND AMBROSIA BEETLES (COLEOPTERA: SCOLYTIDAE) OF DELAWARE, WITH NEW DISTRIBUTIONAL RECORDS ROBERT J. RABAGLIA AND MICHAEL A. VALENTI (RJR) Forest Pest Management, Maryland Department of Agriculture, 50 Harry S. Truman Parkway, Annapolis, MD 21401, U.S.A. (e-mail: rabaglrj@mda.state.md.us); (MAV) Delaware Department of Agriculture, 2320 South DuPont Highway, Dover, DE 19901, ULS:A. Abstract.—A total of 64 species of Scolytidae are reported from Delaware. Fifty-one species are reported for the first time: Hylastes porculus Erichson, H. salebrosus Eichhoff, H. tenuis Eichhoff, Hylurgops rugipennis pinifex (Fitch), Hylesinus aculeatus Say, Den- droctonus frontalis (Zimmermann), D. valens LeConte, Cnesinus strigicollis LeConte, Phloeotribus liminaris (Harris), Phloeosinus dentatus (Say), Carphoborus bifurcus E1- chhoff, Hylocurus rudis (LeConte), H. flaglerensis Blackman, Micracisella opaciicollis (LeConte), Thysanoes fimbricornis LeConte, Ips grandicollis (Eichhoff), /. pini (Say), Orthotomicus caelatus (Eichhoftf), Pityogenes hopkinsi Swaine, Dryoeocetes autographus (Ratzeburg), D. granicollis (LeConte), Lymantor decipiens (LeConte), Xyloterinus politus Say, Ambrosiodmus obliquus (LeConte), A. rubricollis (Eichhoft), A. tachygraphus (Zim- mermann), Euwallacea validus (Eichhoff), Xyleborus atratus Eichhoff, X. celsus Eichhoff, X. ferrugineus (Fabricius), X. pelliculosus Eichhoff, X. pubescens Zimmermann, X. sayi (Hopkins), Xylosandrus crassiusculus (Motschulsky), X. germanus (Blandford), Hypoth- enemus crudiae (Panzer), H. dissimilis (Zimmermann), H. eruditus Westwood, Cono- phthorus coniperda (Schwarz), Corthylus columbianus Hopkins, Gnathotrichus materiar- ius (Fitch), Monarthrum fasciatum (Say), M. mali (Fitch), Pityophthorus cariniceps LeConte, P. confusus Blandford, P. liquidambarus Blackman, P. opaculus LeConte, P. puberulus (LeConte), P. scriptor Blackman, Pseudopityophthorus minutissimus (Zimmer- mann), P. pubescens Blackman. Fourteen of the species are not native to North America. Key Words: | Scolytidae, distribution, Delaware Though its geographical area is compar- atively small, Delaware exhibits an extraor- dinary diversity of tree species and forest associations. Due to its location in a north/ south floral transition zone, many of the forests that typify northern, central, and United States can be found in Delaware (Tatnall 1946, Phillips 1978). Of the approximately 600 species of trees native to the entire United States, about 115 are native to Del- southern regions of the eastern aware (Taber 1939). About one third of the land in Delaware is covered with mostly second and third growth forests represent- ing a fragmented patchwork of the original contiguous forest. The diversity of tree species and forest associations in Delaware is not adequately reflected by the previously documented members of the Scolytidae in the state (Wood and Bright 1992). Such a disparity prompted a preliminary survey of this im- VOLUME 105, NUMBER 2 portant forest insect group in 1997 followed by a more intensive state-wide survey in 1998. In 1997, twelve Lindgren funnel traps baited with 95% ethanol and natural tur- pentine were placed in two forested areas and monitored from April through Septem- ber. In 1998, thirty-six funnel traps were placed at 19 forested sites throughout the state and monitored from March through October. Traps were baited with a variety of lures including ethanol, turpentine, cut tree branches, and various commercially avail- able (PheroTech Inc, Delta, BC, Canada) bark beetle pheromones such as frontalin (Dendroctonus frontalis (Zimmermann) pheromone) and sulcatol (a pheromone for a species of Gnathotrichus). Also in 1998, specimens at the University of Delaware in Newark were examined, as well as speci- mens collected by the Delaware Depart- ment of Agriculture during several years of agricultural pest surveys. Two catalogs of Scolytidae (Wood and Bright 1992, Bright and Skidmore 1997) list only 11 scolytids from Delaware. We report an additional 51 species. Two addi- tional species, Xyleborus californicus Wood and Dryoxylon onoharaensum (Murayama), were reported from Delaware for the first time in recent publications (Vandenberg et al. 2000, Bright and Rabaglia 1999). Al- most all of these new records are for species that have been recorded from neighboring states, but were not recorded from Dela- ware. Hylocurus flaglerensis Blackman was not previously reported from the area, hay- ing been only known from Florida (Rabag- lia 2003, also reported this species as new in’ Maryland). Pityophthorus Blackman is reported for the first time north of North Carolina. Of the 64 species now reported from Del- aware, 14 are not native to North America. Eight of these are ambrosia beetles in the tribe Xyleborini. Atkinson et al. (1990) pointed out the increasing occurrence of these beetles in the eastern United States. Several species recorded from neighbor- scriptor 313 ing states likely will be found with further collecting in Delaware. In addition, Dela- ware’s location on the east coast and in- creasing international trade in the area will result in additional exotic species becoming established. Three exotic species, Tomicus piniperda (L.), Hylastes opacus Erichson, and Xyleborus pfeili Eichhoff, and two western North American species of Hyle- sinus have recently been reported in Mary- land (Vandenberg et al. 2000, Rabaglia and Williams 2002, Rabaglia 2003). Hoebeke (1991) reported the exotic ambrosia beetle, Ambrosiodmus lewisi (Blandford), for the first time in North America in southeastern Pennsylvania, near Delaware. Abbreviations for repositories of speci- mens new to Delaware are: UDCC—Uni- versity of Delaware, Newark DE; RJRC— Robert J. Rabagha collection, Annapolis MD. Numbers of specimens appear in pa- rentheses with abbreviations. published records are from Wood and Bright 1992, Bright and Skidmore 1997. Sources of New DELAWARE RECORDS OF SCOLYTIDAE Subfamily Hylesininae Hylastes porculus Erichson Distribution.—USA: AL, AR, CT, DC, FL, GA, IN, LA, ME, MD, MA, MI, MN, MSs NENT, NY. NG, PAY SG; SD: IN; TX, VT, VA, WV, WI, DE: Kent Co., Little Creek Wildlife Area, 22 April 1998, M. A. Valenti coll. (2, RJRC); Sussex Co., Wood- side, 20 April 1930, J. M. Amos coll. (1, UDEC): Hylastes salebrosus Eichhott Distribution USA: AL, AR, FL, GA, LA, MD, MS, NJ, NC, SC, TX, VA, DE: New Castle Co., Blackbird State Forest, Ty- bout Tract, 27 May 1997, M. A. Valenti coll. (1, RJRC); Sussex Co., Redden State Forest, HQ Tract, 16 May 1997, M. A. Val- enti coll. (2, RJRC). Hylastes tenuis Eichhoff Distribution USA: AL, AZ, AR, CA, DE FLY GA: ID; IN, KYLA, Mb, MA; 314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MS, NV, NM, NY, NC, OR, PA, SC, TN, TX, UT, VA, WV, DE: New Castle Co., Blackbird State Forest, Wright Tract, 27 May 1997, M. A. Valenti coll. (2, RJRC); Sussex Co., Redden State Forest, HQ Tract, 24 April 1998, M. A. Valenti coll. (1, RJRC). Hylurgops rugipennis pinifex (Fitch) Distribution —USA: AL, AZ, CA, CO, DC, GA, ID, ME, MD, MA, MI, MN, MT, NH, NJ, NM, NY, NC, OH, OR, PA, RI, SC, UT, VA, WV, WI, WY, DE: New Castle Co., Blackbird State Forest, Wright Tract, 30 April 1997, M. A. Valenti coll. (1, RJRC); Sussex Co., Redden State Forest, Appenzellar Tract, 24 June 1997, M. A. Valenti coll. (1, RJRC). Hylesinus aculeatus Say Distribution —USA: AR, CO, CT, DC, GA, IL, IN, IA, KS, LA, ME, MD, MA, MI, MN, MS, MO, MT, NE, NH, NJ, NY, NG INDY OH) OK) PAS SG) SDINT EX VA, WV, WI, DE: New Castle Co., New- ark, 3 May 1904, C. O. Houghton coll. (1, UDEE): Dendroctonus frontalis (Zimmermann) Distribution—USA: AL, AZ, AR, DC, FL, GA, KY, LA, MD, MS, NC, OK, PA, SG] UENS DX VASW VIDE Sussex) (Coe Oak Orchard, Jan. 5 1961, D. FE Bray coll. ex. Loblolly pine (9, UDCC). Dendroctonus valens LeConte Distribution USA: AZ, CA, CO, ID, IL, IN, KS, ME, MD, MA, MI, MN, MT, NV, NH, NJ, NM, NY, NC, OH, OR, PA, SD, UT, VT, VA, WA, WV, WI, WY, DE: New Castle Co., Blackbird State Forest, Wright Tract, 30 April 1997, M. A. Valenti coll. (2, RJRC); Kent Co., Blackiston Wild- life Area, 23 April 1998, M. A. Valenti coll. @eRIRE): Cnesinus strigicollis LeConte Distribution —USA: DC, FL, GA, IL, LA, MD, MI, MO, NJ, NY, NC, OH, PA, S@sGENE EXS WAS WIV DEssINewsGastle Co., Blackbird State Forest, White Rd Tract, 27 May 1997, M. A. Valenti coll. (1, RJRC); Kent Co., NG Wilder Wildlife Area, 26 May 1998, M. A. Valenti coll. (1, RJRC). Phloeotribus liminaris (Harris) Distribution USA: CT, DC, FL, IA, KY, MD, MA, MI, MS, NH, NJ, NY, NC, OH, PA, SC, VA, WV, DE: Kent Co. N: Dover, June 1999, M. A. Valenti coll. (1, RJRC); New Castle Co., Newark, April ISIS, UDEe): Phloeosinus dentatus (Say) Distribution —USA: AL, AR, DC, FL, GA, IL, IA, KS, KY, MD, MA, MN, MS, MO, NE, NH, NJ, NY, NC, OH, OK, PA, SC, SD, TN, TX, VA, WV, DE: Sussex Co., Prime Hook Wildlife Refuge, 27 April 1998, M. A. Valenti coll. (3, UDCC). Carphoborus bifurcus Eichhott Distribution —USA: DC, FL, GA, MD, MS, MO, NY, NC, OK, PA, TN, VA, WV, DE: Kent Co., Dover, 15 April 1998, Del Dept of Ag coll. ex. Ipslure-baited trap, (1, RJRC); Sussex Co., Redden State Forest, Appenzellar Tract, 1 May 1997, M. A. Val- enti coll. (1, RJRC). Hylocurus rudis (LeConte) Distribution USA: DC, GA, KS, KY, MD, MI, NJ, OH, PA, TX, VA, DE: Kent Co., Killens Pond SP, 1 July 1998, M. A. Valenti coll. (1, RJRC). Aylocurus flaglerensis Blackman Distribution —USA: FL, MD, DE: Kent Co., Blackiston Wildlife Area, 23 April 1998, M. A. Valenti coll. (1, RJRC). Micracisella opacicollis (LeConte) Distribution USA: DC, IL, GA, KS, MA, MD, MI, MN, MO, NJ, NY, NC, OH, PA, VA, WV, DE: Kent Co., Milford, 5 May 1932 (1, UDCC). VOLUME 105, NUMBER 2 Thysanoes fimbricornis LeConte Distribution —USA: DC, FL, IL, MD, MS, NJ, NC, PA, TX, VA, WV, DE: Kent Co., Milford Neck, 26 May 1998, M. A. Valenti coll. (6, RJRC). Ips grandicollis (Eichhoff) Distribution.—Australia; USA: AL, AR, Ci DEeA ELS GA, lk. IN, LA, ME, MD; MA, MI, MN, MS, MO, MT, NE, NH, NJ, INML ONY, ING, OH; PA, SC, TN, TX; VA, WV, WI, DE: New Castle Co., 24 Aug 1938, J. M. Amos coll. (1 UDCC); Sussex Co., Bethany Beach, 27 August 1974, Frank Boys coll. ex: Japanese black pine (2 WUDGE): Ips pini (Say) Distribution USA: AK, AL, AR, CA, CO, Cia DC) FL, GA, 1D) IL, IN, TA; KS, LA, ME, MD, MS, MI, MN, MS, MT, NE, NiV;, INE INI; NY, NC, OH, OR, PA, SD, TX, UT, VT, VA, WV, WI, WY, DE: New Castle Co., Wilmington, 14 April 1998, DDA coll. ex: ipslure-baited trap (1, RJRC). Orthotomicus caelatus (Eichhoff) Distribution.—Africa; USA: AK, AL, ARS GAYCONCT DG) FL, ‘GA, ID IE, IN, IA, KS, LA, ME, MD, MS, MI, MN, MS, MT, NE, NV, NH, NJ, NY, NC, OH, OR, PA, SD, TX, UT, VT, VA, WV, WI, WY, DE: Kent Co., Little Creek Wildlife Area, 22 April 1998, M. A. Valenti coll. (1, RJRC); Sussex Co., Redden State Forest, HQ Tract, 24 April 1998, M. A. Valenti coll. (1, RJRC). Pityogenes hopkinsi Swaine Distribution.—USA: CT, GA, IL, IN, IA, ME, MD, MA, MI, MN, NH, NJ, NY, NC, OH, PA, SC, TN, VA, WV, WI, DE: New Castle Co., Wilmington, 27 May 1998, DDA coll. ex: Ipslure-baited trap (2, RJRC). 315 Dryocoetes autographus (Ratzeburg) Distribution.—Africa, Asia, Europe; USA: AK, CA, CO, ME, MD, MI, MN, MT, NV, NH, NM, NY, NC, OR, PA, SD, TN, UT, VA, WA, WV, WI, WY, DE: New Castle Co., Blackbird State Forest, Tybout Tract, 13 June 1997, M. A. Valenti coll. (1, RJRC). Dryocoetes granicollis (LeConte) Distribution USA: AR, DC, FL, GA, IL, KY, LA, MD, MO, NJ, NY, NC, OH, PA, TX, VA, DE: New Castle Co., Newark, 2 June 1901) Gs WDEEe). Lymantor decipiens (LeConte) Distribution —USA: IA, KY, ME, MD, MA, MI, MN, MS, NJ, NY, OH, PA, VA, WV, DE: New Castle Co., Blackbird State Forest, Tybout Tract, 7 August 1997, M. A. Valenti coll. (3, RJRC). Xyloterinus politus Say Distribution USA: CT, DC, GA, IL, KY, ME, MD, MA, MI, MO, MS, NH, NJ, NY, NC, OH, PA, VA, WV, WI, DE: Kent Co., Little Creek Wildlife Area, 22 April 1998, M. A. Valenti coll. (1, RJRC); New Castle Co., Newark, 14 March 1958, Don- ald MacCreery coll. (1, UDCC); Sussex Co., Redden State Forest, HQ Tract, 24 April 1998, M. A. Valenti coll. (1, RJRC). Ambrosiodmus obliquus (LeConte) Distribution.—Africa, South America; USA: DC, FL, GA, LA, MD, MS, NC, VA, DE: Kent Co., Blackiston Wildlife Area, 17 June 1999, M. A. Valenti coll. ex: funnel trap (3, RJRC):; Sussex Co., Redden State Forest, Appenzellar Tract, 28 May 1997, M. A. Valenti coll. ex: turpentine and ethanol funnel trap (1, RJRC). Ambrosiodmus rubricollis (Eichhoff) Distribution.—Asia, Australia; USA: AL, CT; FL, MD, PA, SC, TN, VA, DE: Kent Co., Blackiston Wildlife Area, | Oc- tober 1999, M. A. Valenti coll. ex: funnel trap (3, RJRC). 316 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ambrosiodmus tachygraphus (Zimmermann) Distribution USA: AL, DC, FL, GA, IMDS MS; NJ; NG OHS PAY SG) WAG WV, DE: Sussex Co., Assawoman Wildlife Area, 27 April 1998, M. A. Valenti coll. (2, RJRC). Euwallacea validus (Eichhoff) Distribution.—Asia; USA: MD, NY, PA. DELAWARE: Kent Co., Blackiston Wild- life Area, 23 April 1998, M. A. Valenti coll. (2, RJRC); New Castle Co., Claymont, 20 May 1997, C. Bartlett coll. (1, UDCC); Sussex Co., Redden State Forest, HQ Tract, 24 April 1998, M. A. Valenti coll. (1, RJRC). Xyleborus atratus Eichhoff Distribution.—Asia; USA: FL, GA, MD, TN, VA, WV. DELAWARE: Kent Co., Blackiston Wildlife Area, 23 April 1998, M. A. Valenti coll. (5, RJRC); Sussex Co., Redden State Forest, HQ Tract, 24 April 1998, M. A. Valenti coll. (2, RJRC). Xyleborus celsus Eichhoft Distribution USA: AR, CT, DC, FL, GA, IL, IN, IA, KS, LA, MD, MI, MS, MOWNE NYSNG OHS RA. SC. VEX Wal VA, WV. DELAWARE: New Castle Co., Newark, 22 June 1990, C. Bartlett coll. ex: blacklight trap (4, UDCC). Xyleborus ferrugineus (Fabricius) Distribution.—Africa, Asia, Australia, South America; USA: AL, AZ, AR, DC, FL, GA, HI, IL, IN, KY, LA, MD, MA, MI, MS, MO, NJ, NY, NC, OH, PA, SC, TN, TX, VA, WV. DELAWARE: Kent Co., NG Wilder Wildlife Area, 26 May 1998, M. A. Valenti coll. (1, RJRC); New Castle Co., Newark, 21 June 1901 (1, UDCC): Sussex Co., Redden State Forest, Owens Tract, 24 June 1998, M. A. Valenti coll. (1, RJRC). Xyleborus pelliculosus Eichhoff Distribution.—Asia; USA: MD, PA. DELAWARE: Kent Co., Blackiston Wild- life Area, 23 April 1998, M. A. Valenti coll. (1, RJRC). Xyleborus pubescens Zimmermann Distribution.—USA: AL, AR, DC, FL, GA, KY, LA, MD, MS, NJ, NY, NC, PA, SC, TX, VA, WV, DE: Kent Co., Little Creek Wildlife Area, 25 June 1998, M. A. Valenti coll. (1, RJRC): New Castle Co., Newark, 20 May 1907 (2, UDCC); Sussex Co., Bethany Beach, April 9, 1932 (2, UDEC): Xyleborus sayi (Hopkins) Distribution —USA: CT, DC, IL, IN, KY, ME, MD, MI, MO, NJ, NY, NC, OH, PA, TN, VA, WV, DE: Sussex Co., Redden State Forest, HQ Tract, 11 August 1997, M. A. Valenti coll. (1, RJRC). Xylosandrus crassiusculus (Motschulsky) Distribution.—Africa, Asia; USA: FL, GA, LA, MD, MS, NC, SC, TX. DELA- WARE: Kent Co., Little Creek Wildlife Area, 22 April 1998, M. A. Valenti coll. (2, RJRC); Sussex Co., Redden State Forest, HQ Tract, 26 May 1998, M. A. Valenti coll. (2, RJRC). Xylosandrus germanus (Blandford) Distribution.—Asia, Europe; USA: CT, Tz. IN; (KY; NI; IN; INGs OH, PAS Wi: DELAWARE: Kent Co., Blackiston Wild- life Area, 23 April 1998, M. A. Valenti coll. (4, RJRC); New Castle Co., Newark, 21 May 1983, C. Sanford coll. ex: Liroden- dron tulipifera (1, UDCC); Sussex Co., Redden State Forest, HQ Tract, 24 April1998, M. A. Valenti coll. (1, RJRC). Hypothenemus crudiae (Panzer) Distribution.—Africa, Asia, South Amer- ica; USA: AL, AR, DC, FL, GA, IN, KS, Key, IEA; IMDS IMS? NISINY, NG sPAgsEe TN, TX, VA, WV. DELAWARE: Sussex Co., Redden State Forest, Owens Tract, 24 June 1998, M. A. Valenti coll. ex: Lindgren funnel trap with ethanol and yellow poplar branch (1, RJRC). VOLUME 105, NUMBER 2 Hypothenemus dissimilis (Zimmermann) Distribution.—USA: AL, CT, DC, FL, GA, IL, IN, LA, KY, MD, MN, MS, MO, NJFINY NG. OH: PA, SC, TNX VA. WV. DELAWARE: New Castle Co., New- ark, 21 May 1983, C. Sanford coll. ex: Lir- odendron tulipifera (1, UDCC). Hypothenemus eruditus Westwood Distribution.—Africa, Asia, Australia, Europe, South America; USA: AL, AR, CA, DC, FL, GA, IL, LA, MD, MI, MS, INET. NUE INY, NC, PA; SC, TN, TX, VA, WV. DELAWARE: New Castle Co., New- ark, 15 May 1906 (1, UDCC). Conophthorus coniperda (Schwarz) Distribution USA: ME, MD, MA, MI, MN, NH, NJ, NY, NC, PA, VA, WV, WI. DELAWARE: New Castle Co., Wilming- ton, 6 May 1993, J. Plummer coll. ex: Ips- lure-baited funnel trap (1, RJRC). Corthylus columbianus Hopkins Distribution USA: DC, FL, GA, IN, KS, MD, MA, MO, NJ, NC, SC, TN, VT, VA, WV. DELAWARE: Kent Co., Dover, 15 April 1998, Del Dept of Ag. coll. ex: Ipslure-baited trap (1, RJRC); New Castle Co., Iron Hill County Park, 24 August 1998, M. A. Valenti coll. (1, RJRC); Sussex Co., Redden State Forest, Owens Tract, 26 May 1998, M. A. Valenti coll. (1, RJRC). Gnathotrichus materiarius (Fitch) Distribution.—Europe; USA: AL, AR, DC, FL, GA, LA, ME, MD, MA, MI, MN, MS, MO, NE, NH, NJ. NY, NC, OK, PA, SC, TX, VA, VT, WV, WI. DELAWARE: Kent Co., Little Creek Wildlife Area, 22 April 1998, M. A. Valenti coll. (1, RJRC); Sussex Co., Redden State Forest, HQ Tract, 24 April 1998, M. A. Valenti coll. (2, RJRC). Monarthrum fasciatum (Say) Distribution USA: AR, CT, DC, FL, GA, IL, IN, IL, KS, KY, LA, MD, MA, MI, MS, MONI NY ING OHMPA, “SG “IN: 317 TX, VA, WV, WI. DELAWARE: Kent Co., NG Wilder Wildlife Area, 22 April 1998, M. A. Valenti coll. (2, RJRC); New Castle Co., Newark, 20 April 1912 (17, UDCC). Monarthrum mali (Fitch) Distribution USA: AL, AR, CA, CT, DE, FIL; Wb. IN, IL, TA; Key, LA, MEX Mp; MA, MI, MN, MS, NJ, NY, NC, OH, PA, TN, TX, VA, VT, WV, WI. DELAWARE: New Castle Co., Newark, 20 May 1907 (1, UDCC). Pityophthorus cariniceps LeConte Distribution USA: CT, IN, KY, ME, MD, MA, MI, MN, NH, NY, NC, OH, PA, SC, VA, WI. DELAWARE: New Castle Co., Blackbird State Forest, 27 May 1997, M. A. Valenti coll. (3, RJRC). Pityophthorus confusus Blandtord Distribution—USA: AL, AR, FL, GA, LA, MD, MS, NC, PA, SC, TX, VA, WV, DE: New Castle Co., Wilmington, 16 Au- gust 1993 (1, RJRC); Sussex Co., Redden State Forest, HQ Tract, 1 May 1997, M. A. Valenti coll. (2, RJRC). Pityophthorus liquidambarus Blackman Distribution USA: AR, CT, DC, FL, GA, IL, IN, IA, LA, MD, MS, NC, WV, DE: Sussex Co., Redden State Forest, HQ Tract, 28 July 1997, M. A. Valenti coll. (1, RJRC). Pityophthorus opaculus LeConte Distribution —USA: AK, AZ, CA, CO, ID, ME, MD, MI, MN, NV, NH, NM, NY, PA, SD, UT, WV, WY, DE: New Castle Co., Wilmington, 9 June 1993, Del Dept of Ag coll. ex. pinene-baited trap (1, RJRC). Pityophthorus puberulus (LeConte) Distribution —USA: DC, IN, KS, KY, ME, MD, MA, MI, MN, NH, JA, NJ, NY, NC, OH, PA, RI, VA, WV, WI, DE: New Castle Co., Wilmington, 28 April 1998, Del Dept of Ag coll. ex. pinene-baited trap (1, RJRC); Sussex Co., Redden State Forest, 318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Owens Tract, 24 June 1998, M. A. Valenti coll. (1, RJRC). Pityophthorus scriptor Blackman Distribution—USA: GA, MS, NC, OK, TN, TX. DELAWARE: Kent Co., Black- burn Wildlife Area, 23 April 1998, M. A. Valenti coll. (2, RJRC). Pseudopityophthorus minutisstmus (Zimmermann) Distribution —USA: AR, DC, FL, GA, IL, IA, KS, LA, ME, MD, MS, MA, MI, NH, NJ; NY; NG PA, SC, VA, WY, WI, DE: New Castle Co., Newark, 9 February 1984, D. F Bray coll. ex: oak firewood (12, UDCC). Pseudopityophthorus pubescens Blackman Distribution —USA: MD, NC, OK, VA, DE: New Castle Co., Iron Hill County Park, 21 April 1998, M. A. Valenti coll. (1, RJRC). List OF DELAWARE SCOLYTIDAE a = Species newly reported in Delaware b = Exotic species found in Delaware Hylesininae Hylastini Hylastes porculus Erichson—a Hylastes salebrosus Eichhoff—a Hylastes tenuis Eichhoff—a Hylurgops rugipennis pinifex (Fitch)—a Hylesinini Hylesinus aculeatus Say a Tomicini Dendroctonus frontalis (Zimmermann )—a Dendroctonus terebrans (Olivier) Dendroctonus valens LeConte—a Hylurgopinus rufipes (Eichhoff) Bothrosternini Cnesinus strigicollis LeConte—a Phloeotribini Phloeotribus liminaris (Harris)—a Phloeosinini Chramesus hicoriae LeConte Phloeosinus dentatus (Say)—a Polygraphini Carphoborus bifurcus Eichhoftf—a Scolytinae Scolytini Scolytus multistriatus (Marsham)—b Scolytus quadrispinosus (Say) Scolytus rugulosus (Mulier)—b Micracini Hylocurus rudis (LeConte)—a Hylocurus flaglerensis Blackman—a Micracisella opacicollis (LeConte)—a Thysanoes fimbricornis LeConte—a Ipini Ips calligraphus (Germar) Ips grandicollis (Eichhoff)—a Ips pini (Say)—a Orthotomicus caelatus (Eichhoff)—a Pityogenes hopkinsi Swaine—a Dryocoetini Dryocoetes autographus (Ratzburg)—a Dryocoetes granicollis (LeConte)—a Dryoxylon onoharaensum (Murayama)—a Lymantor decipiens (LeConte)—a Xyloterini Xyloterinus politus Say—a Xyleborini Ambrosiodmus obliquus (LeConte)—a Ambrosiodmus rubricollis (Eichhoff)—a, b Ambrosiodmus tachygraphus (Zimmer- mann)—a Euwallacea validus (Eichhoff)—a, b Xyleborinus saxeseni (Ratzeburg)—b Xyleborus affinis Eichhoff Xyleborus atratus Eichhoff—a, b Xyleborus californicus Wood—b Xyleborus celsus Eichhoff—a Xyleborus ferrugineus (Fabricius )—a Xyleborus pelliculosus Eichhott—a, b VOLUME 105, NUMBER 2 Xyleborus pubescens Zimmermann—a Xyleborus sayi (Hopkins )—a Xylosandrus crassiusculus sky)—a, b Xylosandrus germanus (Blandford)—a, b (Motschul- Cryphalini Hypothenemus crudiae (Panzer)—a, b Hypothenemus dissimilis (Zimmermann) Hypothenemus eruditus Westwood—a, b Hypothenemus seriatus (Eichhotf)—b a Corthylini Conophthorus coniperda (Schwarz )—a Corthylus columbianus Hopkins—a Gnathotrichus materiarius (Fitch) Monarthrum fasciatum (Say) Monarthrum mali (Fitch)—a a a Pityophthorus cariniceps LeConte—a Pityophthorus confusus Blandford—a Pityophthorus liquidambarus Blackman—a Pityophthorus opaculus LeConte Pityophthorus puberulus (LeConte) Pityophthorus pulicarius (Zimmermann) Pityophthorus scriptor Blackman—a Pseudopityophthorus minutissimus (Zim- mermann)—a Pseudopityophthorus pubescens Black- man—a a a ACKNOWLEDGMENTS Donald E. Bright identified or verified the identity of many of the specimens, his help is greatly appreciated. We also thank Charles Bartlett, University of Delaware, for the loan of specimens and the Delaware Department of Agriculture, Plant Industries Section, for providing specimens from their USDA Cooperative Agricultural Pest Sur- vey. Funding was provided through a USDA, Forest Service Focus Funding Grant. LITERATURE CITED Atkinson, T. H., R. J. Rabaglia, and D. E. Bright. 1990. Newly detected exotic species of Xyleborus (Co- leoptera:Scolytidae) with a revised key to species in eastern North America. Canadian Entomologist 122: 92-104. Bright, D. E. and R. J. Rabaglia. 1999. Dryoxylon, a new genus for Xyleborus onoharaensis Muraya- ma, recently established in the southeastern Unit- ed States (Coleoptera: Scolytidae). Coleopterists Bulletin 53(4): 333-337. Bright, D. E. and R. E. Skidmore. 1997. A Catalog of Scolytidae and Platypodidae (Coleoptera), Supple- ment | (1990-1994). National Research Council Press, Ottawa, Ontario. 368 pp. Hoebeke, E. R. 1991. An Asian ambrosia beetle, Aym- brosiodmus lewisi, new to North America (Cole- optera: Scolytidae). Proceedings of the Entomo- logical Society of Washington 93: 420-424. Phillips, C. E. 1978. Wildflowers of Delaware and the Eastern Shore. Delaware Nature Education Soci- ety, Inc., Hockessin, Delaware. 303 pp. Rabaglia, R.J. 2003. Annotated list of the bark and ambrosia beetles (Coleoptera: Scolytidae) of Maryland, with new distributional records. Pro- ceedings of the Entomological Society of Wash- ington. 105: 373-379. Rabaglia, R. J. and G. L. Williams. 2002. Two species of western North American Hylesinus Fabricius (Coleoptera: Scolytidae) new to the eastern United States. Proceedings of the Entomological Society of Washington. 104: 1058-1060. Taber, W. S. 1939. Delaware Trees: A Guide to the Identification of the Native Tree Species. Associ- ates International, Inc., Wilmington, Delaware. 254 pp. Tatnall, R. R. 1946. Flora of Delaware and the Eastern Shore. The Society of Natural History of Dela- ware. 313 pp. Vandenberg, N. J., R. J. Rabaglia, and D. E. Bright. 2000. New records of two Xyleborus (Coleoptera: Scolytidae) in North America. Proceedings of the Entomological Society of Washington, 102: 62— 68. Wood, S. L. and D. E. Bright. 1992. A Catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxonomic Index. Great Basin Naturalist Memoirs No. 13. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 320-330 SUBGENERA OF CLADONOTA STAL (HEMIPTERA: MEMBRACIDAE), WITH TWO NEW SPECIES FROM MEXICO DUANE J. FLYNN 1500 E. Garrison Blvd., Gastonia, NC Curator of Life Sciences, Schiele Musuem, 28054, U.S.A. (e-mail: dflynn49 @ yahoo.com) Abstract.—The subgenera of Cladonota Stal (Hemiptera: Membracidae: Hysoprorini) are reviewed. The subgenus Acanthonota Buckton is a new synonym of the subgenus Lobocladisca Stal, and the subgenus Lecythifera Fowler is reinstated. A key to subgenera of Cladonota and a checklist of the 47 valid species are provided. Twenty-four species are referred to different subgenera. Two new species of Cladonota are described and illustrated from Mexico: C. bulbosa and C. yucatanensis. Abstracta.—Se repasa los subgéneros de Cladonota Stal (Hemiptera: Membracidae: Hysoprorini). El subgénero Acanthonota Buckton es sinodnimo con el subgénero Lobo- cladisca Stal y el subgénero Lecythifera Fowler esta reinstalado como valido. Se encuentra una clave para los subgéneros de Cladonota junto con una lista de las 47 espécies validas. No hay combinaciones nuevas sino 24 espécies estan referidas subgéneros diferentes. Dos nuevas especies de Cladonota de México estan descritas e ilustradas: C. bulbosa y C. yucatanensis. Key Words: Members of the membracid genus Cla- donota Stal (1869c) are among the most or- nate of earth’s creatures (Figs. 1—6). The genus includes the largest number of spe- cies of the Tribe Hypsprorini (Membraci- nae) yet almost nothing is known of their biology. Funkhouser (1951a) stated the tri- lobed head (clypeus and two subantennal lobes), ocelli on an imaginary line through the middle of eyes, and free forewings as distinctive generic characters. Peldez (1945a) did a generic study of Cladonota (then known as Sphongophorus Fairmaire 1846a) for Mexico. McKamey (1998) listed 46 valid species in his catalogue. These species were placed in the genus Sphon- gophorus until McKamey (1997) found Sphongophorus to be a junior objective synonym of Hypsauchenia Germar (1835a), Membracidae, Cladonota, subgeneric key, Mexico, new species, treehoppers and McKamey (1997) elevated the subge- nus Cladonota Stal (1869c) to generic sta- tus to accommodate the numerous New World species incorrectly placed in Sphon- gophorus. Fifteeen species of Cladonota had never been placed in subgenera (McKamey 1998). The inclusion of the two new species, described below, increases the number of valid species in Cladonota to 47. Three characters appear to be useful for delimiting subgenera within Cladonota: (1) the presence or absence of a thorn or knob extension on the posterior margin of the an- terior process—10 of the 47 species have a tubercle or thorn on the posterior margin of the anterior process (Lobocladisca Stal) as in Figs. 3, 37 species do not (as in Fig. 2); (2) the presence or absence of an interme- diate process; 42 of the 47 species have an VOLUME 105. NUMBER 2 intermediate process (as in Fig. 4), five do not (Falculifera McKamey) as in Fig. 1; (3) the length of the anterior process when an intermediate process is present; 32 species have an intermediate process and the pos- terior margin of the anterior process is en- tire. In sixteen, the anterior process sur- passes the intermediate process (Cladonota Stal); and in 16 it does not (Lecythifera Fowler). Although no subgenus is distin- guished by one feature by itself, these dis- tinctions are useful, at least until a cladistic analysis of the species is undertaken. Goding (1928e) synonymized the sub- genus Lecythifera Fowler (1894c) with the subgenus Cladonota (as Sphongophorus) Stal (1869c). Fowler (1894c) had separated Lecythifera Fowler from Cladonota Stal by the length of the anterior process when an intermediate process is present: in Cladon- ota the anterior process surpasses the inter- mediate process; in Lecythifera Fowler the anterior process does not surpass the inter- mediate process. The aforementioned fea- tures delimit two distinct subgeneric groups. I therefore propose reinstating Le- cythifera Fowler as a valid subgenus. Buckton (1903a) created a new subgenus Acanthonota to accommodate one new spe- cies, Cladonota livida. The spatulate ante- rior process is the only character that sep- arates livida from species of the subgenus Lobocladisca Stal. I therefore propose Acanthonota Buckton (1903a) as a new synonym of Lobocladisca Stal (1869c). KEY TO SUBGENERA OF CLADONOTA (MODIFIED FROM FOWLER (1894c), BUCKTON (1903a)) 1. Pronotum with erect process present at approx- imately mid length (‘intermediate process”) in the form of a ball, knobbed extension or erect horny (Bis. Sy" Seyrw 3 aches cypetse sats es eee 2 — Pronotum without intermediate process (Fig. |) Me tenete Ra aaswtke tet derek. Falculifera McKamey 2. Posterior margin of anterior process with small tooth-like extension (Fig. 3)... Lobocladisca Stal — Anterior process entire, lacking process on pos- terior margin (Fig. 2) 3. Anterior process does not surpass the inter- mediate process (Fig. 4) ... Lecythifera Fowler Ww to — Anterior process surpasses the intermediate process! (Fig. 2)) 5.02 se eye ce ee Cladonota Stal The following are descriptions of two new species of Cladonota (Lecythifera) from Mexico. As for most species in the genus, host plant data are lacking for these species. Cladonota (Lecythifera) bulbosa Flynn, new species (Figs. 5—7) Type locality—MEXICO. Jalisco, 7 km N. Malacque. Diagnosis.—Cladonota bulbosa is the only species in the genus with a large bulb attached to a stalked intermediate process. Description.—Female: Unknown. Male: Head (Fig. 6): Ocelli closer to eyes than to each other, bisected by an imaginary line that passes through center of eyes; su- pra antennal lobes elongate, rounded, not attaining apex of clypeus; clypeus nearly twice as long as wide, clypeus apex round- ed and pilose. Thorax: Pronotum with an- terior process arching posteriorly, extend- ing above and nearly touching top of large pronotal bulb of intermediate process, slightly compressed at mid length on pos- terior face of anterior pronotal process, apex heart shaped and glabrous (Fig. 7); intermediate process with three pronotal bulbs in one plane (Fig. 5): small bulb di- rected anteriorly, large subspherical bulb directed dorsoposteriorly and a small bulb formed by a constriction on anterior face of large bulb; large bulb approximately 3 times longer (in lateral view) and about 20 times the volume than small basal anteri- orly directed bulb of intermediate process; midline of large bulb defined by thin, nar- row groove; pronotal surface cancellate— reticulate; humeral angles blunt and ob- tuse; Sparse erect setae on pronotum longer on large posterior pronotal bulb; posterior pronotal process ends in bulb nearly equal in size to small basal posteriorly directed bulb of intermediate process, not attaining apex of forewing. Forewing: Coriaceous with broad apical limbus. 322 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. 1, Cladonota (Falculifera) apicalis, habitus, lateral view (Sphongophorus ballista redrawn from Pelaez 1945a). 2, C. (Cladonota) latifrons, habitus, lateral view (redrawn from Pelaez 1945a). 3, C. (Lobocladisca) biclavatus, habitus, lateral view (redrawn from Striimpel 1973a). 4, C. (Lecythifera) yucatanensis, habitus, lateral view. Color: Head with frontoclypeus reddish brown, frequently covered with white to- mentose; eyes brownish yellow. Thorax with anterior pronotal process from meto- pidium to half length of anterior process brownish yellow; remainder of anterior process to apex and posterior process (pos- terad of intermediate process) reddish brown; large bulb with sunken surface dark red brown, raised surface black. Fore- eS) im) es) VOLUME 105, NUMBER 2 Figs. 5-7. Cladonota (Lecythifera) bulbosa. 5, Habitus, lateral view. 6, Head, anterior view. 7, Detail of apex of anterior pronotal process, dorsal view. 324 wing reddish brown. Legs: Tibiae brown- ish yellow; tarsi yellowish. Abdomen: Reddish brown. Dimensions (mm): Length from anterior margin of eyes to apex of forewing in re- pose: 5.5—6.5 (holotype: 6.5); height from dorsal margin of eyes to apex of anterior process: 6.0—7.0 (holotype: 7.0); height from posterolateral margin of pronotum to dorsal margin of intermediate process: 4.5— 4.9 (holotype: 4.9); width of large pronotal bulb (with small constricted bulb in lateral view): 3.5—4.0 (holotype: 4.0) Material examined.—Holotype (¢) and three paratypes (d) from the University of Georgia Entomological Collection are la- beled: ““MEX. Jalisco 7/km N. Malacque/ 16, 19 July 1990/J. E. Wappes.” Holotype male additionally labeled ““HOLOTYPE/ Cladonota/bulbosa Flynn” deposited in Na- tional Museum of Natural History, Smith- sonian Institution, Washington, DC; two paratypes (d) additionally labeled *“‘-PARA- TY PE/Cladonota/bulbosa Flynn” deposited in University of Georgia Entomological Collection and one paratype male deposited in author’s collection. All type placements are with permission of the University of Georgia Entomological collections. Distribution.—Mexico. Discussion.—This new species is closely allied to Cladonota inflata (Pelaez) and Cladonota plummeri (Peldez), each of which also have a large bulb on the pro- notal posterior process. Cladonota bulbosa differs from the aforementioned species in having the bulbs attached to a stalked ver- tical process as opposed to lying directly on, or being an extension of, the pronotal posterior process. Cladontoa bulbosa is named for the large diagnostic bulb on the intermediate process of the species. Cladonota (Lecythifera) yucatanensis Flynn, new species (Figs. 8-10) Type locality—MEXICO: Yucatan, km E. Chichén Itza. Diagnosis.—Cladonota yucatanensis 1s No PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON distinguished by an anterior pronotal pro- cess that does not arch posteriorly as far as the base of an intermediate process that has three lobes. Description.—Male: Unknown. Female: Head (Fig. 9): Ocelli on bases directed lat- erad toward eyes, closer to eyes than each other, situated on an imaginary line that passes through center of eyes; supra-anten- nal lobes elongate, rounded, not attaining apex of clypeus; clypeus apex truncate, pi- lose with white tomentose on the underside. Thorax: Pronotum with anterior process weakly recurved with trilobed posteriorly directed intermediate process (Fig. 8); base of anterior process tubular with raised can- cellate-reticulate surfaces prominent; apex of anterior process expanded, heart-shaped with center deeply cleft (Fig. 10); interme- diate process almost equal to height of an- terior process with three lobes: one small lobe directed dorsoanteriorly, one small lobe directed ventroposteriorly and large lobe directed dorsoposterorly; lobes cancel- late reticulate; pronotal surface cancellate reticulate; humeral angles blunt and obtuse; pronotum covered with short, erect setae on pronotum longer along inferior margin of pronotum; posterior process ends in bulb, raised vertically over half distance to ven- troposterorly directed intermediate process bulb, bifurcate on top, not attaining tip of forewings. Forewing: Coriaceous with clear cell below knob at tip of posterior pro- cess. Color: Head with frontoclypeus reddish brown grading to dark brown toward me- topidium, becoming lighter toward clypeus: ocelli yellowish; eyes brownish yellow with outermost facets reddish. Thorax with pron- otum reddish brown, covered in white to- mentum; intermediate process darker; raised surfaces reddish brown with deeper surfaces black; tip of anterior pronotal pro- cess black. Forewing: Dark reddish brown. Legs: Tibiae brownish yellow; tarsi darker. Abdomen reddish brown with white tomen- tum. Dimensions (mm): Length from anterior VOLUME 105, NUMBER 2 es) ) Figs. 8-10. Cladonota (Lecythifera) yucatanensis. 8, Habitus, lateral view. 9, Head, anterior view. 10, Detail of apex of anterior pronotal process, lateral view. Nn 326 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON edge of eyes to tip of the forewing: 6.0; height from top of eyes to apex of anterior pronotal process: 7.0; height from postero- lateral margin of pronotum to dorsal margin of intermediate process: 5.6; width across eyes: 125! Material examined.—Holotype (2) from the University of Georgia Entomological Collection labeled: ““MEXICO: Yucatan/2 km E. Chitzen Itza/16 June 1990/R. Turn- bow” and additionally labeled: ““HOLO- TY PE/Cladonota/yucatanensis Flynn,” de- posited in the National Museum of Natural History, Smithsonian Institution, Washing- ton, DC with permission of the University of Georgia Entomological collections. Distribution.—Mexico. Discussion.—Cladonota yucatanensis most resembles Cladonota hoffmanni (Pe- laez) but differs in having (1) the apex of the anterior process anterior to, rather than above, the intermediate process and (2) an intermediate process with three lobes (Cla- donota hoffmanni has four). Cladonota yu- catanensis is named after the state of the type location. LIST OF SPECIES OF CLADONOTA The letter suffixes following citation dates are those used in the major biblio- graphic references for the Membracoidea based on year of citation: Prior to 1956: Metcalf and Wade (1965a); 1956-1980: Deitz and Kopp (1987a); 1981-1987: Deitz (1989a); 1988-1998: McKamey (1998). Synonomies are given for each species, cit- ing the first usage of a name combination. All the subgenera have unique combina- tions of the features mentioned in the sub- generic review. If each of these features evolved once, some of the subgenera may eventually become synonyms. In the ab- sence of a cladistic estimate for the genus, the subgeneric classification and place- ments proposed here are expected to be sta- ble. An asterisk (*) indicates species with new subgeneric assignment. Subgenus Lobocladisca Stal (1869c) bennetti (Kirby). Venezuela, Trinidad. Sphongophorus (Lobocladisca) ben- netti Kirby 1829a: 20. Cladonota (Lobocladisca) bennetti (Kirby); McKamey 1997a: 192. biclavata (Westwood). Mexico to Panama. Sphongophorus (Lobocladisca) bicla- vatus Westwood 1840a: 432. Centrotus (Lobocladisca) biclavatus Westwood; Duncan 1843a: 286. Sphongophorus (Lobocladisca) gueri- nii Fairmaire 1846a: 262. Sphongophorus spatulatus Fairmaire 1846a: 262. Hysauchenia spatulata Fairmaire 1846c: 13. Hypsauchenia guerinii Fairmaire 1846c: 13. Sphongophorus bivexilliffer Costa 1862: 150; Funkhouser 1951a: 53. Lobocladisca guerinii Fairmaire; Buckton 1903a: 82. Lobocladisca bivexilliffer Buckton 1903a: 82. Sphongophorus (Lobocladisca) dorsa- lis Buckton 1903a: 80. Sphongophorus parvulus 1903a: 80. Spongophorus [sic] guerini [sic] Fair- maire; Comstock 1940a: 404. Cladonota (Lobocladisca) biclavata (Westwood); McKamey 1997a: 193. clavaria (Fairmaire). Brazil, Ecuador. clavaria Fairmaire Costa; Buckton Hypsauchenia 1846c: 13. Sphongophorus (Lobocladisca) cla- varius Fairmaire; Stal 1869c: 275. Lobocladisca clavaria Fairmaire; Buckton 1903a: 82. Sphongophorus clavaria Fairmaire; Funhouser 1927f: 67. Cladonota (Lobocladisca) clavaria (Fairmaire) McKamey 1997a: 193. *livida (Buckton). Brazil, Peru. Sphongophorus (Acanthonota) lividus Buckton 1903a: 81. VOLUME 105, NUMBER 2 Cladonota livida (Buckton); Mce- Kamey 1997a: 190. lobulata (Stal). Colombia. Sphongophorus (Lobocladisca) lobu- latus Stal 1869c: 276. Lobocladisca lobulatus Stal; Buckton 1903a: 82. Cladonota (Lobocladisca) lobulata (Stal); McKamey 1997a: 193. *occidentalis (Striimpel). Colombia. Sphongophorus occidentalis Striimpel 1973a: 335. Cladonota occidentalis (Striimpel): McKamey 1997a: 189. *orientalis (Striimpel). Colombia. Sphongophorus orientalis Strtimpel 1973a: 338. Cladonota orientalis McKamey 1997a: 189. rigida (Stal). Colombia. Sphongophorus (Lobocladisca) rigi- dus Stal 1869c: 275. Lobocladisca_ rigidus Stal; Buckton 1903a: 82. Cladonota (Lobocladisca) rigida (Stal); McKamey 1997a: 193. spatulata (Fairmaire). Brazil. Sphongophorus spatulatus Fairmaire 1846a: 262. Hysauchenia 1846c: 13. Spongophorus [sic] spatulatus Fair- maire; Funkhouser 1927f: 67. Sphongophorus (Lobocladisca) spatu- latus Fairmaire; Stal 1869c: 275. Cladonota (Lobocladisca) spatulata (Fairmaire); McKamey 1997a: 193. vexillifera (Goding). St. Vincent Island, WI. Sphongophorus (Lobocladisca) vexil- lifera Goding 1893b: 53. Sphongophorus intermedius Buckton 1903a: 80. Lobocladisca_ vexillifera Buckton 1903a: 82. Sphongophorus (Lobocladisca) vexil- liferus Goding: Metcalf & Wade 1965a: 1384. (Strtimpel); spatulata Fairmaire Goding; 327 Cladonota (Lobocladisca) vexillifera (Goding); McKamey 1997a: 193. Subgenus Falculifera McKamey (1997) apicalis (Stal 1869c). Mexico to Panama. Hypsauchenia balista Amyot & Ser- ville 1843a: 535. Sphongophorus ballista Amyot & Ser- ville; Fairmaire 1846a: 261. Sphongophorus (Sphongophorus) bal- ista [sic] Amyot & Serville; Goding 1930b: 8. Spongophorus (Sphongophorus) apt- calis Stal 1869c: 273. Spongophorus (Sphongophorus) amy- oti Metcalf & Wade 1965: 1368 [er- ror]. Cladonota (Falculifera) apicalis Stal; McKamey 1997a: 192. *bolivari (Pelaez 1945a). Mexico. Sphongophorus bolivari Pelaez 1945a: 66. Cladonota bolivari Pelaez; McKamey 1997a: 189. clavigera (Stal 1864a). Mexico to Costa Rica. Sphongophorus (Sphongophorus) cla- viger Stal 1864a: 68. Sphongophorus (Sphongophorus) cla- vigerus Stal; Goding 1893a: 467. Cladonota clavigera Stal; McKamey lOO Tas 192: *luctuosa (Pelaez 1945a). Mexico. Sphongophorus luctuosus Pelaez 1945a: 68. Cladonota luctuosa Pelaez; McKamey 1997a: 189. *zeledoni (Pelaez 1967a). Costa Rica. Sphongophorus zeledoni Pelaez 1967a: 209. Cladonota zeledoni Pelaez; McKamey 1997a: 190. Subgenus Lecythifera Fowler (1894c) *affinis (Fowler 1894c). Guatemala. Sphongophorus (Lecythifera) affinis Fowler 1894c: 29. Sphongophorus (Cladonota) affinis 328 Fowler; Metcalf & Wade 1965a: 1372. Cladonota (Cladonota) affinis Fowler; McKamey 1997a: 190. *brunnea (Fallou 1890a). Brazil. Sphongophorus 1890a: 354. Cladonota brunnea Fallou; McKamey 1997a: 189. bulbosa Flynn, n. sp. Mexico. *championi (Fowler 1894c). Mexico, Gua- temala. Sphongophorus (Lecythifera) cham- pioni Fowler 1894c: 28. Sphongophorus (Cladonota) cham- pioni Fowler; Metcalf & Wade 1965a: 1373. Cladonota (Cladonota) championi Fowler; McKamey 1997a: 190. *costata (Buckton 1903a). St. Vincent Is- land., W.1. Hypsoprora costata Buckton 1903a: ol. Sphongophorus costata Funkhouser 1927f: 67. Sphongophorus (Cladonota) costatus Fowler; Goding 1928e: 228. Cladonota (Cladonota) costata Buck- ton; McKamey 1997a: 191. *falleni (Stal 1862e). Brazil. Sphongophorus (Cladonota) falleni Stal 1862e: 24. brunneus Fallou Buckton; Cladonota_ falleni Stal; Buckton 1903a: 83. *eonzaloi (Pelaez 1945a). Mexico. Sphongophorus gonzaloi Pelaez 1945a: 68. Cladonota gonzaloi Pelaez; McKamey 1997a: 189. *hoffmanni (Pelaez 1940a). Mexico. Sphongophorus hoffmanni Pelaez 1940a: 285. Cladonota hoffmanni Pelaez; Mc- Kamey 1997a: 189. *inflata (Fowler 1894c). Guatemala to Pan- ama. Sphongophorus (Lecythifera) inflatus Fowler 1894c: 30. (Cladonota) Sphongophorus inflatus PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fowler; Metcalf & Wade 1965a: 1374. Cladonota (Cladonota) inflata Fowler; McKamey 1997a: 191. */ocomotiva (Breddin 1901a). Ecuador. Sphongophorus locomotiva Breddin 1901a: 201. Sphongophorus (Cladonota) locomo- tivus Breddin; Goding 1928e: 228. Cladonota (Cladonota) locomotiva Breddin; McKamey 1997a: 191. *machinula (Breddin 1901a). Ecuador. Sphongophorus machinula Breddin 1901a: 201. Sphongophorus (Lobocladisca) mach- inulus Breddin; Goding 1928e: 229. Cladonota (Lobocladisca) machinula Breddin; McKamey 1997a: 193. *pieltaini (Pelaez 1945a). Mexico. Sphongophorus 194Sa: 68. Cladonota pieltaini Pelaez; McKamey 1997a: 189. *plummeri (Pelaez 1945a). Mexico, Gua- temala. pieltaini Pelaez Sphongophorus plummeri Pelaez 1945a: 67. Cladonota plummeri Pelaez: Mc- Kamey 1997a: 189. *robustula (Fowler 1894c). Guatemala to Panama. Sphongophorus (Lecythifera) robustu- lus Fowler 1894c: 29. Sphongophorus (Cladonota) robustu- lus Fowler; Metcalf & Wade 1965a: 1378. Cladonota (Cladonota) robustula Fowler; McKamey 1997a: 192. *siparuna (Striimpel 1973a). Colombia. Sphongophorus siparuna Striimpel 1973a: 344. Cladonota siparuna Strtimpel: Mc- Kamey 1997a: 190. yucatanensis Flynn, n. sp. Mexico. Subgenus Cladonota Stal (1869c) albofasciata (Goding 1893b). St. Vincent Island, W.I. VOLUME 105, NUMBER 2 Sphongophorus ciata Goding 1893b: 54. Sphongophorus (Cladonota) albofas- ciatus Goding 1893a: 467. Cladonota albofasciata Goding; Buck- ton 1903a: 83. Cladonota (Cladonota) albofasciata Goding; McKamey 1997a: 190. amazonica (Andrade 1978a). Brazil. Sphongophorus (Cladonota) amazoni- cus Andrade 1978a: 1. Cladonota (Cladonota) amazonica Andrade; McKamey 1997a: 190. atrata (Fonesca 1936a). Brazil. Sphongophorus (Cladonota) atratus Fonesca 1936a: 162. Cladonota (Cladonota) atrata Fones- ca; McKamey 1997a: 190. cinerea (Fonesca 1933a). Brazil. Sphongophorus (Cladonota) cinereus Fonesca 1933a: 445. Cladonota (Cladonota) cinerea Fones- ca; McKamey 1997a: 191. crassepunctata (Sakikabara 1971b). Brazil. Sphongophorus (Cladonota) crasse- punctatus Sakikabara 1971b: 185. Cladonota (Cladonota) crassecpuntata Sakikabara; McKamey 1997a: 191. foliata (Funkhouser 1922a). Brazil. Sphongophorus (Cladonota) foliatus Funkhouser 1922a: 1. Cladonota (Cladonota) foliata Funk- houser; McKamey 1997a: 191. *fritzi (Sakikabara 1976e). Brazil. Sphongophorus fritzi Sakikabara 1976e: 159. Cladonota fritzi Sakikabara; Kamey 1997a: 189. gracilis (Sakikabara 1971b). Brazil. Sphongophorus (Cladonota) gracilis Sakikabara 1971b: 187. Cladonota (Cladonota) gracilis Saki- kabara; McKamey 1997a: 191. *euimaraest (Sakikabara 1981b). Brazil. Sphongophorus guimaraesi Sakika- bara 198 1a: 85. Cladonota guimaraesi Sakikabara; McKamey 1997a: 189. (Cladonota) albofas- Mc- 329 latifrons (Stal 1869c). Mexico, Guatemala, Brazil. Sphongophorus (Cladonota) latifrons Stal 1869c: 274. Sphongophorus (Cladonota) nodosus Buckton 1903a: 79. Sphongophorus inelegans Buckton 1903a: 82. Sphongophorus nodosis |sic| Buckton; Funkhouser195 1a: 54. Cladonota (Cladonota) latifrons Stal; McKamey 1997a: 191. *lopezi (Strumpel 1973a). Colombia. Sphongophorus lopezi Strtimpel 1973a: 346. Cladonota lopezi Striimpel; McKamey L99OTa: 189. mirabilis (Pairmaire 1846a). Trinidad, Ve- nezuela, Brazil. Sphongophorus mirabilis Fairmaire 1846a: 261. Hypsauchenia 1846a: 13. Sphongophorus (Cladonota) mirabilis Fairmaire; Stal 1869c: 273. Cladonota mirabilis Fairmaire; Buck- ton 1903a: 82. Cladonota (Cladonota) mirabilis Fair- maire; McKamey 1997a: 191. paradoxa (Germar 1821a). Ecuador, Boliv- ia, Brazil, Argentina. Membracis paradoxa Germar 182 1a: 26. Hypsauchenia 1835a: 231. Sphongophorus paradoxa Germar; Fairmaire 1846a: 261. Sphongophorus (Cladonota) paradox- us Germar; Stal 1869c: 273. Sphongophorus (Cladonota) facetus Walker 1858a: 64; Funkhouser 192QT£: 67. Cladonota acetus Buckton 1903a: 82. Cladonota (Cladonota) paradoxa Ger- mar; McKamey 1997a: 191. ridicula (Walker 1858a). Mexico, Brazil. Walker mirabilis Fairmaire paradoxa Germar Sphongophorus — ridiculus 1858a: 64. 330 Sphongophorus (Cladonota) ridiculus Walker; Stal 1869c: 273. Cladonota (Cladonota) ridicula Walk- er; McKamey 1997a: 191. rufescens (Fonesca 1933a). Brazil. Sphongophorus rufescens Fonesca 1933a: 444. Sphongophorus (Cladonota) rufescens Fonesca; Metcalf & Wade 1965a: 1378. Cladonota (Cladonota) rufescens Fo- nesca; McKamey 1997a: 192. *trilobosa (Fonesca and Diringshofen 1969a). Argentina. Sphongophorus trilobosa Fonesca and Diringshofen 1969a: 151. Cladonota trilobosa Fonesca and Di- ringshofen; McKamey 1997a: 190. undulata (Walker 1851la). Brazil, Ecuador, Mexico. Sphongophorus (Cladonota) undulatus Walker 1851a: 498. Sphongophorus ludicrus Walker; Stal 1869c: 273. Cladonota undulatus Walker; Buckton 190 1a: 82. Cladonota ludicrous Walker; Buckton 1901a: 83. ACKNOWLEDGMENTS I thank Cecil Smith of the University of Georgia Entomological Collections, Ath- ens, GA, for loaning membracid specimens for identification, yielding two new species. I also thank Jim Craig of the Schiele Mu- seum, Gastonia, NC, for illustrating the new species; Lewis Deitz of North Carolina State University and Stuart McKamey of the Systematic Entomology Laboratory, U.S. Department of Agriculture, Washing- ton, DC, for helpful suggestions on the manuscript; and Ken Davis for the Spanish abstract translation. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Buckton, G. B. 1903a. A Monograph of the Membra- cidae. Lovell, Reeve & Co., Limited, London, England. Plates I-LX, 206 pp. Deitz, L. L. 1989a. Bibliography of the Membracoidea (Homoptera: Aethalionidae, Biturritiidae, Mem- bracidae, and Nicomiidae) 1981—1987. North Car- olina Agricultural Research Service Technical Bulletin 290, 32 pp. Deitz, L. L. and D. D. Kopp. 1987a. Bibliography of the Membracoidea (Homoptera: Aetalionidae, Bi- turritiidae, Membracidae, and Nicomiidae) 1956— 1980. North Carolina Agricultural Research Tech- nical Bulletin 284, 40 pp. Fairmaire, L. M. H. 1846a. Revue de la tribu des Membracides. Annales de la Société Entomolo- gique de France (2)4: 235-320. Fowler, W. W. 1894c. Biologia Centrali-Americana. Rhynchota. Hemiptera-Homoptera, Vol. II. Family Membracidae, xvili—ix, 1-173, Tables 1—10. Funkhouser, W. D. 1951a. Homoptera. Fam Membra- cidae. Genera Insectorum. Fascicle 208. Brussels, Belgium. Plates I-XIV, 383 pp. Germar, E. F 1835a. Species Membracidum Musae E.F Germari et dispostitio generum Membraci- dum. Revue Entomologique publiée par Silber- mann 3: 223-261 et suppl. 307-311. Goding, F W. 1928e. Membracidae of South America and Antilles, I. Subfamily Membracinae. Journal of the New York Entomological Society 36: 201— 234. McKamey, S. H. 1997. Nomenclatural changes in the Membracidae and Aetalionidae (Hemiptera: Membracoidae): species—group names and Sphon- gophorus Fairmaire, revised status. Steenstrupia 22: I-11. . 1998. Taxonomic Catalogue of the Membra- coidea (Exclusive of Leafhoppers), Vol. 60. Mem- oirs of the American Entomological Institute, Gainesville, Florida. 377 pp. Metcalf, Z. P. and V. Wade. 1965a. General Catalogue of the Homoptera. A Supplement to Fascicle I— Membracidae of the General Catalogue of Hemip- tera. Membracoidea. In Two Sections. North Car- olina State University, Raleigh. 1,552 pp. Pelaez, D. 1945a. Estudios sobre membracidos. V. Las espécies mexicanas del género Sphongophorus Fairmaire (Hem. Hom.). Anales de la Escuela Ciencias Biologicas 4: 53-146. Stal, C. 1869c. Bidrag till membracidernas kannedom. Ofversigt af Kongliga Svenska Vetenskaps-Aka- demiens Forhandlingar 1869: 231—300. Strimpel, H. 1973a. Die Membraciden-Fauna Kolum- biens 2. Die Gattung Sphongophorus Fairmaire, 1846. Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 4(80): 327-350. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 331-347 RE-EVALUATION OF THE ELAPHRIA FESTIVOIDES (GUENEE) SPECIES COMPLEX (LEPIDOPTERA: NOCTUIDAE) MICHAEL G. POGUE AND J. BOLLING SULLIVAN (MGP) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, Smithsonian Insti- tution, PO. Box 37012, MRC-168, Washington, DC 20013-7012, U.S.A. (e-mail: mpogue @sel.barc.usda.gov): (JBS) 200 Craven St., Beaufort, NC 28516-2119, U.S.A. Abstract.—Elaphria alapallida, new species, is described and illustrated. Elaphria fes- tivoides (Guenée) is re-evaluated and distinguished from EF. alapallida by the structures of the male vesica and female appendix bursa. In the male genitalia of E. festivoides, the cornuti in the vesica are larger and more numerous than in E. alapallida. In the female genitalia of E. festivoides, the appendix bursa is curved behind the corpus bursae, and in E. alapallida it projects vertically from lateral margin of corpus bursae. The distribution of collecting dates suggests E. festivoides is at least double brooded and E. alapallida is single brooded. Key Words: The Elaphria festivoides (Guenée 1852) species complex was evaluated in a previ- ous paper (Saluke and Pogue 2000) and at that time a new species, E. cornutinis Sa- luke and Pogue was described. After study- ing additional material and re-evaluating what Saluke and Pogue (2000) recognized as E. festivoides, we determined that an ad- ditional species is present. In this paper, we establish the identity of E. festivoides based on type comparison and describe a third species in the festivoides complex. Elaphria festivoides (Guenée) (Figs. 1, 3, 5-8, 13-14, 17, 20) Celaena festivoides Guenée in Boisduval and Guenée 1852: 220. Erastria varia Walker 1857: 808.—Draudt 1926: 220. Oligia festivoides: Smith 1889: 147.—But- ler 1891: 240.—Smith 1891: 43.—Dyar 19023 12% Monodes _ festivoides: Hampson — 1909: Elaphria alapallida, Elaphria festivoides, taxonomy, distribution 483.—Barnes and McDunnough 1917: 68.—Draudt 1926: 268. Elaphria festivoides: Kimball 1965: 99.— Franclemont and Todd 1983: 141.— Poole 1989: 351.—Poole and Gentili 1996: 739.—Saluke and Pogue 2000: 234, 235 (fig. 2) Diagnosis.—The forewing size is slightly larger in festivoides than in alapallida, which is evident when comparing a series of both species as there is overlap in the absolute measurements. The overall color- ation of the forewing is more drab in fes- tivoides than in alapallida because of the lack of rufous and white scales in festivoi- des, though some specimens of alapallida also lack the brighter scales. Markings of the forewings are generally less defined in festivoides than in alapallida, for example, the claviform spot is indistinct to absent in festivoides and is well developed in alapal- lida. The hindwing in festivoides is darker than in alapallida. In the male genitalia the 332 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. U.S.America. Povvleday gXxt'- 46 —11o. TYPE, > rae ‘ ‘" 4 lena peotivoideo ia 2 VEE Colourslile Z2& 7, EpastRra VARIA, laeid Tubdesat E. Flexida ax Sohas Olof: w Doubleday, ANS aT 2q. ®. : >, 4 Type specimens. 1, ¢ Holotype of Celaena festivoides Guenée. 2, Labels from holotype of Celaena festivoides Guenée. 3, 2 Holotype of Erastria varia Walker. 4, Labels from holotype of Erastria varia Walker. number and size of cornuti is diagnostic. In festivoides there are more (range 35—50, mean = 42.4, n = 7) cornuti and the longest cornutus is longer than in alapallida (range 0.45—0.70 mm, mean = 0.579 mm, n = 7). In alapallida the cornuti are fewer (range 21—35, mean = 27.0, n = 7) and the longest cornutus is shorter than in festivoides (range 0.35-0.50 mm, mean = 0.393 mm, n = 7). In the female genitalia, the appendix bursae curves around to the dorsal side of corpus bursae in festivoides; in alapallida the ap- pendix bursae projects vertically and is not curved around the corpus bursae. Description.—Male: Head: Labial palp with basal and median segments dark gray to black with ventral margin and interior cream, dorsal apex of median segment cream, apical segment cream with a few black and/or rufous Frons” with closely appressed scales cream, longer dor- sal scales pale rufous to rufous with a vary- ing amount of black scales forming a spot on eye margin where appressed and longer scales. VOLUME 105, NUMBER 2 scales meet. Vertex concolorous with longer scales of frons. Antennal scape white ven- trally, with varying amounts of black and rufous dorsally; flagellum gray to black. Thorax: Patagium pale rufous to brown mixed with varying amounts of white with a few darker scales. Tegula white with a few darker scales. Dorsal tuft between pa- tagia with cephalic half brown to dark ru- fous extending laterally to form a margin on tegula, rest white. Mesothorax tan. Metathorax tan with gray apex. Foreleg with femur gray speckled with white; tibia black with median band cream to pale ru- fous, this lighter medial area can vary in amounts of lighter colored scales; tarsi black with apical rings cream to cream mixed with pink. Midleg with femur white speckled with brown scales; tibia with lon- ger narrower scales in basal two-thirds cream mixed with gray to pale rufous, shorter scales in basal third black with white apical ring; tarsi black with white api- cal rings. Hindleg with femur and tibia mostly white mixed with gray scales; tarsi gray with white apical rings. Forewing: Length 10.5—14 mm (n = 11). Basal third of wing ground color very light brown; bas- al line black (indistinct to absent); anteme- dial line black, sinnuate (indistinct); orbic- ular spot, indistinct to very faint, round, a few darker scales in center, bordered by black scales nearest reniform spot; clavi- form spot absent; median line black, bor- ders reniform spot to postmedian line; a trapezoidal area between orbicular and re- niform spots may be completely filled with black; area between antemedial and post- median lines and below orbicular and re- niform spots light brown to brown, some specimens with a dark rufous patch; reni- form spot very light brown with darker scales internally; postmedial line straight, indistinct, pale, with black scales along proximal border; basal third very light brown to brown; veins outlined with black and white for a short distance from post- median line; gray along outer margin; fringe with basal half gray and apical half 333 light brown. Hindwing: Ground color gray; fringe white with darker basal line. Abdo- men: Dorsal tuft on first abdominal seg- ment absent; dorsum gray with an indistinct basal border of white scales; venter a mix- ture of white and gray scales. Genitalia (Figs. 13-14): Uncus curved, widest at proximal end, narrowing after curve to a hooked apex. Tegumen with an enlarged penicillum bearing an elongate, dense pen- iculum; a shorter, less sparse peniculum at inner base of penicillum. Pleurite C-shaped with a small muscle plate. Valve elbow shaped, curved, vertical, apex round; am- pulla arising from near base of valve, curved, constricted medially, apex pointed, a small triangular process just proximal to medial constriction. Juxta rectangulate with proximal margin produced medially into a pointed projection, chevron-shaped. Vin- culum V-shaped. Aedoeagus straight, wider at base than at apex; with an elongate, pointed, ventro-distal projection. Vesica in- tially curves ventrad then curves back around itself and ends dorsal to the apex of the aedoeagus; cornuti arranged in two par- allel rows, a dorsal row runs along edge of vesica then curves off this edge apically, a second semicircular lateral row with fewer cornuti; cornuti numerous (range 35—50, mean = 42.4, n = 7), with longest cornutus (range 0.45—0.70 mm, mean = 0.579 mm, n—a/): Female: As in male except forewing length 11.0—14.0 mm (n = 10). Hindwing slightly darker. Genitalia (Fig. 17): Papillae anales with round apex. Ventral plate of os- tium bursae an elongate rectangle; distal margin straight. Ductus bursae_ short, straight, sclerotized; wall invaginated; spic- ulate; basal projection prominent. Appendix bursae curved behind corpus bursae (Fig. 17). Corpus bursae ovate, narrowest at apex. Signum absent. Type material.—Celaena festivoides Guenée (Figs. 1-2). The 6d holotype [BMNH] is labeled: 1) Syn-type, 2) U.S. America. [printed], Doubleday, 46—110. 334 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5-8. Adults of Elaphria festivoides. 5, 2, Florida, Highlands Co., Archbold Biol. Sta., 12 Mar. 1997, M. G. Pogue, USNMENT 00063105. 6, 2, Florida, Putnam Co., Welaka, University Conservation Reserve, 12 Mar. 1962, D. C. Ferguson, USNMENT 63150. 7, d, North Carolina, New Hanover Co., MOTSU Buffer, Peter’s Point, 13 Sep. 1994, J. B. Sullivan, R. Broadwell, B. Smith. 8, d, Alabama, Baldwin Co., Bon Secour NWR, 15 Oct. 1986, T. L. Schieter, USNM 00063680. {handwritten], 3) Celaena festivoides [handwritten]. Erastria varia Walker (Figs. 3-4). The 2 holotype [BMNH] is labeled: 1) Type, 2) 2, 3) 432 [handwritten, yellow label], 4) glued on abdomen [handwritten], 5) colour slide, 224 [hand written], 6) 12. Erastria varia.[printed], 7) E. Florida, St Johns Bluff., E. Doubleday., 41.-7—29., 15 [hand- written]. Distribution (Fig. 20).—This species has principally a southeastern United States dis- tribution. The collection localities range from southeastern Kansas and central Mis- souri, south to southern Texas, east to Flor- ida, and north to North Carolina. Material examined.—137 ¢ and 184 @. U.S.A.: ALABAMA: BALDWIN CO., 1 mi E Oyster Bay, T9S, R4E, sec 6, sand dunes, 10-14 Mar. 1990 (4 6,5 2), Brown ?, genitalia slide MGP 1140; 1 mi E Oyster Bay, T9S, R4E, sec 7 NW, 13 Oct. 1990 (1 3), Brown; Bon Secour NWR, | Apr. 1994 (1 3), 17 Apr. 1994 (1 3), Pollock, 14 May 1994 (1 2), Seymour; Bon Secour NWR, foredunes, 17 Apr. 1993 (1 2), Brown, 15 Oct. 1996 (7 3, 3 2), Bon Secour NWR, hind dunes, 2 Apr. 1994 (1 dg, 1 2), Pol- lock; Bon Secour NWR, T9S, R2E, Sec. 24, 11 Mar. 1990 (1 2), Brown, 12-14 Oct. 1991 (1 2), Brown and Pollock: Bon Se- cour NWR, TOS) *R2Es Secs 25. 1490ct 1990 (2 36, 2 ¢), Brown; Bon Secour NWR, T9S, R2E, Sec. 25N, 11 Oct. 1991 (1 6,3 2), Brown and Pollock; Bon Secour NWR, T9S, R2E, Sec. 25S, 12—16 Oct. 1991 (9 3, 4 2), Brown and Pollock; Bon Secour NWR, T9S, R2E, Sec. 25S, dunes, 12-16 Oct. 1991 (1 do, 1 2), Brown and Pollock: Bon Secour NWR, T9S, R3E, Sec. VOLUME 105, NUMBER 2 1ONW, 11-15 Oct. 1991 (1 2), Brown and Pollock: nr. Bon Secour NWR, oak-pine forest, 17 Apr. 1993 (1 2), Brown and Pol- lock. DEKALB CO., DeSoto St. Pk., T6S, RIOE, Sec. 19SE—-20SW, 1,360—1,460 ft., 19 May 1990 (1 2), Brown and Pollock; DeSoto St. Pk., T6S, RIOE, Sec. 31SW, 1,240 ft., 20-24 May 1990 (1 6, 1 @), Brown and MacGown; DeSoto St. Pk., 25— 29 Sep. 1992 (1 6, 1 2), Brown and Pol- lock. LEE CO., 2.5 mi E I-85 & Hwy 29, 4 May 1995 (1 ¢), Brown and Vogt. JEF- FERSON CO., Pinson, 30 Mar. 1986 (1 @), Brown. MONROE CO., Haines Island Park, 4-5 Apr. 1995 (2 36, 2 2), MacGown. ARKANSAS: LOGAN CO., Cove Lake, TIN, R25W, sec. 35SE, 1,020 ft., 14-20 May 1989 (1 2), Brown. FLORIDA: [no locality], (1 3d, 2 2), 2 genitalia slide JFG 1757. DUVAL CO., St. Johns Bluff, 20 Apr. 1962 (1 6, 1 2), 21 Apr. 1962 (1 9), Ferguson. HAMILTON CO., Suwannee River St. Pk., 10 Apr. 1987 (2 @), Steiner. HIGHLANDS CO., Archbold Biological Station, 12 Mar. 1997 (1 3, 12 2), 2gen- italia slide USNM 46658, Pogue, 25 Mar. 1962 (1 @), Ferguson; Archbold Biological Station, Lake Placid, 17 Feb. 1985 (2 3), lSskeb. 1985e(1..2)3 19Feb. 1985 (1 9), 20sFebs 1985a(1"o); 21 Febs 1985 Cd od); DIGREDsA MOSS. 2), 2Z5keb. 1985 | G); 28 Mar. 1962 (1 6), Ferguson. INDIAN RIVER CO., Vero Beach, Dec. (1 2), Mal- lock. LEON CO., Tall Timbers Research Station, 26-27 Sep. 1972 (1 @), Todd. MARTIN CO., Jonathan Dickinson State Park, 20 Mar. 1963 (1 2), Covell. PUT- NAM CoO., University Conservation Re- serve, Welaka, 10 Mar. 1962 (1 @), 12 Mar. I9G28 Cigar ll 3S) 115 Mars 196252 d)5 21 Mar. 1962 (1 3), Ferguson; University Re- serve, Welaka, 8 Apr. 1962 (1 d), 9 Apr. 1962 (1 6), 10 Apr. 1962 (1 3d, 1 2), o genitalia slide USNM 46243, 2 genitalia slide USNM 46244, 12 Apr. 1962 (2 2), 15 Apr. 1962 (1 @), 19 Apr. 1962 (1 @), Fer- guson. VOLUSIA CO., Cassadega, 24 Mar. 1965 (1 2), Fuller. WAKULLA CO., Och- lockonee River State Park, 10 Oct. 1983 (1 335 3,1 2), Miller. GEORGIA: CHARLTON CO., Okefenokee National Wildlife Refuge, 20; Apr 1962 71d 2), 21) June 1991 (i 2), 20 May 1991 (1 @), Ferguson; Okefenokee Refuge Headquarters, nr. Folkston, 2 May 1981 (1 2), 3 May 1981 (1 3), Ferguson. KANSAS: CRAWFORD CO., 4 mi E Pitts- burg, 1-15 Sep. 1972 (1 6, 1 2), 3 geni- talia slide USNM 46260, Todd. DOUGLAS CO., Kansas University, 25 July 1937 (1 3), Fritz, LABETTE CO., Oswego, 2 Sep. 1965 (1 2), Hevel. KENTUCKY: CAL- LOWAY CO), Kenlake St: Pk., 16, Sep. 19795) 2)5 17 Sep. 1979 (i -2); Cornett. MCCRAKEN CO., vic. Paducah, 24 July 1976 (1 2), Covell. LOUISIANA: BOSS- IER PARISH: Barksdale Air Force Base, 18 Apr. 1996 (1 3), 24 Apr. 1996 (1 3), Pol- lock; Barksdale Air Force Base, calcareous prairie, 9 May 1996 (1 2), 14 Sep. 1996 (1 3), Pollock; Barksdale Air Force Base, shortleaf-pine forest, 24 Apr. 1996 (1 ¢), 1 May 1996 (1 3), 6 May 1996 (1 6, 1 &), 15 May 1996 (1 ¢), Pollock. MISSISSIPPI: FRANKLIN CO., Trib. of McGehee Crk., TON, R4E, Sec. 26 SW, 8 Apr. 1992 (1 9), MacGown and Schieter; GEORGE CO., 4.5 mi NNW Lucedale, TIS, R6W, Sec. 6 SE, 11 Mar. 1991 (1 @), Pollock, 29-30 Oct. 1989 (1 3, 4 ¢), Schiefer. HARRI- SON CO., Long Beach, 16 Oct. 1990 (1 2), 21 Oct. 1992 (1 6), Kergosien. HINDS CO., Clinton, 12 Apr. 1992 (1 3), Roshore, 23 Apr. 1966 (1 6), Mather; Jackson, 18 Mar. 1961 (1 2). JACKSON CO., 1 mi W Hwy 90 & 57, T7S, R8W, Sec. 25, 15 Apr. 1989 (1 36), Brown. LEE CO., Tombigee State Park, 1-23 Apr. 1993 (1 6, 2 @), 24 Apr.-7 May 1993 (1 2), Kergosien. LOWNOES CoO., TI9N, RI6E, Sec. 30 SE, 17 Apr. 1991 (1 3), Pollock. OKTIBBEHA CO., Starkville, 13 May 1983 (1 3d), Porter, 6 mi SW Starkville, 25 Apr. 1985 (1 @), 30 Apr. 1987 (1 2), Brown; A & M College, 29) Apral93ind 6); 18: June 1931 (1 2); Hutchins; TI8N, RI4E, Sec. 23, 18 Sep. 1987 (1 3), Brown. STONE CO., Sweetbay Bogs, T2S, R13W, Sec. 34 SW; 12 Mar. 1991 (1 2), Pollock. TISHOMINGO CO., 336 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 11 Figs. 9-12. 12 Adults of Elaphria alapallida. 9, 3, Connecticut, New Haven Co., West Rock Park, 24 May 1986, D. C. Ferguson, USNMENT 00063085. 10, 2, Connecticut, New Haven Co., Hamden, 14 June 1967, D. C. Ferguson, USNMENT 00063074. 11, d, Nova Scotia, Halifax Co., West Dover, 28 May 1955, D. C. Ferguson, USNMENT 00063370. 12, 2, North Carolina, Ashe Co., Mt. Jefferson State Park, 4.150 ft., hairpin turn, | June 2000, J. B. Sullivan, USNMENT 00164817. Tishomingo State Park, 11—12 Apr. 1986 (1 2), MacDonald. NORTH CAROLINA: BLADEN CO., Doe Island at Duck Pond (Bayfields), near White Oak, 9 Apr. 1994 (1 3), Broadwell. BRUNSWICK CO., 1 mi S Pretty Pond, Boiling Springs Lakes, 19 Apr. 1994 (2 ¢), Sullivan, Broadwell, Smith; Green Swamp, 29 Sep. 1994 (1 d, 1 @), Sullivan, Broadwell, Smith. CAR- TERET CO., Fort Macon State Park, brack- ish marsh, 5 Apr. 1997 (1 3), 3 genitalia slide MGP 1135, Sullivan; Fort Macon State Park, hardwoods, 2 May 1996 (1 2), 2 genitalia slide MGP 1136, Sullivan; Fort Macon State Park, brackish marsh, 13 Sep. 1996 (1 @), 14 Oct. 1996 (1 ¢), 3 genitalia slide MGP 1139, Sullivan. COLUMBUS CO., Lake Waccamaw, 10 May 1985 (1 &), Steiner, Gerberichh DARE CO., 18 May 1977 (1 ¢), Gifford. MOORE CO., Wey- mouth Woods Natural Area, 4 Apr. 2000 (2 2), Hartley, Helms, Hall; Niagara, 24 Apr. 1954 (1 2), 25 Apr. 1954 (1 2), 30 Apr. 1954 (2 2): Southern Pines, 8 May 1986 (1 3), Steiner. NEW HANOVER CO., Caro- lina Beach State Park, E of Dow Rd., coast- al fringe sandhill, 28 Apr. 1994 (1 d), Sul- livan, Broadwell, Smith; Carolina Beach State Park, Limesink Area, coastal fringe sandhill, 23 Mar. 1994 (1 ¢), d genitalia slide MGP 1133, 20 Sep. 1995 (1 6, 1 @), Sullivan, Broadwell, Smith; Fort Fisher Maritime Forest, coastal fringe evergreen forest, 13 Sep. 1994 (1 d), d genitalia slide MGP 1138; MOTSU Buffer, Peter’s Point, coastal fringe sandhills, 7 Mar. 1995 (1 d), 3 genitalia slide MGP 1134, 20 Apr. 1994 (1 2), 2 genitalia slide MGP 1137; 13 Sep. 1994 (1 dc), 22 Sep. 1994 (1 3), 27 Sep. 1994 (1 @), Sullivan, Broadwell, Smith. ONSLOW CO., Camp Lejune, Corn Land- ing, Juncus march, 2 May 1996 (1 2), Sul- livan. POLK CO., Tryon, 30 Apr. 1904 (1 O) 21) SMay Cli eo) Sakiske= Goll: VOLUME 105, NUMBER 2 337 Figs. 13-16. Elaphria species, 3 Park, maritime shrub, 14 Oct. 1996, J. B. Sullivan, genitalia slide MGP 1139. 14, aedoeagus and vesica of same specimen. 15, E. alapallida, West Virginia, Hardy Co., Lost River State Park, 29 June 1968, J. EF G. Clarke, USNM genitalia slide 46626. 16, aedoeagus and vesica of same specimen. OKLAHOMA: SEQUIA CO., Lake Ten- killer, 2 mi NW Blackgum, 25-29 Aug. 1982 (1 6,3 @), do genitalia slide USNM 46684, 14-16 July 1983 (1 @), Davis. SOUTH CAROLINA: CHARLESTON CO., Wedge Plantation, McClellanville, 26 Apr. 1980 (3 6, 2 2), 3 genitalia slide USNM 46261, Ferguson; Charleston, 28 Apr. 1962 (1 @), Ferguson. HORRY CO., Myrtle Beach, 21 Sep. 1940 (1 2), Mc- Elvare, 28 Sep. 1954 (1 2), McEvans, 30 Sep. 1953 (1 6). TEXAS: [no specific lo- cality], (3 d, 3 2), d genitalia slide JGF 1758. ANDERSON CO., Tennessee Colo- ny, 5 Mar. 1967 (1 3, 1 2), ¢ genitalia slide USNM 46680, 12 Mar. 1962 (1 2), genitalia. 13, E. festivoides, North Carolina, Carteret Co., Fort Macon State Blanchard. BASTROP CO., Bastrop State Park, 27 Feb. 1965 (1 ¢), 3d genitalia slide AB 666, 28 Feb. 1965 (1 6, 1 2), 3 gen- italia slide USNM 46679, Blanchard. BLANCO CO., Pedernales Falls State Park, 4 May 1973 (1 6), Blanchard. BOSQUE CO., Clifton, (1 d), Belfrage; Laguna Park, 24 Sep. 1970 (3 5, 2 2), 3d genitalia slide USNM 46677, Blanchard. BOWIE CO., 5 mi N New Boston, 17 Sep. 1972 (1 3), Todd; 10 mi NW New Boston, 18 Sep. 1972 (1 3), Todd. BRAZOS CO., College Station, June (1 @), @ genitalia slide USNM 46661, Glick, 10-17 Aug. (1 2), Sep. (1 2). BROWN CO., Lake Brown- wood State Park, 7 Sep. 1963 (1 6, 1 @), 338 Blanchard. CAMERON CO., Brownsville, 1-7 Mar. (1 6d), 8-15 Mar. (1 3),16 Mar. 1923 (1 3), 16-23 Mar. (2 3). CHAM- BERS CO., Black Jack Springs, (1 d). CO- MAL CO., Guadalupe River, nr. New Braunfels, 12 Mar. 1972 (1 6), Blanchard. DALLAS CO., Dallas, (1 2), vic. Irving, 29 Mar. 1997 (1 ¢), 14 Apr 1981 (1 Q), 12 Sep: 1981 G2); 20) Sepilosirdads 1 2), d genitalia slide USNM 46659, Rahn. HARRIS CO., Houston, (1 @), Doll; Spring, 9 Mar. 1965 (1 d), Blanchard. HAYS CO., Freeman Ranch, 6 km NW San Marcos, 13 Mar. 1995 (1 2), 2 genitalia slide USNM 46660, Steiner, Swearingen, Ott, Silverfine. JASPER CO., Town Bluff (Dam B), 10 Mar. 1965 (1 3), ¢ genitalia slide USNM 46239, Blanchard. KERR CO., Kerrville, 8 Mar. (1 2), 2 genitalia slide JGF 1759, Lacy, (4 2), 2 genitalia slide JGF 1761, Sep. 1902 (1 3), 3 geni- talia slide JGF 1760, Lacy, (5 3), 6 geni- talia slide USNM 46264. KIMBLE CoO., Junction, 16 June 1972 (1 2), 17 June 1972 (2) 518 June 1972 °C d572 2); Fereuson; 18 Sep. 1968 (1 2), Blanchard. LAMPA- SAS CO., 6.5 mi E Lampasas, 12 Mar. 1982 (8 3, 23 2), 2 o genitalia slides USNM 46263, 46678, Poole. SAN SABA CO., Bend, 11 Mar. 1982 (5 6, 2 2), 6 genitalia slide USNM 46682, Poole. ST. AUGUSTINE CO., 2 mi SW Broaddus Lake Sam Rayburn, 20 Sep. 1972 (1 6, 3 2), Todd. TARRANT CO., Fort Worth, 9 Sep. 1963 (1 2), 30 Sep. 1963 (3 2), Jackh. TRAVIS CO., Bee Cave, 14 Apr. 1964 (2 2), 2 genitalia slide USNM 46240, Blan- chard. UVALDE CO., Sabinal, Mar. 1910 (1 @), 10 May 1910 (1 3), 1 S genitalia slide USNM 46683, Pratt. Discussion.—Elaphria festivoides and E. alapallida are difficult to separate from lo- calities where both occur unless comparing large series. In the extreme south (Florida and Alabama), only festivoides occurs and in the north (Pennsylvania north) only ala- pallida. In other areas of the south and mid- Atlantic region both occur however, in North Carolina festivoides is found in xeric, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sandy areas along the coast and inland to the sandhills and has multiple broods (prin- cipally April-May and September—Octo- ber). Elaphria alapallida is in hardwood habitats from 2,000—6,000 feet and is single brooded (late April—June). Whether these habitat preferences hold true across the en- tire range of either species is yet to be de- termined. The collection dates (Fig. 19) show that E. festivoides is principally double brooded. Dates indicate that the first brood occurs from mid-February to the end of May with the second brood from late August to the end of October. The data shown are from the entire geographic range of EF. festivo- ides. These broods are certainly more de- fined if viewed from a more restricted range. For example, the dates in Florida range from mid-February to mid-April for the first brood and the only records are ear- ly October for the second brood. In Ala- bama the first brood is from mid-March to the end of May and the second brood is from the end of September to mid-October. In North Carolina the first brood is from end of March to mid-May and the second brood is from mid-September to mid-Oc- tober. Elaphria alapallida Pogue and Sullivan, new species (Figs. 9-12, 15-16, 18, 21) Elaphria festivoides: Rockburne and Lafon- taine 1976: 83 [misidentification].—Cov- ell 1984: 135 [misidentification].—Rings et al. 1992: 100 (fig. 30, 35) [misidenti- fication].—Saluke and Pogue 2000: 235 (figs. 1, 3-4), 237 (fig. 9), 238 (figs. 11— 13), 239 (figs. 17-18), 240 (figs. 21-22), 241 (figs. 25, 27) [misidentification]. Diagnosis.—See the previous diagnosis section regarding the differences between E. alapallida and E. festivoides. Description.—Male: Head: Labial palp with basal and median segments black with a few scattered rufous scales, ventral mar- gin and interior cream, apical segment VOLUME 105, NUMBER 2 ab 17 Figs. 17-18. ab r 18 Elaphria species, @ genitalia. 17, E. festivoides, Missouri, Benton Co., 4 mi NW Warsaw, along MO State UU, 7 May 1970, J. R. Heitzman, USNM genitalia slide 46699. 18, EF. alapallida, West Virginia, Tucker Co., Blackwater Falls State Park, 24 June 1968, J. EF G. Clarke, USNM genitalia slide 46272. with a few black and/or rufous Frons with closely appressed scales cream, longer dorsal scales pale rufous to rufous with a varying amount of black scales forming a spot on eye margin where appressed and longer scales meet. Vertex concolorous with longer scales of frons. Antennal scape white ventrally, varying amounts of black and rufous dorsally; fla- gellum gray to black. Thorax: Patagium variable, can be rufous, brown, or light brown and mixed with varying amounts of black scales. Tegula white with a few dark- er scales. Dorsal tuft between patagia with cephalic half black to brown extending lat- erally to form a margin on tegula, rest white. Mesothorax very light brown. Meta- thorax very light brown with brown to gray apex. Foreleg with femur gray speckled with white; tibia gray with median band or spot cream to rufous, this lighter medial area may vary in amounts of lighter colored scales; tarsi gray with apical rings cream to cream scales. cream mixed with pink. Midleg with femur white speckled with brown scales; tibia with longer narrower scales in basal two- thirds cream mixed with pale rufous to pink, shorter scales in basal third black with white apical ring; tarsi black with white api- cal rings. Hindleg with femur and tibia mostly white mixed with gray scales; tarsi gray with white apical rings. Forewing: Length 10-13 mm (n = 15). Basal third ground color white; basal line black (indis- tinct to absent); antemedial line white, sin- nuate; orbicular spot, round, a few darker scales in center, bordered by black scales nearest reniform spot; claviform spot white to cream variably bordered by a few black scales; median line black, borders reniform spot to postmedian line; a trapezoidal area between orbicular and reniform spots can be completely filled with black; area be- tween antemedial and postmedian lines and below orbicular and reniform spots a vary- ing mixture of rufous, light brown, and gray 340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Number of Specimens Collection Dates Fig. 19. Distribution of collecting dates of Elaphria festivoides and E. alapallida from the material examined. scales patches; reniform spot white with darker scales internally; postmedial line straight, white, with a few black scales along proximal border; basal third with an irregular white area bordering postmedian line and posterior margin; a white apical patch; gray along outer margin and rest with scattered rufous scales; fringe gray. Hindwing: tinct pale gray margin; fringe white with Dorsal tuft on absent: Ground color white with indis- darker basal line. Abdomen: first gray with an indistinct basal border of white abdominal segment dorsum scales; venter a mixture of white and gray scales. Genitalia (Figs. 15—16): Essentially same as E. festivoides except: cornuti fewer (range 21—35, mean = 27.0, n = 7), longest cornutus (range 0.35—0.50 mm, mean = 0.393 mm, n = 7). Female: As in male except forewing length 10-13 mm (n = 15). Genitalia (Fig. 18): Essentially the same as festivoides ex- cept: appendix bursae projects vertically from lateral margin of corpus bursae. Type material—The 3 _ holotype [USNM] is labeled: 1) Hamden, New Ha- ven Co., CONNECTICUT; 4. VI. 1966; D. C. Ferguson; 2) USNM ENT 00063094; 3) Holotype; Elaphria alapallida; Pogue and Sullivan. Paratypes: 265 ¢ and 188 @. CANADA: BRITISH COLUMBIA: Kaslo Creek, (1 6), d genitalia slide JGF 1764, Dyar. MANITOBA: Miniota, (1 ¢), Gib- bon. NEWFOUNDLAND): South Branch, 4 July 1949 (1 3), ¢ genitalia slide USNM 46311, Krogerus. NOVA SCOTIA: Bad- dock, Cape Breton Is., 30 July 1970 (2 3), Ferguson; Digby, (1 ¢), d genitalia slide JGF 1762, 18 June 1908 (1 2), 25 June 1908 (1 2), 26 June 1908 (1 2), 2 genitalia slide JGF 1763, Russell. ANNAPOLIS CO., South Milford, 23 June 1950 (1 @), Ferguson. HALIFAX CO., Bog E of Big Indian Lake, Halifax Watershed, 24 June VOLUME 105, NUMBER 2 1962 (2 3), 24 June 1963 (1 3), 24 June 1968 (1 3), 26 June 1968 (1 &); Boulder- wood, 26 June 1959 (1 3), 26 July 1959 (1 3); Halifax Watershed Area, 2 June 1955 (1 3), 5 June 1955 (1 3), ¢o genitalia slide USNM 46259, Ferguson; near Big Indian Lake, Halifax watershed, 24 June 1963 (1 6), Ferguson; Peggy’s Cove, 11 July 1953 (1 2), Ferguson; Purcell’s Cove, 6 June 1960 (1 2), 12 July 1962 (1 3), Ferguson; West Dover, 28 June 1955 (2 3), 3 geni- talia slide USNM 46258, 30 June 1968 (1 3), Ferguson. KINGS CO., Aylesford, 29 June 1962 (1 d), Ferguson; cattail swamp near Aylesford, 20 June 1963 (1 d), Fer- guson; top of North Mt., near Garland, 11 June 1954 (1 2), Ferguson. LUNEN CO., Green Bay, | July 1956 (1 2), Ferguson. QUEENS CO., Lake Kejimukujik, 27 May 1958 7Gise)s 17, sune: 1957) Gad, 2)53 July 1968 (1 d), Ferguson. YARMOUTH CO., Argyle, 7 June 1954 (1 6), Ferguson. ONTARIO: Hymers, 1-7 Aug. (1 @). U.S.A.: COLORADO: Vermeyjo R., 25 June (1 @), Packard. BOULDER CO., Mt. Flagstaff, 2,280 m, 20 June 1988 (1 d, 1 2), Peigler and Weissman. GARFIELD CO., Glenwood Springs, 16—23 July (1 d), 3 genitalia slide USNM 46252. JEFFER- SON CO., Chimney Gulch, 16 June 1920 (1 6), Oslar; Clear Creek, Oslar. CON- NECTICUT: NEW HAVEN CO., Bethany, 17 May 1968, (1 @), Ferguson; Bethany Center, 17 June 1967 (1 36, 1 @), Ferguson; Hamden, 4 June 1967, (2 ¢), 9 June 1967 (1 3d), 14 June 1967 (1 2), Ferguson; West Rock Park, 24 May 1966 (15 6, 1 @), Fer- guson; Yale Preserve, 9 May 1964 (2 3), 21 May 1964 (1 d), 7 June 1964 (2 gd, 1 2), d genitalia slide USNM 46247, @ gen- italia slide USNM 46248. DISTRICT OF COLUMBIA: (1 ?), 30 June (1 ?). DEL- AWARE: NEW CASTLE CO., Newark, 30 May 1983 (1 &), Bray. ILLINOIS: COOK CO., Evanston, 27 May 1895 (1 2). MA- CON CO., Decatur, 8 May 1915, (1 d). IN- DIANA: TIPPECANOE CO., Lafayette, 15 May 1916 (1 2), 9 Aug. 1915 (1 @), 13 Aus) 19155 (2) SKENTUCKYS BELT 341 CO., Pine Mountain State Park, 3 May 1974 (1 3), 13 May 1980 (1 6d), Cornett. BULLITT CO., Bernheim Forest, 17 Apr. 1977 (2 3, 1 &), Brownell; 20 May 1999 (1 2), 21 May 1999 (1 6, 1 @), Vander- pool. Mt. Washington, 23 May 1966 (1 °), 28 July 1988 (2 3), Covell. CARTER CO., Carter Caves, 29 Apr. 1972 (1 2), Covell. JEFFERSON CO., Valley Station, 9 May 1974 (1 2), 15 May 1974 (1 6, 1 2), 25 May 1974 (1 2), 9 Aug. 1974 (1 9), 19 Aug. 1974 (1 2), Scholz. POWELL CO., Tunnel Ridge Rd., 21 May 1988 (1 3), Covell. RUSSELL CO., Lake Cumberland State Park, 18 July 1972 (1 2), Cornett. WHITLEY CO., Cumberland Falls State Park, 16 May 1980 (1 ¢), Cornett. MAS- SACHUSETTS: [County unknown], Bev- erly Farms, 10 June 1913 (1 6), Reiff; HAMPDEN CO., Chicopee, 2 June 1899 (1 2), Knab. MIDDLESEX CO., Auburndale, 13 June 1974 (1 6), Covell; Concord, 3 June 1913 (1 2), Reiff. MARYLAND: ANNE ARUNDEL CO., Southaven, 17 May 1991 (1 @), Stevenson. CALVERT CO., Camp Bay Breeze, nr Lusby, 8 June 1963 (1 &), Flint; Flag Ponds, 9 May 1991 (1 2), Stevenson. CECIL CO., Elkton, 29 May 1983 (lo12)y Brays | PRINCE GEORGE’S CO., Patuxent National Wild- life Research Center, 12 May 1979 (1 @), d genitalia slide USNM 46267, Ferguson. MAINE: [no county], Oguossoc, 6 July (1 2); top Mt. Bigelow, 7 July 1937 (1 @). AROOSTOOK CoO., Ashland, 3 July 1943 (1 2); Houlton, 10 July 1945 (1 d). HAN- COCK CO., Bar Harbor, 7 June 1937 (2 3), Brower, 22 June 1933 (1 2), 28 June 1933 (1 do), 1 July 1934 (1 3), Brower, 4 July 1934 (1 3), 10 July 1935 (1 @), Brower, 17 July 1933 (1 ¢), Brower, 19 July 1934 (1 od), 24 July 1933 (1 2), Brower; Mt. Desert ‘Is., 13° June® 1932 (1S), 23) June 1932 (1 2). KENNEBEC CO., Mt. Vernon, 10 June 1967 (1 2), Dearborn. PENOB- SCOT CO., Lincoln, 5 July (1 2), 6 July (1 2), 7 July 1935 @ G), 10 July 1935 (1 6, 1 @), 18 July 1937 (1 3); Passadum- keag, | June 1936 (1 ¢), d genitalia slide 342 USNM 46253, 18 June 1938 (1 3); Patten, 26 July (1 2). PISCATAQUIS CO., Che- suncook, 13 July (1 2); Mt. Katahdin, 9 July 1939 (1 2); Sebec Lake, 24 June 1930 (3 6,2 2), 16 June 1923 (1 3). SOMER- SET CO., Pittston Farm, 9 July (1 <). MICHIGAN: CRAWFORD CoO., 5 mi W of Roscommon, 19 May 1938 (2 6), 29 May 1938 (5 3d). MONTGOMERY CO., Atlanta, 31 May 1936 (1 6, 1 @), 31 May 1938 (1 3), McAlpine. OAKLAND CO., Franklin, 24 May 1936 (1 6,1 @), dgen- italia slide USNM 46252, 2 June 1934 (1 G01) 2) 2 une 1932 7e2)s 4 June 1937 (1 3), McAlpine; Royal Oak, 14 May 1915 (1 36), McAlpine. OSCODA CO., Luzerne, 4 June 1944 (1 d), Rawson. NORTH CAR- OLINA: ALLEGHANY CO., 7 mi. N. Sparta, 2,700 ft., 10 June 1974 (1 d, 1 &), Sullivan; New River State Park, Oliver Farm, 2,600 ft., 1-4 June 2000 (2 6, 2 9), Sullivan. ASHE CO., Mt. Jefferson State Park, hairpin turn, 4,150 ft., 1 June 2000 (14 6,7 @), Sullivan; Mt. Jefferson State Park, park offices, 3,080 ft., 2-3 May 2000 (6 6, 1 2), 1 June 2000 (3 3G), Sullivan; Mt. Jefferson State Park, summit, 4,600 ft., 1 June 2000 (18 3,8 2), Sullivan. AVERY CO., Grandfather Mountain, 4,850—5,150 ft., 30-31 May 2000 (1 6, 1 @), Sullivan; Grandfather Mountain, Visitor Ctr, 5,150 ft., 25-27 June 2000 (1 3), Sullivan; Moore Mt. 4,000 ft., 30-31 May 2000 (3 6, 2 @), Sullivan. HAYWOOD CO., Wagon Gap, 4533 ft., 20 May 2001 (10 3), 26-27 June 2001 (6 3d, 5 @), Sullivan; Black Balsam Mt., 5,800 ft., 26-27 June 2001 (5 ¢), Sul- livan; mile post 415, Blue Ridge Pkwy. 4,800 ft., 27 June 2001 (2 6, 2 2), Sulli- van; 155 Mt. Pisgah Rd., 2,900 ft., 20 May 2001 (2 d), 25 June 2001 (2 CG), Sullivan and Deutschman; US 276 below Wagon Gap Rd., 3,776 ft, 23 April 2001 (2 3), 20 May 2001 (1 6,3 @), 26-27 June 2001 (5 3,3 2), Sullivan and Deutschman; Wagon Gap Rd. on US 276, 20 May 2001 (4 6, 5 2), 23 June 2001 (5 3d, 2 2), 26—27 June 2001 (2 @), Sullivan and Deutschman. HENDERSON CO., Bat Cave, 26 May PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1970 (1 &), Covell. SWAIN CO., Chero- kee, Big Cove Road, Great Smoky Moun- tains National Park, 2,062 ft., 21 May 2001 (2 2), Sullivan and Deutschmann, 30 May 2001 (1 &), Sullivan; Thomas Divide trail- head, Great Smoky Mountains National Park, 4,580 ft., 21 May 2001 (4 3d), Sulli- van; Great Smoky Mountains National Park, Big Cove Rd., 2,064 ft., 30 May 2001 (1 2), Sullivan and Deutschman. NORTH DAKOTA: EMMONS CO., 16 mi E Lin- ton, 20 June 1975 (3 3), Lago. NORTH CASS CO., Red River, Fargo, 9 June 1961 (1 3), Poole. NEBRASKA: CHERRY CO., Fort Niobrara, 14 June 1983 (1 ¢), Fergu- son; Hackberry Lake, Valentine NWR, 18 June 1983 (1 6), Ferguson. NEW HAMP- SHIRE: Squam Lake, July 1925 (1 @), Al- len. BELKNAP CO., Center Harbor, (1 2). HILLSBOROUGH CoO., Manchester, (1 2). ROCKINGHAM CO., Hampton, 3 June 1902 (1 3), 4 June 1940 (1 3G), 21 June 1931 (1 2), Shaw. NEW JERSEY: BUR- LINGTON CO., Medford Township, Lake Pine, 7 June 1973 (3 6,2 @), 16 Aug. 1974 (1 2), Rentz. HUNTERDON CO., Woodg- len, 13 June 1971 (1 @), 22 June 1971 (1 ?). OCEAN CO., Lakehurst, (1 ¢), May 1929 (1 36, 1 2), Lenmer, May 1925 (1 @); Wrangle Brook Rd., Lakehurst, 30 May 1956 (1 6, 3 2), 2 2 genitalia ‘slides USNM 46268, 46271, 31 May 1956 (2 6, 3 2), 3 June 1956 (4 6, 2 &), dgenitalia slide USNM 46250, 7 June 1956 (2 3), 8 June 1956 (2 6, 3 2), Ferguson. NEW MEXICO: SAN MIGUEL CO., Las Vegas, (2 3), d genitalia slide USNM 46255. NEW YORK: [no specific locality], (2 d, 3 @), 2 genitalia slide JGF 1756. Centre, 21 May 1877 (1 6d), dgenitalia slide JGF 1755, Hill. CATTARAUGUS CO., Allegany State Park, 10 Sep. 1940 (1 @), Shadle. CHEN- ANGO CO., Greene, 2500 ft., (1 d), June (2 3d), Howard. KINGS CO., Brooklyn, 18 June’ 1902 (1d); 20° June. 1901" 6); d genitalia slide USNM 46265. QUEENS CO., Woodhaven, Long Island, (1 d). SUF- FOLK CO., Amagansett, Long Island, 30 May 1913 (1 @), Englehardt; East, Long VOLUME 105, NUMBER 2 Figs. 20-21. Island, 14 May 1902 (1 2), 22 May 1902 (1 @), 5 June 1900 (2 2), Weeks, 5 June 1901 (1 3), Weeks; Eatons Neck, Long Is- land, 10 June 1971 (1 2), Jackh; Penniquid Barrens, Coram, Long Island, 20 Aug. 1920 (1 2). SULLIVAN CO., Debruce, 9 July 1916 (1 3d). TOMPKINS CO., Ithaca, 22 343 Collecting localities. 20, Elephria festivoides. 21, E. alapallida. May 1965 (2 ¢), Ferguson; McLean Bogs Reserve, 15 June 1957 (1 @), Davis. OHIO: ATHENS CO., Waterloo Township, Sec. 33, at a beaver dam, 23 May 1981 (1 °@), Metzler. LUCAS CO., Holland, 10 May 1936 (1 3), Rawson. PENNSYLVANIA: BEAVER CO., New Brighton, 25 May 344 1902 (1 2), 31 May 1902 (1 2), 1 June 1L902NQ 5S. eS) 2elune TOMS Tes) ale June 1901 (1 3), Merrick. CENTRE CO., Howard, 31 May 1979 (1 2), Haas. LAN- CASTER CO., Lititz, Aug. 1924 (1 @), do genitalia slide JGF 1765, Heiserman. SOUTH DAKOTA: LAWRENCE CO., Cheyenne Crossing, 5,600 ft., 1 July 1997 (1 d), Rahn. PENNINGTON CO., Hill City, 5 July 1964 (1 6, 1 2), Ferguson; Slate Creek, 9 mi NW of Hill City, Black Hills, 6,000 ft., 4 July 1964 (1 6, 2 9), Ferguson; Upper Spring Creek, near Hill City, Black Hills, 27 June 1964 (2 6,5 2), 2 genitalia slide USNM 46246, 28 June 1964 (5 3, 2 2), dgenitalia slide USNM 46245, Ferguson. VIRGINIA: [no specific locality], (1 3d). BATH CO., 2.6 mi E of Warm Springs, 8 June 1956 (1 2), Hevel. FAIRFAX CO., Alexandria (Rose Hill), 24 July 1976 (1 3), Opler. MONTGOMERY CO., Brush Mountain, 14 May 1963 (1 ¢d), 18 May 1963 (1 3), Covell. CITY OF VIR- GINIA BEACH, Seashore State Park, 1—4 June 1975 (3 6), Davis. WISCONSIN: MARATHON CO., Township of Stettin, 4 June 1975 (1 3), 17 June 1975 (1 @), Rahn. WEST VIRGINIA: TUCKER CO., Black- water Falls State Park, 23 June 1968 (1 6, 9 2), 24 June 1968 (4 3, 22 2), d genitalia slide USNM 46262, 2 genitalia slide USNM 46272, Clarke. HARDY CO., Lost River State Park, 29 June 1968 (1 6), d genitalia slide USNM 46626, Clarke; near Lost River State Park, 4—6 July 1980 (1 3d), Miller. MORGAN CO., 4 mi SW Grand Cacapon, 19 Aug. 1989 (1 @), Clarke, McIntyre. Paratypes were deposited in the following institutions: American Museum of Natural History, New York, NY; The Natural History Museum, London, Great Britain; Field Museum of Natural History, Chicago, IL; Los Angeles County Museum of Natural History, Los Angeles, CA; Mis- sissippi State University, Mississippi State, MS. Distribution (Fig. 21).—Most records are from the northeastern U.S. and Nova Sco- tia, Canada. It extends as far south as the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mountains of North Carolina and west to western British Columbia and south to Col- orado and New Mexico. Discussion.—Elaphria alapallida has been masquerading as EF. festivoides in the North American literature (Rockburne and Lafontaine 1976, Covell 1984, Rings et al. 1992). After examination of the types of festivoides and varia and comparing these with similar Florida specimens it was de- termined that the southern species was E. festivoides and the more northern species was E. alapallida. The collection dates (Fig. 19) show that E. alapallida is single brooded. Dates range from the end of April to mid-August across its geographical distribution. The data shown are from the entire geographic range of E. alapallida. These broods are certainly better defined if viewed from a more re- stricted range. For example, the dates in North Carolina are from late April to the end of June, in Connecticut from mid-May to mid-June, in Maine from early June to the end of July, and in Nova Scotia from the end of May to the end of July. Etymology.—The specific epithet, ala- pallida, refers to the pale hindwing with the Latin terms ala referring to the hindwing and pallida reterring to the pale coloration as compared to the darker hindwing of E. festivoides. CONCLUSIONS With the recognition of alapallida the festivoides complex of the genus Elaphria consists of three very similar species. Ac- curate determination of the three species by maculation is extremely difficult without a known series of each species. However, males are easily placed by dissection of the genitalia. Elaphria cornutinis is the only species of the complex with ventral abdom- inal pouches and coremata (Saluke and Po- gue 2000). Males of the remaining two spe- cies can be distinguished by forewing size and the number and size of the cornuti on the inflated vesica. They can also be sepa- rated by the color of the hindwing which is VOLUME 105, NUMBER 2 whitish in alapallida and gray in festivoi- des. The hindwing of cornutinis is inter- mediate but closer to gray. The female of festivoides is distinguished by the longer appendix bursa. Females of alapallida and cornutinis are extremely difficult to sepa- rate on characters of the genitalia. However, the female of alapallida has a_ lighter hindwing color and the female forewing of cornutinis has a more mottled pattern. In North Carolina, the three species are largely allopatric. Elaphria festivoides is limited to xeric, white sand habitats in eastern North Carolina where it occurs with such uncom- mon species as Catocala jair Strecker, Zale declarens (Walker) (Noctuidae) and Heter- ocampa varia Walker (Notodontidae). It is common in the early spring and late fall with stragglers throughout the summer. It overlaps E. cornutinis in the sandhills, where both occur at Weymouth Woods Nat- ural Area in Moore County. There festivoi- des is in the dryer, sandy habitats and cor- nutinis iS 1n mesic woodlands. Elaphria cornutinis 18 Common in mesic woodlands across the piedmont of North Carolina and up to an elevation of about 2,200 feet in the mountains. There it overlaps with E. ala- pallida, but the species appear to be in dif- ferent habitats. Elaphria cornutinis is in the more xeric woodlands where Vaccinium species are common. At altitudes below 2,500 feet E. alapallida is infrequent but becomes abundant from 3,000—5,000 feet throughout the montane forests of the state and has been collected as high as 5,800 feet. Elaphria alapallida is single brooded in the spring while E. cornutinis has mul- tiple broods beginning in April-May with a strong flight and showing. straglers throughout the summer but no strong fall flight. The food plant of E. alapallida was re- ported to be Acer negundo Linnaeus by Rockburne and Lafontaine (1976) but the distribution of that plant species does not coincide with the distribution of alapallida in North Carolina (Radford et al. 1968). It does match the distribution of E. cornutinis but that species does not reach Canada, the origin of the food plant record. The food plants of species of Elaphria in the eastern U.S. are poorly known in spite of the abun- dance of adults. Wagner et al. (1997) reared E. versicolor on algal layers on tree bark. If this feeding strategy occurs in other spe- cies of the genus, it may explain why larvae are seldom located. The genus Elaphria as currently con- ceived, contains 127 species (Poole 1989, Saluke and Pogue 2000, and E. alapallida added here). The genus is largely Neotrop- ical but one species occurs in Europe, one in Africa, one in China, three in India, and 14 in North America. The genus has never been revised nor delineated in any detail al- though Forbes (1954) briefly describes ge- neric characters. Dissections of some 25 species from the Americas indicate that the genus is polyphyletic. The type species is FE. grata (Hiibner), which appears to have rather typical and unspecialized genitalia. The genitalia of the species in the festivoi- des complex are somewhat specialized, par- ticularly the two parallel rows of cornuti in the sinnuate vesica in the male and the short ductus bursae in the female. Perhaps the closest relative to the complex is E. agro- tina which has a similar maculation, ab- dominal pockets and coremata, two short rows of cornuti on a sinnuate vesica and a short sclerotized ductus bursae. Saluke and Pogue (2000) suggested that the presence of abdominal pockets and cor- emata in E. cornutinis could indicate that this species reveals conspecifics chemically by the use of pre-mating pheromones. The geographic range of E. cornutinis largely divides E. festivoides from E. alapallida. The distribution of abdominal pockets and coremata 1s sporatic in Elaphria species and may represent a pattern of expression rather than absolute genetic gain or loss of a com- plex character (Poole 1995). It may be in- volved in rapid speciation, perhaps even sympatric speciation. The presence or ab- sence of such a pheromone system is also found in other very closely related species 346 pairs of noctuids (Euagrotis, Lacinipolia and probably others). One can imagine al- leles at very low frequencies which allow the expression in males of the pheromone apparatus or the reception of the pheromone in females. In time, the occurrence of the alleles in individuals of both sexes may oc- cur at a single locality producing offspring which could utilize the pheromone system as an isolating mechnism, and over time al- low for habitat specialization to avoid direct competition in the larval stages. The festi- voides complex may offer a convenient model for the study of rapid speciation and pheromone dependency. ACKNOWLEDGMENTS For critically reviewing a draft of this mauscript, we thank Terry L. Erwin, De- partment of Systematic Biology, Smithson- ian Institution, Washington, DC, and N. J. Vandenberg and D. R. Smith, Systematic Entomology Laboratory, U.S.D.A., Wash- ington, DC. We are grateful to James Ad- ams, Dalton, GA, R. L. Brown, Mississippi State University, Mississippi State, MS, and Bryant Mather, Clinton, MS, Charles V. Covell, Jr, University of Louisville, Lou- isville, KY, Steve Hall, North Carolina Her- itage Program, Raleigh, NC, Don DeFoe and Keith Langdon, Great Smoky Moun- tains National Park, Gatlinburg, TN, for al- lowing us to borrow, examine, and dissect specimens from their collections. For loan- ing us the types of E. festivoides and E. varia, we thank Ian J. Kitching of The Nat- ural History Museum, London, United Kingdom. For permission to collect Lepi- doptera in restricted areas we thank the fol- lowing: Vic French and Randall C. Wilson, North Carolina Wildlife Resources Com- mission; A. Este Stiefel, The Nature Con- servancy; Steve Miller, Grandfather Moun- tain Preserve; Keith Langdon, Great Smok- ey Mountain National Park; Chris Ulrey, Blue Ridge Parkway National Park; Erich L. Hoffman, Ft. Bragg Military Reserva- tion; John Townsend, Camp Lejune Mili- tary Reservation; Scott Hartley, Jody Mer- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ritt, Tommy Wagoner, and Jay Wild, North Carolina Department of Environmental Re- sources. LITERATURE CITED Barnes, W. and J. McDunnough. 1917. Checklist of the Lepidoptera of Boreal America. Herald Press, Decatur, Illinois. 392 pp. Boisduval, J. B. A. D. de and A. Guenée. 1852. His- toire Naturelle des Insectes. Species Général des Lépidopteres. Tome Cinquiéme. Noctuélites. Tome I. Roret, Paris, 407 pp. Butler, A. G. 1891. Notes on the synonymy of Noctuid moths. The Entomologist 24: 237-242. Covell, C. V., Jr. 1984. A Field Guide to the Moths of Eastern North America. Houghton Mifflin Co., Boston. xv, 496 pp., 64 plates. Draudt, M. 1926. Family Noctuidae, pp. 15-412. In Seitz, A. Die Gross-Schmetterlinge der Erde. Ab- teilung I]. Amerikanischen Faunengebietes. Band 7. Eulenartige Nachtfalter. Stuttgart. Alfred Ker- nen. 508 pp., 96 plates. Dyar, H. G. 1902 [1903]. A list of North American Lepidoptera and key to the literature of this order of insects. Bulletin of the United States National Museum, No. 52, xix + 723 pp. Forbes, W. T. M. 1954. Lepidoptera of New York and Neighboring States, Part II] Noctuidae. Memoir 329. Cornell University Agricultural Experiment Station, Ithaca, New York, 433 pp. Franclemont, J. G. and E. L. Todd. 1983. Noctuidae, pp. 120-159. In Hodges, R. W. et al., eds. Check List of the Lepidoptera of America North of Mex- ico. E.W. Classey Ltd. and The Wedge Entomo- logical Research Foundation, London. Hampson, G. FE 1909. Catalogue of the Lepidoptera Phalaenae in the British Museum, Vol. 8. Taylor and Francis, London. 583 pp, xiv, plates 123-136. Kimball, C. P. 1965. The Lepidoptera of Florida an annotated checklist. Division of Plant Industry, State of Florida. Department of Agriculture, Gainesville, Florida. 363 pp., xxvi plates. Poole, R. W. 1989. Lepidopterorum Catalogus (New Series), Fascicle 118. Noctuidae. E.J. Brill and Flora and Fauna Publications, Leiden. 1,313 pp. . 1995. Noctuioidea, Noctuidae (part). Jn Dom- inick, R. B., et al. The Moths of America North of Mexico, fasc. 26.1, 249 pp. Poole, R. W. and P. Gentili, eds. 1996. Nomina Insecta Nearctica. A Check List of the Insects of North America, Vol. 3. Diptera, Lepidoptera, Siphonap- tera. Entomological Information Services, Rock- ville, Maryland. 1,143 pp. Radford, A. E., H. E. Ahles, and C. R. Bell. 1968. Manual of the Vascular Flora of The Carolinas. University of North Carolina Press, Chapel Hill, 1,183 pp. VOLUME 105, NUMBER 2 Rings, R. W., E. H. Metzler, F J. Arnold, and D. H. Harris. 1992. The Owlet Moths of Ohio Order Lepidoptera Family Noctuidae. Ohio Biological Survey Bulletin. New Series Vol. 9 No. 2. Vi + 219 pp, 16 pls. Rockburne, E. W. and J. D. Lafontaine. 1976. The Cut- worm Moths of Ontario and Quebec. Research Branch Canada Department of Agriculture, Pub- lication 1593. 164 pp. Saluke, S. V. and M. G. Pogue. 2000. Resolution of the Elaphria festivoides (Guenée) species complex (Lepidoptera: Noctuidae). Proceedings of the En- tomological Society of Washington 102: 233-270. Smith, J. B. 1889. Contributions toward a monograph 347 of the Noctuidae of temperate North America: Ge- nus Oligia Hiibner. Entomologica Americana 8: 143-152. . 1891. List of Lepidoptera of Boreal America. American Entomological Society, Philadelphia, Pennsylvania. 124 pp. Wagner, D. L., V. Giles, R. C. Reardon, and M. L. McManus. 1997. Caterpillars of Eastern Forests. United States Department of Agriculture, Forest Service, FHTET 96-34, 113 pp. Walker, FE 1857 [1858]. List of specimens of Lepidop- terous Insects in the collection of the British Mu- seum, Vol. 12. Edward Newman, London, pp. 764-982. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 348-351 THE DISCOVERY OF THE GENUS ECNOMIOS MASON (HYMENOPTERA: BRACONIDAE) IN CHINA, WITH DESCRIPTION OF A NEW SPECIES XUEXIN CHEN AND J. B. WHITFIELD (XC) Institute of Applied Entomology, Zhejiang University, Hangzhou 310029, China & Department of Entomology, University of Hlinois, 320 Morrill Hall, 505 S. Goodwin Ave., Urbana, IL 61801, U.S.A.; (JBW) Department of Entomology, University of Illinois, 320 Morrill Hall, 505 S. Goodwin Ave., Urbana, IL 61801, U.S.A. (e-mail: jwhitfie@ life.uiuc.edu) Abstract.—A new species of braconid wasp, Ecnomios flavus, is described. It repre- sents the first record of the genus Ecnomios Mason as well as the subfamily Ecnomiinae in China. Key Words: The rare genus Ecnomios was first de- scribed by Mason (1979) from Papua New Guinea with the type species E. papuensis Mason 1979. Thirteen years later a second species of this genus was described from Australia by Austin and Wharton (1992) and, in 1993, another two species were re- ported from Vietnam by Belokobylskij (1993). Achterberg (1995) provided a key to the species of Ecnomios with descrip- tions of an additional three species from In- donesia. Currently seven species of the ge- nus have been described with an obvious Indo-Australian distribution before this study. When the first author visited the Shang- hai Institute of Entomology, Academia Sin- ica in 2000, a female specimen of Ecnomios was encountered, and further examination proved it to be a new species. It represents the first record of the genus Ecnomios Ma- son as well as of the subfamily Ecnomiinae in China. The subfamily Ecnomiinae was erected by Achterberg (1985) to hold the aberrant genus Ecnomios. A second genus, Korec- nomios Park and Achterberg 1994, of this Ecnomios, new species, Braconidae, China subfamily was reported from Korea, with only the type species known (Park and Achterberg 1994). The biology of the sub- family is unknown. Mason (1979) originally placed his new genus Ecnomios in the tribe Orgilini Foers- ter 1862 (now subfamily Orgilinae), al- though more recent research suggests it may belong to the ““microgastroid lineage” (Quicke and Achterberg 1990). The most prominent shared character with this line- age is the large plical cell of the hind wing, with a distinct cleft distally. A large plical cell appears to be plesiomorphic within the Hymenoptera (Quicke and van Achterberg 1990), but in phylogenetic analysis within Braconidae (Quicke and van Achterberg 1990, Wharton et al. 1992), its presence ap- pears as a reversal. Other possible synapo- morphies with the microgastroid lineage are the shape of the first discal cell of fore wing, the comparatively long fore spur, the absence of the lateral carina of the mesos- cutum, and the position of vein SRI of the fore wing. The vertical position of vein fr, the oblique vein 1-CU1, the comparatively slender first discal cell and the reduced vein VOLUME 105, NUMBER 2 2-1A of the fore wing suggest some rela- tionship to the subfamily Dirrhopinae with- in the microgastroid lineage. Autoapomor- phies of the Ecnomiinae within the micro- gastroid lineage are the anteriorly wide pronotum, the short labial palp (in compar- ison with the maxillary palp), and the sin- uate vein 2-M of hind wing. It is to be hoped that suitable material of this group will become available to allow analysis of DNA sequence data, to test these possible relationships further in the context of ex- panding understanding of braconid, espe- cially microgastroid, phylogeny (Whitfield 1997, Belshaw et al. 1998, Dowton and Austin 1998, Whitfield 2002). The morphological (including wing vein) terminology used in this paper follows Ach- terberg (1993). Ecnomios flavus Chen and Whitfield, new species (Figs. 1—4) Female.—Body length 2.7 mm, fore wing length 2.4 mm. Color: Brownish yellow, metasoma after first tergite light brown; palp pale yellow; antenna brownish yellow, apical half dark- er; legs yellowish; pterostigma brown, pale basally; veins unpigmented. Head: Antennal segments 24, length of first flagellar segment 1.5 times second fla- gellar segment, first, second and penulti- mate flagellar segments 2.3, 1.7 and 1.6 times their width, respectively, apical seg- ment with a spine; occipital carina com- plete; length of maxillary palp 0.7 times height of head; length of eye in dorsal view 1.7 times temple; temple strongly and rounded behind eyes, smooth; OOL: OD: POL = 10:10:11; frons smooth and slightly concave medially; face with a wide medio- longitudinal ridge, nearly smooth with long setae, its width 1.7 times its height, as long as height of eye; clypeus distinctly convex, smooth, its width about 2 times its height; length of malar space 1.7 times basal width of mandible. 349 Mesosoma: Length of mesosoma 2.0 times its height; side of pronotum coarsely crenulate anteriorly and posteriorly, rest al- most smooth; precoxal sulcus narrow, cren- ulate-rugose, anteriorly absent, rest of me- sopleuron smooth, shiny; metapleuron coarsely rugose; notauli shallow but dis- tinct, rugose, joining in a broad, distinct and rugose area posteriorly before scutellar su- ture, with a medio-longitudinal carina in this rugose area; mesoscutum other than no- tauli largely smooth with setae; scutellar su- ture with 5 flat and smooth, glabrous, without lateral carina; propodeum completely rugose with a dis- tinct transverse carina in posterior half, forming a pair of latero-posterior tubercles. Wings: Fore wing: length of pterosigma 1.2 times vein I-R1, 2.5 times its width; r: 3-SR+SR1: 2-SR: m-cu = 14:73:18:17; 2-CU1 much lower than M+CU1; SRI sin- uate; 1-CU1: 2-CU1 = 13:20, 1-CUlI:cu-a = 13:6. Hind wing: M+CU: I-M: Irm = carinae: scutellum 36:18:15; base of SR unsclerotised; 2- SC+R much shorter than Im, nearly quadrate. Legs: Hind coxa with some distinct ca- rinae anteriorly, rest largely smooth; all tar- sal claws simple and rather slender; length of femur and basitarsus of hind leg 4.3 and 5.6 times their width, respectively; length of hind tibia 1.25 times hind tarsus; length of hind tibial spurs 0.57 and 0.39 times hind basitarsus; basitarus and second tarsal seg- ment with a distinct ventral carina. Metasoma: Length of first tergite 0.8 times its apical width, distinctly and line- arly widened apically, its surface coarsely rugose, weaker near apical margin where is almost smooth, its dorsal carina absent; sec- ond and following tergites smooth; second suture absent; length of ovipositor sheath 0.8 times hind basitarsus, 0.08 times fore wing; Ovipositor sheath glabrous. Male.—Unknown. Material examined.—Holotype, 2, Chi- na: Yunnan, Hekou, Xiaonanxi, 1956.vi.7, Huang Keren, kept in Shanghai Institute of Entomology, Academia Sinica, Shanghai. 350 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Ecnomios flavus, holotype. 1, Wings. 2, Head, dorsal view. 3, Head, frontal view. 4, Mesoscutum, dorsal view. VOLUME 105, NUMBER 2 Etymology.—The specific name ‘‘fla- vus’ refers to the body color of the new species being almost completely yellowish. Notes.—This new species runs to couplet 6 in the key of Achterberg (1995), but can be separated from the most closely similar species, Ecnomios caophongi Belokobylskij 1993 in having the vein 1-CU1 of fore wing much longer, almost 2.2 times vein cu-a (1.2 times in E. caophongi); vein 2-SC+R of hind wing shorter, nearly quadrate (dis- tinctly longitudinal in E. caophongi); notau- li more distinct with a broad joining area posteriorly (notauli very shallow and _ ru- gulose, joining in a small and weakly reti- culo-puctate area posteriorly in E. caophon- gi), face smooth with a broad medio-lon- gitudinal ridge (this area much narrower in E. caophongi), and hind femur more slen- der, 4.3 times as long as wide (3.5 times in E. caophongi). ACKNOWLEDGMENTS The first author thanks Dr. Haisheng Ying for his hospitality during his stay at the Shanghai Institute of Entomology, Shanghai, and for the loan of the specimen. We thank C. van Achterberg (Leiden, the Netherlands) and S. A. Belokobylskij (St. Petersburg, Russia) for reviewing the first draft. The project was partly supported by the National Scientific Foundation of China (NSFC number: 39970099) to the first au- thor. LITERATURE CITED Achterberg, C. van. 1985. Notes on Braconidae V—VI. V. The systematic position of the genera Ecnomios Mason and Pselaphanus Szepligeti (Hymenop- tera). Zoologische Mededelingen, Leiden 59: 341— 348. . 1993. Hlustrated key to the subfamilies of the 351 Braconidae (Hymenoptera: Ichneumonoidea). Zoologische Vergandelingen Leiden 283: 1-189. . 1995. New taxa of the subfamilies Betylob- raconinae, Cenocoeliinae, Economiinae, Homolo- binae, and Sigalphinae (Hymenoptera: Braconi- dae) from East Indonesia. Zoologische Medede- lingen, Leiden 69: 307-328. Austin, A. D. and R. A. Wharton. 1992. New records of subfamilies, tribes and genera of Braconidae (Insecta: Hymenoptera) from Australia, with de- scription of seven new species. Transactions of the Royal Society of South Australia 116: 41-65 Belokobyskij, S. A. 1993. New taxonomic data on the braconid fauna (Hymenoptera: Braconidae) of Vi- etnam. Russian Entomological Journal 2(2): 37— 67. Belshaw, R., M. Fitton, E. Herniou, C. Gimeno, and D. L. J. Quicke. 1998. A phylogenetic reconstruc- tion of the Ichneumonoidea (Hymenoptera) based on the D2 variable region of 28S ribosomal RNA. Systematic Entomology 23: 109-123. Dowton, M. and A. D. Austin. 1998. Phylogenetic re- lationships among the microgastroid wasps (Hy- menoptera: Braconidae): combined analysis of 16S and 28S rDNA genes. Molecular Phylogeny and Evolution 10: 354-366. Mason, W. R. M. 1979. A new genus and species of Orgilint (Hymenoptera: Braconidae) from New Guinea. Proceedings of Entomological Society of Washington 81: 640-644. Park, J. S. and C. van Achterberg. 1994. A new genus of the subfamily Ecnomiinae van Achterberg (Hy- menoptera: Braconidae) from Korea. Zoologische Mededelingen, Leiden 68: 49-54. Quicke, D. L. J. and C. van Achterberg. 1990. Phy- logeny of the subfamilies of the family Braconi- dae (Hymenoptera: Ichneumonoidea). Zoologis- che Vergandelingen, Leiden 258: 1—95. Wharton, R. A., S. R. Shaw, M. J. Sharkey, D. B. Wahl, J. B. Woolley, J. B. Whitfield, P.M. Marsh, and W. Johnson. 1992. Phylogeny of the subfam- ilies of the family Braconidae (Hymenoptera: Ich- neumonoidea): a reassessment. Cladistics 8: 199— 235). Whitfield, J. B. 1997. Molecular and morphological data suggest a single origin of the polydnaviruses among braconid wasps. Naturwissenschaften 84: 502-507. . 2002. Estimating the age of the polydnavirus/ braconid wasp symbiosis. Proceedings of the Na- tional Academy of Sciences of the USA 99: 7508— (pyler PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 352-361 FIVE NEW SPECIES OF XANTHINA ALDRICH (DIPTERA: DOLICHOPODIDAE) FROM MEXICO AND CENTRAL AMERICA HAROLD ROBINSON Botany, Department of Systematic Biology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0166, U.S.A. Abstract.—Five new species of Xanthina are described from Mexico and Costa Rica, X. squamifera, X. flagellifera, X. schildi, X. turrialbae, and X. attenuata, showing various distinctive antennal or palpal forms. The large palpal scales of X. squamifera and the flagelliform palpal setae in X. flagellifera are particularly noted. Key Words: Xanthina Aldrich is an American genus credited in the Neotropical Catalogue (Rob- inson 1970) with four species, X. flavus (Al- drich) and X. plumicauda Aldrich from the Lesser Antilles, and X. nigromaculata Van Duzee and X. subcurva Van Duzee from Panama. Four species were described by Robinson (1975) the Lesser Antilles, X. X. domini- censis, X. persetosa, and X. rubromargin- ata. Another supposedly undescribed spe- cies was mentioned in Robinson (1975) from Puerto Rico, that has been reexamined and seems to be a minor variant of X. per- setosa. At the time of the Dominica treat- ment, at additional remarkable species already was collected from Mexico, and representatives of four other Central from Dominica in acuticornis, least one American species, collected from the Tur- rialba area of Costa Rica in 1922 or ca. 1922 by Pablo Schild, had been sorted out of the Melander collections at the National Museum of Natural History, Smithsonian Institution (USNM). It is these Mexican and Central American species that are the topic of the present paper, especially the remark- able Mexican species with large scales on its palpi. Like the more temperate Achalcus Loew, Xanthina, new species, Dolichopodidae, Mexico, Costa Rica, palpi, ESEM Xanthina is distinctive in its yellowish color with mostly black bristles, and in having the scape of the antenna bare above, the posterior slope of the mesoscutum some- what flattened, and the wings with the vein M usually scarcely deflected at the cross- vein dm-cu, and vein R,,; often becoming subparallel with M more distally. However, the femora of Xanthina lack true antero- dorsal preapical bristles. In the Catalogue, Robinson (1970) states,““The genus seems very close to Achalcus except for the lack of true preapical bristles on the femora. The palpi are often highly developed and or- nate.’ The palpus character is especially true of some species described below, but it seems that the labella are also often in- terestingly modified. In one new species, X. turrialbae, modified labella occur in both sexes. The new species with scales on the palpi, Xanthina squamifera, and a related new species with narrow trichomes on the palpi, X. flagellifera, share a distinctive short ab- domen with the fifth tergite hiding the hy- popygium. The two species have the first flagellomere deeply notched at the apex, unlike most NXanthina; a smaller less dis- tinct notch is seen in one of the other new VOLUME 105, NUMBER 2 species, X. schildi. Establishment of a new subgenus for the two most discordant spe- cies has been considered, but decided against at this time. For illustration of the new species, the new Philips XL30 ESEM Scope of the Na- tional Museum of Natural History, that al- lows imaging without metallic coating, has been used. This has allowed highly detailed imaging that includes some substructure of the scales of Xanthina squamifera. In the course of the study, material of two probable additional species has been seen, including one from North Carolina. Such material is too limited or too damaged for description at this time, however. The new species described here are based on limited material, each with only one male and often with no female. As a result, no effort has been made to dissect out hidden genital capsules, and some legs and wings were not available for description. Still, each of the species described below has at least one unquestionably distinctive feature by which the males, and in one case the females, can be easily identified. Xanthina squamifera H. Robinson, new species (Fig. 1) Male.—Length 2.2 mm; wing 2.7 by 0.8 mm. Setae mostly black. Eyes nearly contiguous at lower Y, of face, facial triangles brown pollinose with pale reflections; front dark brown, scarcely dulled with sparse pollen. Palpus (Fig. 1B) narrowly ovate, acute, at midlength with numerous large stipitate white scales with dark bases, with flattened whitish setae nearer tips. Labella tonglike, pale yellow; long-acute at front, narrowed to apical peg. Antenna dark yellow, first flagellomere darker distally, oblong, with broad deep apical notch to 7 of length (Fig. 1A), longer lobe below notch; arista from middle of notch. Thorax yellow; 6 or 7 pairs of acrosti- chals; 6 pairs of distinct dorsocentrals. Legs yellow. Fore coxa with pale hairs 353 anteriorly, with small black bristles along apical margin; femora not distorted, without highly modified setae; middle femur with slightly longer posteroventral hairs in distal ¥Y.. Tibia IT with anterodorsals near %, 4, posterodorsal near ¥, 3 apicals; tibia III with anterodorsals near ¥,, ¥,, %, postero- dorsals at %, %, 7%, ventrals at 7, ¥, and 3 apicals. Lengths of tibiae and tarsomeres as follows: tibia I: 0.6 mm, tarsi: 0.47, 0.22, 0.16, 0.08, 0.08 mm; tibia II: 1.11 mm, tar- si: 0.36, 0.28, 0.19, 0.14, 0.11 mm; tibia II: 1.55 mm, tarsi: 0.22, 0.30, 0.25, 0.14, 0.11 mm. Wing narrowly oval, slightly fuscous. Vein M scarcely bent at crossvein dm-cu; vein dm-cu nearly as long as CuA,; tip of CuA, ending slightly before margin. Knob of haltere pale brownish. Abdomen brown on tergites, yellow be- low. Tergites 1—5 exposed, with black bris- tles, longer near margins. Genital capsule mostly hidden in tergite 5, brown, rounded apex with small black bristles; cercus pale, small, ovate to subquadrate. Female.—Unknown. Holotype.—d. Mexico: Oaxaca, above Valle Nacional, 16 May 1963, Robinson (USNM). Remarks.—Males of Xanthina squami- fera and the following species are unusual for the genus by the deep apical notch in each first flagellomere, the elaborated tri- chomes of each palpus, only five visible ter- gites of the abdomen, and the short cerci of their genital capsules. In neither case has any effort been made to dissect the genital capsules of the single males of each species. Perhaps more material will provide struc- tural details that will justify a separate sub- generic status for the two species. The present species has been noted by the author since the time of its collection because of the large scales on the palpi, but the description has awaited a fitting method of illustration. The new Philips XL30 ESEM Scope of the Museum has proven perfect, requiring neither coating with metal nor notable moistening of the specimens. 354 PROCEEDINGS OF THE Fig. 1. of proboscis showing acute palpi with broad and narrow scales, and showing prolonged, tong-like tips of labella. ENTOMOLOGICAL SOCIETY OF WASHINGTON Xanthina squamifera. A, First flagellomeres of antennae with deep apical sinuses. B, Frontal view C, Enlarged palpus with broad scales. D, Palpal scale showing some substructure. E, Edge of palpal scale with structural details. Scales or scalelike structures occur in some other Dolichopodidae such as the Sciapo- dinae, along the costa of the wing or along tibiae or basitarsi. The scales of the present species, however, are the largest seen in the family (Figs. |B— E). In the SEM scans the scales show some transparency, and high magnification shows some of the reticulate substructure. Such structure is slightly rem- iniscent of that in the scales of Lepidoptera, but it is not nearly as intricate. The flattened bristles on the more distal parts of the palpi are evidently scales that are scarcely broad- ened. The labella of the proboscis also seem rather odd in their tonglike form and their peglike tips. Xanthina flagellifera H. Robinson, new species (Fig. 2) Male.—Length ca. 2 mm; wings broken near base. Setae mostly black. VOLUME 105, NUMBER 2 » | Xanthina flagellifera. A, Inside view of first flagellomere of right antenna showing deep apical sinus 30 nm pum Fig. 2. B, Palpi and labella showing flagelliform setae of palpi and acuminate tips of labella. C, Enlarged nearly glabrous distal half of one labellum showing acuminate tip. D, Inside view of first flagellomere of left antenna showing apical sinus. E, Enlarged minute setulae of palpus showing truncated tips. Eyes nearly contiguous at lower /, of face, facial triangles brownish pollinose; front dark brownish, not or scarcely dulled. Palpus (Fig. 2B) brownish, triangular with oblique tip, with many long bristles having flagelliform tips, with many small peglike hairs; labella (Figs. 2B, C) pale yellow, very thin, nearly with acuminate tip. Antenna yellowish, first fla- gellomere slightly darker at tips, oblong, apical notch to Y, of length (Figs. 2A, D), lower lobe longer. Thorax yellow; 6 or 7 pairs of acrosti- chals, 6 pairs of distinct dorsocentrals. Legs yellow. Fore coxa with pale hairs glabrous, slender anteriorly, with small black bristles along apical margin. or unusual bristles. Femora without distortions Tibia I with anterodor- sals at ¥%, ¥5; posterodorsal at ¥;, apicals 3; tibia III with anterodorsals at ¥,, /, %, pos- terodorsals at 4%, %, ¥%,, ventral near %, %, ca. 4 erect ventral hairs in 2nd Y,, apicals 3. Lengths of tibia and tarsomeres as follows: tibia [2°0:6.mmy» ‘tarsi: 10.38, 0:2, 0.16; 0:1, 0.1 mm; tibia II: 0.88 mm, tarsi: 0.36, 0.2, 0:16, (ON12, (0:12 mm: tibia Ti: 1:28 mm, 5; 0:24, 0.22;0:16; 0.12 mm. Wings slightly fuscous, broken near base. tarsi: 0. Knob of haltere yellow. Abdomen brown on tergites, yellow be- low. Tergites 1—5 exposed, with black bris- tles, mostly hidden by tergite 5, drical, ish, thin, Female.—Length ca. 3 longer near margins. Genital capsule brown, cylin- with short black hairs; cercus whit- short, ovate, with black hairs. mm; wings 2.8 mm long, ca. | mm wide. First flagellomere ovate, not notched, arista subapical on outer surface. Palpus brownish-yellow, with black hairs; labella plain, without pointed tip. Tibia I] with anterodorsals at /,, ¥;, pos- terodorsal at ’,, apicals 3; tibia III antero- 356 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. two sizes of setae. D, Abdomen with genital capsule and cerci, showing comb of blunt setae of hind margin of Xanthina schildi. Sth sternite. dorsals at Y,, %, ¥,, posterodorsals at ¥;, ¥,, apicals 3. Abdominal tergites more yellowish on sides. Wing slightly fuscous; vein M scarcely bent at crossvein CuA; R,,; curving back to become subparallel with vein M. Holotype.—d. Costa Rica: Cartago. La Suiza de Turrialba, Aug 1922?, Pablo Schild (USNM, from Melander Collection). Paratypes; 2. Costa Rica: Higuito, San Ma- teo, s.d., 1922?, Schild (USNM); 2, Car- tago, Suiza de Turrialba, July 1922?, Schild (USNM). Remarks.—Xanthina flagellifera shares with X. mostly enclosed in the 5th tergite, the short squamifera the genital capsule cerci, and the deeply notched first flagello- mere of the antenna, and the two are evi- dently They unique forms of trichomes on their palpi. In closely related. also share A, First flagellomere of antenna showing small apical sinus. B, C, Palpi showing the present case (Fig. 2B) the trichomes are slender flagellate bristles whose ends reach to the margin of the labella. At the highest magnification the flagellate tips show mi- nute appendages, a detail recalling the mi- nute substructure of the scales in X. squam- ifera. The identity of the females is based on similar chaetotaxy of the legs and the pale knobs of the halteres, but the females are somewhat larger with larger bristles on the tibiae. The females lack the pointed tips of the labella seen in the male. Xanthina schildi H. Robinson, new species (Fig. 3) Male.—Length ca. 1.5 mm; wing ca. 1.6 mm long, torn behind R, ;. Setae mostly black. Eyes contiguous in lower /;, facial triangles brown pollinose, lower triangle al- VOLUME 105, NUMBER 2 most absent; front blackish, not dulled. Pal- pus (Figs. 3B, C) brownish, small, with black bristles and minute hairs; labella of proboscis brownish, small. Antenna yellow- ish, darker distally; first flagellomere ovate, subacute, with easily overlooked small api- cal notch bearing arista. Thorax brownish above, yellowish on sides; 6 or 7 pairs of acrostichals, 6 pairs of distinct dorsocentrals. Legs yellow. Fore coxa with mostly pale hairs, with | black bristle on distal margin. Fore and middle femora plain, hind femur with ventral row of more erect black bris- tles, longest near distal Y, of femur, , as long as width of femur. Tibia IT with anter- odorsals at ¥;, 5, no posterodorsals, apicals 3; tibia II] with small anterodorsal at /,, larger at %, ¥,; posterodorsal near /,; apicals 3. Lengths of tibiae and tarsomeres as fol- lows: tibia I: 0.5 mm, tarsi: 0.3, 0.14, 0.08, 0.06, 0.06 mm; tibia II: 0.65 mm, tarsi: 0.28, 0.14, 0.1, 0.07, 0.08 mm; tibia HI: 0.9 mm, tarsi: 0.15, 0.2, 0.14, 0.09, 0.09 mm. Wing slightly fuscous, torn, missing be- hind R, ;. Knob of haltere brown. Abdomen with tergites brown, sternites yellow. Tergites 1—6 exposed, bristles dark with pale reflections. Genital capsule (Fig. 3D) pale brownish, facing a comb of stiff, curved, blunt bristles apparently borne on a preceding sternite; cercus as long as abdo- men, filiform. Female.—Unknown. Holotype.—d, Costa Rica: Cartago. La Suiza de Turrialba, July 1922?, Pablo Schild (USNM) from the Melander Collec- tion. Remarks.—Xanthina schildi is more like West Indian members of the genus, with six tergites exposed in the abdomen, a more ex- posed genital capsule, and long cerci. The first flagellomere of the antenna also looks more like the typical West Indian type that has no apical notch, but the present species does have a small notch (Fig. 3A). A comb of stiff sternal bristles facing the genital capsule, of the type seen in X. schildi, has thus far not been seen in any other member 357 of the genus. The cerci are the longest and narrowest thus far seen in the genus. Xanthina turrialbae H. Robinson, new species (Figs. 4—5) Male.—Length 2.3 mm; wing 2.65 by 1.0 mm. Setae mostly black. Eyes nearly contiguous in lower Y, to 1; of face, facial triangles yellowish pollinose; front brown, scarcely dulled with pollen. Palpus (Fig. 4D) brownish, oval, with black bristles; labella with slender attenuate tips. Antenna with first flagellomere brownish yellow, oblong, slightly emarginate at tip (Figs. 4A, B); arista nearly twice as long as face, at lower magnification appearing gla- brous throughout, thicker in basal Y, with cover of minute hairs (Fig. 4C), becoming very slender distally with sparse minute fimbriae (Fig. 4B). Thorax yellow; 6 or 7 pairs of acrosti- chals, 6 pairs of dorsocentrals. Legs yellow. Fore coxa with black hairs anteriorly, with black setae along distal margin. Femora without distortions or un- usual bristles. Tibia I with anterodorsal at Y.; tibia Il broken; tibia III with bristles strong, anterodorsals at 4, 4, ¥%; postero- dorsals at ca. ¥;, 4%, ¥,, smaller ventrals at V,, 73; few erect posteroventral hairs in basal ¥,—Y,; apicals 3. Lengths of tibiae and tar- someres as follows: tibia I: 0.83 mm; tarsi: 0.4, 0.23, 0.2, 0.13, 0.12 mm; tibia II and tarsi missing; tibia HI: 1.57 mm; tarsi: 0.27, 0.37 mm, other tarsi broken. Wing slightly fuscous; vein M scarcely bent at dm-cu, R,; bent distally and be- coming subparallel with M, dm-cu as long as last of CuA,, CuA, fading before margin. Knob of haltere. brownish-yellow. Abdomen brown on tergites, yellow on sternites; five tergites exposed, with mar- ginal bristles long, black, others black with yellowish reflections. Genital capsule most- ly hidden by tergite 5, brownish with small black hairs; cercus short. Female.—Length ca. 2.7 mm; wing 3 by | mm. 358 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. flagellomere of antennae. C, Tip of first flagellomere showing slight sinus and base of minutely haired arista. Xanthina turrialbae, male. A, View of head showing antennae, face, and base of proboscis. B, First D, Proboscis with right palpus, part of laterally directed left palpus, and pointed tips of labella. E, Arista showing nearly hairless appearance. Eyes contiguous. Palpus smaller but most- ly as in male (Fig. 5A); labella pointed as in male. First flagellomere shorter than high, slightly emarginate at apex; arista about 1.5 times as long as face, hairy (Fig. 5B). Tibia I with no dorsal; tibia II with an- terodorsals at /,, /,; posterodorsal at ¥.; tibia III with bristles large, anterodorsals at 4, 4;, 7;, apicals 3. Lengths of tibiae and tarsomeres as follows: tibia I: 0.77 mm; tarsi: 0.37, 0.23, 0.2, 0.13, 0.17 mm; tibia I: 1.1 mm; tarsi: 0.43, 0.27, 0.2, 0.15, 0.1 mm; tibia HI: 1.57 mm; bas- itarsus: 0.27 mm, other tarsi broken. Holotype.—d, Costa Rica: Cartago, La Suiza, Apr 1922, Schild (USNM). Allotype 2. Costa Rica: Cartago, La Suiza de Tur- ,, 43 posterodorsals at Y, rialba, Aug 1922, Schild (USNM). Both from the Melander Collection. Remarks.—Xanthina turrialbae differs generally by its large size, the larger bristles on its hind tibiae, and the black rather than pale hairs on the front surfaces of the fore coxae. Of more particular interest are the nearly bare aristae of the male antennae and the pointed labella of both the male and the female probosci. The species has the abdo- men with only five visible abdominal tergites, the nearly hidden genital capsule, and the short cerci of the group of species including X. squamifera and X. flagellifera, but the first flagellomere of the antenna does not have the large apical notch and the palpi do not have strikingly distinctive setae or scales like those found in the two species described above. Xanthina attenuata H. Robinson, new species (Fig. 6) Male.—Length 1.8 mm; wings broken. Setae mostly black. VOLUME 105, NUMBER 2 Fig. 5. Xanthina turrialbae female. A, Proboscis showing short palpi and pointed labella. B, Antennae showing reduced first flagellomeres with small apical sinuses and pubescent aristae. Eyes contiguous in lower , of face; fa- cial triangles mostly pale brownish _ polli- nose; front blackish brown scarcely dulled with pale brownish pollen. Palpus very thin, pale, broadly obdeltoid (Fig. 6B), with scattered pale hairs; labella broad, thin, pale, obtusely pointed with minute apiculus (Fig. 6B). Antenna mostly yellowish; first 360 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON a 25 pm ia B Fig. 6. Xanthina attenuata. A, View of head showing triangular first flagellomeres of antennae with single acuminate tips. B, Proboscis with face view of right palpus in center and edge view of left palpus to right, labella showing short-apiculate tips. VOLUME 105, NUMBER 2 flagellomere darker distally, with obtuse an- gle above near basal ), (Fig. 6A), with long, slender, hairy, attenuate tip below; arista from distal /, of upper margin, slightly lon- ger than first flagellomere, hairy. Thorax yellow, 6 or 7 pairs of acrosti- chals, 6 pairs of dorsocentrals. Legs yellow. Fore coxa with pale hairs anteriorly, with black bristles distally; fe- mur I plain; femur IT with row of erect, fine hairs ventrally, some as long as ); width of femur, small black apical on midanterior surface; hind femora missing. Tibia I with anterodorsals at ,, ¥%, small posterodorsal at ¥;, 3 apicals. Lengths of tibiae and tar- someres as follows: tibia I: 0.5 mm, tarsi: 0.33, 0.14, 0.1, 0.09, 0.063 mm; tibia II: 0.8 mm); basitarsus: 0.3 mm, other tarsi broken. Hind tibiae and tarsi missing. Wings broken near base. Knob of haltere dark yellow. Abdomen brown above on most tergites, yellow on most of Ist tergite, sides of most other tergites and on sternites. Tergites 1—6 showing, with black bristles, marginal bris- tles longer. Genital capsule mostly hidden, brownish; cercus appearing small. Female.—Unknown. Holotype.—d, Costa Rica: Cartago, La 361 Suiza de Turrialba, Aug 1922, Schild (USNM) from the Melander Collection. Remarks.—Xanthina attenuata 1s distinct among the species with dorsally inserted aristae by the greater length versus width and more tenuous tip of the first flagello- mere. Relationship may be closest to X. dominicensis H. Rob. of the Lesser Antilles because of the row of erect hairs or setae ventrally on the middle femur and the short form of the cerci. The palpi of the new spe- cies are very thin and totally lack any brownish or blackish setae. ACKNOWLEDGMENTS The Philips XL30 ESEM Scope was op- erated by Scott Whitaker, manager of the National Museum of Natural History SEM laboratory. Marjorie Knowles, Technician in Botany, is thanked for work with the specimens and illustrations. LITERATURE CITED Robinson, H. 1970. Family Dolichopodidae, pp. |—92. In Papavero, N. ed. A Catalogue of the Diptera of the Americas South of the United States. Sao Pau- lo: Secretaria da Agricultura, vol. 40. . 1975. Bredin—Archbold—Smithsonian Biolog- ical Survey of Dominica, the family Dolichopod- idae with some related Antillean and Panamanian species (Diptera). Smithsonian Contributions to Zoology 185: i-iv, 1-141. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 362-372 A NEW GENUS AND NEW SPECIES OF COLPURINI (HETEROPTERA: COREIDAE) FROM NEW GUINEA HARRY BRAILOVSKY AND ERNESTO BARRERA Departamento de Zoologia, Instituto de Biologia, UNAM, Apdo Postal No. 70153, México 04510, D.E, México (e-mail: coreidae @ servidor.unam.mx ) Abstract.—The new genus Ullrihygia and five new species Ullrihygia iriana, Heisshy- gia tafa, Lothygia sordida, Lothygia consocia, and Sciophyroides splendidula collected in New Guinea are described in the tribe Colpurini (Coreidae). Dorsal habitus illustrations and drawings of the head, male genital capsule, and female genital plates are provided to distinguish each species. A key to the Lothygia species 1s given. Key Words: Guinea The island of New Guinea contains a re- markable and highly endemic assemblage of genera and species 1n the tribe Colpurini, many of which are still undescribed. The tribe Colpurini is represented by a number of species that show various bizarre morphological specializations. The striking features of these genera are the re- markable reduction of the hemelytra, the development of the head including the eyes and tylus, and the shape of male genital capsule and female genital plates (Brailov- sky 1996, 2000). The current paper is a further contribu- tion to this study. Provided are descriptions for one new genus and five new species and a key to the species belonging to the genus Lothygia Brailovsky. The following abbreviations are used for the institutions cited in this paper: AMNH (The American Museum of Natural History, New York); BPBM (Bernice P. Bishop Mu- Honolulu, Hawaii); BMNH (The Natural History Museum, London, Eng- land); UNAM (Instituto de Biologia, Univ- ersidad Nacional Autonoma de México); WUCO (Wolfgang Ullrich collection, Ger- most seum, Insecta, Heteroptera, Coreidae, Colpurini, new genus, new species, New many); ZSM_ (Zoologische Staatssa- mmlung, Miinchen, Germany). All measurements are given in millime- ters. Ullrihygia Brailovsky and Barrera, new genus Diagnosis.—Ullrihygia is related to Bau- mannhygia Brailovsky (1996), Nishihygia Brailovsky (2000), and Wygohygia Brailov- sky (1993a). Similar to Baumannhygia without ocelli, head quadrangular and wider than long, antenniferous tubercle armed with long robust spine, eyes small and glob- ular, buccula with sharp projection, femora unarmed, micropterous, abdomen elliptical, and connexivum higher than terga. Bau- mannhygia known only from the Philippine Republic, is recognized by having the tylus unarmed, the postocular tubercle protuber- ant, antennal segment I longer than III and shorter than maximum length of the head, humeral angles rounded and not raised, and scutellum apically acute. In Ullrihygia, re- corded from Irian Jaya, the tylus is apically acute and upturned to form a median horn, the postocular tubercle is absent, the length VOLUME 105, NUMBER 2 of antennal segment I is equal than II, and equal than maximum length of head, the humeral angles are raised, and the scutel- lum is apically rounded. This new genus resembles Nishihygia in having the tylus apically acute, antenniferous tubercle armed, ocelli absent, eyes globular, humeral angles slightly raised, femora unarmed, mi- cropterous, apex of scutellum rounded, ab- domen elliptical, and connexivum higher than terga. In Nishihygia, the frontal angles are rounded and not exposed, the posterior lobe of pronotum depressed, postocular tu- bercle moderately protuberant, and buccula rounded, without sharp acute projection. Wygohygia is like Ullrihygia with the eyes small and protrudent on short stalks, head quadrate, buccula with sharp projection, an- tenniferous tubercle armed, and frontal an- gles produced forward as rounded lobes. Wygohygia is recognized by the following combination of characters: head longer than wide, tylus apically globose, ocelli present, postocular tubercle protuberant, femora armed, apex of scutellum acute, and sub- macropterous reaching the posterior third of abdominal segment VI. Nishihygia and Wygohygia are recorded from New Guinea. Generic description.—Head: Wider than long, quadrangular, and dorsally slightly convex; tylus apically upturned to form a subacute median horn, extending anteriorly to and laterally higher than juga; juga un- armed, thickened, and apically barely acute: antenniferous tubercle armed with long ro- bust spine; antennal segment I stout, thick- est, slightly curved outward and equal than maximum length of head; segments IT and Ill cylindrical and slender; segment IV fu- siform; antennal segment II longest, IV shortest, and I equal to III: ocelli absent: preocellar pit deep, circular; eye small, globular, and protruding on short stalk; pos- tocular tubercle absent; buccula rounded, short, not projecting beyond antenniferous tubercle, with sharp middle projection; ros- trum reaching anterior margin of abdominal sternite Ill; mandibular plate absent. 363 Thorax: Pronotum wider than long, tra- peziform, bilobed, and non declivent; an- terior lobe slightly longer than posterior lobe, and each margin convexly rounded; collar weakly apparent; frontal angles pro- duced forward as conical lobe; humeral an- gles rounded, raised, and directed upward; posterolateral borders and posterior border straight; calli weakly convex. Anterior lobe of metathoracic peritreme reniform, poste- rior lobe sharp, small. Legs: Unarmed; tibiae cylindrical and sulcate. Scutellum: Wider than long, flat, mod- erately rectangular; apex rounded. Hemelytron: Micropterous, reaching an- terior third of abdominal segment I; wings reduced to small pads, widely separated; clavus and corium fused, and abdomen ex- posed mesially; membrane absent. Abdomen: Conspicuously elliptical; con- nexivum higher than terga; connexival mar- gin entire, with posterior angle of each seg- ment not extending on a short spine; terga Il to VI barely convex. Integument: Body surface dull, almost glabrous; antennal segments and legs with long erect setae; head with circular, grayish white farinose punctures; pronotum, scutel- lum, hemelytron, abdominal segments, tho- rax, and abdominal sterna strongly punc- tate; connexivum and upper margin of pleu- ral abdominal sterna smooth. Male genitalia: Posteroventral edge of genital capsule strongly produced forward as robust conical projection (Figs. 7—8). Female, unknown. Etymology.—Named after Wolfgang UII- rich, a distinguished German entomologist. Gender feminine. Type species.—Ullrihygia iriana Brai- lovsky, new species. Ullrihygia triana Brailovsky and Barrera, new species (Figs. 7-8, 18) Description.—Measurements: Male: Head length 1.48; width across eyes 1.84; interocular space 1.32: preocular distance 364 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-8. 1-6, Lothygia spp., male genital capsule in caudal view. 1, L. papuana. 2, L. cordigera. 3, L. enarotadi. 4, L. consocia. 5, L. sordida. 6, L. sandaracine Brailovsky. 7-8, Ullrihygia iriana, male genital capsule. 7, Caudal view. 8, Lateral view. 1.34; length antennal segments: I, 1.48; II, 2.02; HI, 1.48; IV, 1.08. Pronotum: Total length of anterior lobe 0.94; total length of posterior lobe 0.89; total width of anterior lobe 2.02; total width of posterior lobe 2.92. Scutellar length 0.40; width 1.32. Maxi- mum width of abdomen 4.80. Body length 8.95. Dorsal coloration: Head black; antennal segments I to III reddish brown, with basal joint of Ill ochre orange; antennal segment IV dark chestnut orange with basal third reddish brown; anterior lobe of pronotum black and posterior lobe reddish brown; an- terolateral margin and upper margin of hu- meral angles ochre orange; scutellum red- dish brown with wide ochre-orange longi- tudinal stripe at middle third; hemelytron and abdominal segments reddish brown; connexivum reddish brown with posterior margin of connexival segments V and VI dark orange. Ventral coloration: Head black; rostral segments dark yellow; thorax, abdominal sterna, and genital capsule dark orange yellow with punctures reddish brown to black; coxae black; trochanters yellow; femora yellow with apical third and few and scattered discoidal spots reddish VOLUME 105, NUMBER 2 brown; tibiae reddish brown with two yel- low rings one subbasal, the other one near middle third: tarsi chestnut orange. Type material.—Holotype: d, Indonesia: Irian Jaya, Paniai Province, N Mulia Trail to Dowome, 2,200—2,500 m, 8 July 1994, A. Riedel (Ent. Expl. New Guinea, 1990— 1994) (WUCO). Paratype: 1 d, same label data as holotype (UNAM). Etymology.—The name is derived from the type locality. Heisshygia tafa Brailovsky and Barrera, new species (Figs. 14, 16) Description.—Measurements: Male: Head length 1.60; width across eyes 1.56; interocular space 0.94; interocellar space 0.32; preocular distance 1.04; length anten- nal segments: I, 1.20; II, 1.80; HI, 1.24; IV, 0.88. Pronotum: Length 1.32; width across frontal angles 1.60; width across humeral angles 2.76. Scutellar length 0.88; width 0.96. Maximum width of abdomen 4.39. Body length 8.85. Female: Head length 1.60; width 1.60; interocular space 0.96; interocellar space 0.34: preo- cular distance 1.09; length antennal seg- ments: I, 1.28; II, 2.00; II, 1.40; IV, 0.92. Pronotum: Length 1.60; width across fron- tal angles 1.68; width across humeral angles 2.76. Scutellar length 0.84; width 1.06. Maximum width of abdomen 5.30. Body length 10.00. Male: Head: Pentagonal, longer than wide; antenniferous tubercle barely diverg- ing anteriorly; ocelli obscure; rostrum reaching anterior border of abdominal ster- nite VI (Fig. 16). Thorax. Pronotal disc flat, with posterior median depression. Scutellum: Wider than long, with apex rounded. Hemelytron: Micropterous, reaching posterior third of abdominal segment I; cla- vus and corium fused into coriaceous pad, and wings widely separated from each oth- er, leaving abdomen totally exposed mes- ally; membrane represented by small flap. across eyes 365 Genital capsule: Posteroventral edge with open and deep “*V” concavity, en- closed by two broad rounded lobes (Fig. 14). Coloration: Ground color bright reddish orange with head darker; antennal segments I to I bright reddish orange, and IV orange yellow with basal third bright reddish or- ange; apex of scutellum dark yellow; upper margin of connexivum and rostral segments I to IV pale chestnut orange; posterior mar- gin of acetabulae yellow; coxae, trochan- ters, and fore and middle femora pale or- ange yellow; hind femur dark chestnut or- ange with two yellow rings, one basal, the other one near middle third; tibiae pale chestnut orange with two yellow rings, one subbasal, the other one near middle third; tarsi chestnut orange; anterior lobe of meta- thoracic peritreme pale yellow, and poste- rior lobe reddish orange. Female: Head as longer as wide; rostrum reaching posterior third of abdominal ster- nite V. Coloration: similar to male. Connex- ival segments VIII and LX, dorsal abdomi- nal segments VIII and [X, and genital plates bright reddish orange. Type material.—Holotype d, Papua New Guinea: Mt. Tafa, 8,500’, February 1934, L. E. Cheesman (BMNH). Paratypes: | d, | 2, same data as for holotype (BMNH, UNAM). Discussion.—Among New Guinea Heis- shygia species (Brailovsky 1995), H. tafa, is superficially most similar to H. novogut- nensis Brailovsky (1993b), but may be sep- arated by the following characters: In H. tafa, the posteroventral edge of the male genital capsule has an open and deep V- shaped concavity, enclosed by two broad rounded lobes (Fig. 14), the rostrum reach the posterior third of abdominal sternite V or anterior border of VI, the antenniferous tubercle are unarmed, and the hemelytra reaching the posterior third of abdominal segment I. In H. novoguinensis (Fig. 17), the posteroventral edge of male genital cap- sule has two short lateral projections, sur- rounding a broad middle plate (Fig. 15), the 366 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9-17. 9-13, Sciophyroides spp. 9-11, Male genital capsule in caudal view. 9, S. splendidula. 10, S. acuminatus. 11, S. pumilus. 12-13, Female genital plates in lateral view. 12, 8. acuminatus. 13, S. splendidula. 14-17, Heisshygia spp. 14-15, Male genital capsule in caudal view. 14, H. tafa. 15, H. novoguinensis. 16-17, Head in dorsal view. 16, H. tafa. 17, H. novoguinensis. VOLUME 105, NUMBER 2 367 Fig. 18. Dorsal view of Ullrihygia iriana, male. rostrum reach the anterior third of abdom- inal sternite V, the antenniferous tubercle is armed, and each lobe raised and diverging anteriorly, and the hemelytra reaching the posterior third of abdominal segment II. Etymology.—This species is named for its occurrence in the Tafa mountains. Lothygia sordida Brailovsky and Barrera, new species (Figs. 5, 19) Description.—Measurements: Male: Head length 1.96; width across eyes 1.84; interocular space 1.04; interocellar space 0.48; preocular distance 1.28; length anten- 368 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nal segments: I, 1.40; I, 2.08; TI, 1.36; IV, 1.42. Pronotum: Length 2.12; width across frontal angles 1.56; width across humeral angles 3.80. Scutellar length 1.48; width 1.60. Body length 10.95. Female: Head length 2.16; width across eyes 2.00; inter- ocular space 1.16; interocellar space 0.52; preocular distance 1.48; length antennal segments: I, 1:52; I, 2-16: UM, 1:40; IV, 1.44. Pronotum: Length 2.44; width across frontal angles 1.68; width across humeral angles 4.16. Scutellar length 1.64; width 1.96. Body length 12.30. Male: Head: Longer than wide; tylus apically truncated; antenniferous tubercle with small lobelike projection; eyes spher- ical, elevated, and protruding on_ short stalks; postocular tubercle protuberant, and acutely expanded; rostrum reaching middle third of abdominal sternite VII. Thorax: Pronotum wider than long; fron- tal angles produced forward as_ conical teeth; humeral angles rounded, not exposed. Hemelytron: Macropterous, reaching apex of last abdominal segment. Genital capsule: Posteroventral edge lat- erally produced on a short blunt lobes, and between them a deep “U” concavity (Fig. 5) Dorsal coloration: Head shiny black with tylus bright reddish brown; antennal segment I bright reddish brown; segments II to IV dark chestnut orange; dorsal face of postocular tubercle dark yellow; anterior lobe of pronotum black, and posterior lobe dark reddish brown; scutellum dark reddish brown; clavus and corium pale reddish brown; hemelytral membrane dark brown; connexivum pale orange yellow with pos- terior third or posterior border of segments Ill to VIL black; abdominal segments III to VI dark orange and VII dark orange with extensive black marks. Ventral coloration: Head black; rostral segments dark to pale chestnut orange; thorax, abdominal sterna and genital capsule bright reddish brown with pleural abdominal segments III to VII pale orange yellow; anterior lobe of meta- thoracic peritreme yellow and_ posterior lobe black; coxae, trochanters and femora pale reddish brown; tibiae and tarsi pale chestnut orange. Female: Genitalia: Gonocoxae I en- larged dorso-ventrally, in caudal view closed, in lateral view barely convex, with upper border almost straight. Coloration: similar to male. Rostral segments I and I dark chestnut orange and III and IV pale chestnut orange; legs bright pale reddish brown with subbasal yellow ring at tibiae; connexival segments VIII and IX black with anterior third dark yellow; abdominal segments VIII and IX dark brown with dark yellow reflections; gonocoxae I bright red- dish brown with upper margin dark yellow; paratergite VIII] and IX bright reddish brown. Type material—Holotype d, Indonesia: Irian Jaya, Wamena Pass Valley, 1,900 m, 15-16 September 1990, A. Riedel (ZSM). Paratypes: 1 d, Indonesia: Irian Jaya, Wa- mena Llugwa Melanggama, 1,800—2,000 m, 9-12 September 1990, A. Riedel (UNAM). 1 6, 1 &, Indonesia: Irian Jaya, Wamena Pass Valley, 1,900 m, 15—16 Sep- tember 1990, A. Riedel (ZSM). Discussion.—This species is diagnosed mostly on basis of the shape of the poster- oventral edge of male genital capsule (Figs. 1—6), the presence of an acutely produced postocular tubercle, and by the shape of the eyes that are spherical, elevated, and pro- truding on a short stalks. In L. cordigera Brailovsky (1994), the postocular tubercle is not acutely produced, the eyes are spher- ical and weakly protrudent, and the poster- oventral edge of male genital capsule is heart-shape (Fig. 2). Etymology.—From the Latin sordidus, meaning dirty and referring to the blackish color. Lothygia consocia Brailovsky and Barrera, new species (Fig. 4) Description.—Measurements: Male: Head length 1.80; width across eyes 1.64; interocular space 0.96; interocellar space 369 NUMBER 2 By, VOLUME 10 e. Dorsal view of Lothygia sordida, mal 19: ‘ig. 370 0.48; preocular distance 1.20; length anten- nal segments: I, 1.36; I, 1.76; HI, 1.16; IV, 1.36. Pronotum: Length 2.04; width across frontal angles 1.60; width across humeral angles 3.36. Scutellar length 1.36; width 1.64. Body length 10.00. Female: Head length 1.92; width across eyes 1.72; inter- ocular space 0.98; interocellar space 0.47; preocular distance 1.36; length antennal seoments: 1 51-40; I 1883 IL, 24: LV; 1.36. Pronotum: Length 2.24; width across frontal angles 1.60; width across humeral angles 3.52. Scutellar length 1.48; width 1.64. Body length 11.30. Male: Head: Longer than wide; tylus apically truncate; antenniferous tubercle un- armed, with apex truncate; eyes spherical, and barely protrudent; rostrum reaching middle third of abdominal sternite VII; pos- tocular tubercle conically protuberant. Thorax: Pronotum wider than long; fron- tal angles produced forward as_ conical teeth; humeral angles rounded, not exposed. Hemelytron: Submacropterous, reaching middle third of abdominal sternite VII. Genital capsule: Posteroventral edge lat- erally produced in a short blunt lobes, and between them with large concave plate (Fig. 4). Dorsal Head black; tylus, and juga shiny dark reddish brown; anten- nal segments dark chestnut orange; dorsal surface of postocular tubercle dark yellow; anterior lobe of pronotum black, and pos- terior lobe and scutellum dark reddish brown; clavus and corium pale reddish brown; hemelytral membrane dark brown, with veins and basal angle dark orange; connexivum bright orange yellow, with posterior third of segments III to VII black; abdominal segments orange. Ventral col- Head black; rostral segments bright chestnut; thorax, abdominal sterna, and genital capsule bright reddish brown with pleural margin of abdominal sterna III to VII bright orange yellow; anterior lobe of metathoracic peritreme yellow and pos- terior lobe black; legs reddish brown, with dark chestnut orange reflections at femora. coloration: oration: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Female: Tylus apically with small spiny projection; hemelytra submacropterous, reaching posterior margin of abdominal segment VII. Genitalia. Gonocoxae I en- larged dorso-ventrally, in caudal view closed, in lateral view barely convex, with upper border almost straight. Coloration: similar to the male. Upper margin of me- tapleuron with orange yellow rectangular spot close to middle third; connexival seg- ments VIII and IX orange yellow, with pos- terior border black; abdominal segments VIII and IX dark orange: gonocoxae I bright reddish brown with internal angle and apical margin yellow; paratergite VIII and IX dark chestnut orange, with basal third black. Type material.—Holotype ¢, Indonesia: Irian Jaya, Paniai Province, S Mulia Trail to Wuyuneen, 1,900—2,000 m, 6—7 July 1994, A. Riedel (Ent. Expl. New Guinea, 1990-1994) (WUCO). Paratypes: 1 @, same data as for holotype (UNAM). 1 2, Indonesia: Irian Jaya, Paniai Province, N Mulia trail to Dowome, 2,200—2,500 m, 8 July 1994, A. Riedel (Ent. Expl. New Guin- ea, 1990-1994) (WUCO). Discussion.—Lothygia consocia, 1s easi- ly recognized by the shape of the poster- oventral edge of male genital capsule (Fig. 4), and by the submacropterous condition of the hemelytra which reach the middle third of abdominal sternite VII. On the pre- vious known species of Lothygia, the hem- elytra exhibit a macropterous condition, reaching or extending beyond the apex of last abdominal segment (Brailovsky 1994). Etymology.—From the latin consocius, a companion: one of several species occur- ring sympatrically in Irian Jaya. Key TO LOTHYGIA SPECIES (MODIFIED FROM BRAILOVSKY 1994) 1. Abdominal segments I to VI black; gonocoxae [iinvcaudaliview Opened Mir n:ssarecueten-s nyse 2 — Abdominal segments I to VI orange yellow; gonocoxae I in caudal view closed ........ 3 2. Tylus apically with a short spiny projection; posteroventral edge of male genital capsule VOLUME 105, NUMBER 2 with short median plate (Fig. 1) ........ SE Deo ELD int D SLRS ets L. papuana Brailovsky — Tylus apically globose, truncate; posteroventral edge of male genital capsule with large median plate (Fig. 3) 3. Frontal angles poorly developed; male genital Bie Steds sete L. enarotadi Brailovsky capsule with posteroventral edge sinuate, and laterally barely exposed (Fig. 6) ........ Oe ON ty ati he een L. sandaracine Brailovsky — Frontal angles well developed, produced for- ward as conical expansions ............. 4 4. Submacropterous, reaching middle third of ab- dominal sternite VII; posteroventral edge of male genital capsule with large concave plate Gigs. Saad. eee Ee — Macropterous, reaching or extending beyond the apex of last abdominal segment; posterov- consocia new species entral edge of male genital capsule without large*concave plate (Figs. 15) 25 os s...... 5 . Eyes spherical, elevated, and protruding on a nn short stalk; posteroventral edge of male genital capsule with deep ““U” concavity (Fig. 5)... L. sordida, new species — Eyes spherical, weakly protrudent; posterov- entral edge of male genital capsule heart- Shapedi(Bis% 1) oicem seas, - L. cordigera Brailovsky Sciophyroides splendidula Brailovsky and Barrerra, new species (Figs. 9, 13) Description.—Measurements: Male: Head length 2.79; width across eyes 2.17; interocular space 1.14: interocellar space 0.48; preocular distance 1.88; length anten- nal segments: I, 3.28; I, 4.52; II, 2.74; IV, 1.98. Pronotum: Length 3.59; width across frontal angles 2.04; width across humeral angles 5.64. Scutellar length 3.10; width 2.48. Body length 18.38. Female: Head length 2.82; width across eyes 2.23; inter- ocular space 1.20; interocellar space 0.53; preocular distance 1.82; length antennal segments: I, 3.03; II, 4.27; HI, 2.66; IV, 2.04. Pronotum: Length 3.53; width across frontal angles 2.10; width across humeral angles 6.07. Scutellar length 3.34; width 2.66. Body length 19.43. Male: Head: Elongate, longer than wide, and dorsally flat; tylus projecting an- teriad of juga, upturned to form sharp bifid horn; antenniferous tubercle unarmed; ocel- li with large diameter, and clearly tubercu- late; antennal segment II the longest, IV the 371 shortest, and I longer than HI; mandibular plate armed with sharp and small tubercle, visible in dorsal view; postocular tubercle protuberant; rostrum reaching middle third of abdominal sternite V. Thorax: Pronotum wider than long; fron- tal angles produced forward as small coni- cal teeth; humeral angles rounded, not ex- posed. Legs: Femora unarmed, and ventrally with tiny denticles; tibiae cylindrical and sulcate. Scutellum: Triangular, flat, and conspic- uously longer than wide; apex subacute. Hemelytron: Macropterous, reaching posterior margin of last abdominal segment. Genital capsule: Posteroventral edge tri- furcate, each lobe robust and wide; central lobe the longest, directed upward and back- ward, and lateral lobes remarkably folded downward (Fig. 9). Dorsal coloration: Head dark reddish brown with the space between ocelli and eye, and dorsal view of postocular tubercle dark yellow; antennal segments I to II dark chestnut orange with basal joint of III yel- low; antennal segment IV dark chestnut to dark yellow with basal and apical third dark reddish brown; pronotum, scutellum, cla- vus, and corium pale reddish brown with external margin of humeral angles and apex of scutellum yellow; middle third of calli dark reddish brown; hemelytral membrane dark ambarine; connexivum dark reddish brown with posterior margin yellow; dorsal abdominal segments bright orange with posterior margin of VII dark reddish brown. Ventral coloration: Ground color dark yel- low with punctures dark to pale reddish brown; rostral segments I and I dark chest- nut orange, III pale chestnut orange, and IV dark yellow; anterior lobe of metathoracic peritreme yellow and posterior lobe dark reddish brown; coxae and trochanters red- dish brown; femora reddish brown with ir- regular yellow ring close to middle third; tibiae reddish brown with two yellow rings, one subbasal, the other one close to middle third; tarsi pale chestnut orange; pleural ab- dominal sterna reddish brown with poste- rior margin yellow; genital capsule reddish brown. Female: Abdominal sternite VIL without plica and fissura; gonocoxae I enlarged dor- so-ventrally; in caudal view closed, in lat- eral view with upper third convex, and in- ner angle produced on a median lobe (Fig. 13). Coloration: similar to male. Connexi- val segments VIII and IX dark reddish brown with posterior margin yellow; dorsal abdominal segments VIII and IX and me- sosternum dark reddish brown; genital plates dark reddish brown with inner mar- gin of paratergite VIII and upper margin of gonocoxae I dark yellow. Type material.—Holotype ¢, Papua New Guinea, Owen Stanley Range, Goilala Bome, 1,950 m, 24 February—7 March 1958, W. W. Brandt (BPBM). Paiatypes: 1 2, Papua New Guinea, Eastern Highlands District No. 6, Pengage Camp, East Slopes Mt. Wilhelm, 2,770 m, 19-20 September 1959, L. J. Brass (Sixth Archbold Expedi- tion to New Guinea) (AMNH); 1 6, NE Papua New Guinea, East Highlands, Aiyu- ra, 1,800—1,900 m, 6 January 1965, J. Sed- lacek (UNAM). Discussion.—Brailovsky and Barrera (1996) described the genus Sciophyroides and includ 10 species, of them only S. ac- uminatus Brailovsky and Barrera, and _ S. pumilus Brailovsky and Barrera have the middle third of the posteroventral edge of male genital capsule produced upward (Figs. 10-11). On the other species the mid- dle third has a pronounced or relatively small U-shaped or V-shaped concavity, lat- erally enclosed by robust or tiny lobes. In Sciophyroides acuminatus the central lobe is elongate and acute (Fig. 10), in S. pumilus stout and shorter than lateral lobes (Fig. 11), and in S. splendidula Brailovsky, New Species, is stout and clearly longer than lateral lobes which are folded down- ward. The female of S. acuminatus has gon- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ocoxa I in lateral view remarkably exposed, with inner third widely concave (Fig. 12), and in the new species gonocoxa I in lateral view is barely convex, with inner third weakly concave (Fig. 13). The female of S. pumilus is unknown. Etymology.—The species name is de- rived from the Latin splendidulus, meaning diminutive and formed from splendidus, and refers to the remarkable shape of the male genital capsule. ACKNOWLEDGMENTS For access to the collections under their respective care or for the loan of specimens, we thank Randall T. Schuh (AMNH), Mick Webb (BMNH), Gordon Nishida (BPBM), Klaus Schonitzer (ZSM), and Wolfgang Ullrich. We also thank Albino Luna (UNAM) and Jesus Contreras (UNAM) for the dorsal view illustrations. LITERATURE CITED Brailovsky, H. 1993a. New genera and new species of Colpurini (Heteroptera: Coreidae) from the Fiji Is- lands and New Guinea. Proceedings of the Ento- mological Society of Washington 95(3): 435-448. . 1993b. New genera and new species of mi- cropterous Colpurini from Buru Islands and New Guinea (Heteroptera: Coreidae). Pan-Pacific En- tomologist 69(4): 281-289. . 1994. A new genus and four new species of Colpurini (Hemiptera: Heteroptera: Coreidae) from New Guinea. Journal of the New York En- tomological Society 102(2): 142-153. 1995. Description of three new species of Heisshygia Brailovsky from New Guinea (Hemip- tera: Heteroptera: Coreidae: Colpurini). Journal of the New York Entomological Society 103(4): 364-373. . 1996. New genera and new species of Col- purini (Insecta: Heteroptera: Coreidae) from Phil- ippines, Borneo, and Malaya. Annals of the Car- negie Museum 65(4): 383-391. . 2000. Nishithygia: a New Genus of Colpurini from New Guinea (Hemiptera: Coreidae: Corei- nae). Annals of the Entomological Society of America 93(1): 28-30. Brailovsky, H. and E. Barrera, 1996. Revision del complejo Sciophyrus (Hemiptera: Coreidae: Col- purini). Folia Entomologica Mexicana 96: 15— 106. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 373-379 ANNOTATED LIST OF THE BARK AND AMBROSIA BEETLES (COLEOPTERA: SCOLYTIDAE) OF MARYLAND, WITH NEW DISTRIBUTIONAL RECORDS ROBERT J. RABAGLIA Forest Pest Management, Maryland Department of Agriculture, 50 Harry S. Truman Parkway, Annapolis, MD 21401, U.S.A. (E-mail: rabaglrj @ mda.state.md.us) Abstract.—A total of 110 species of Scolytidae are reported from Maryland. Thirty species reported new to Maryland are: Hylastes opacus Erichson, Dendroctonus valens LeConte, Tomicus piniperda (L.), Phloeotribus dentifrons (Blackman), Carphoborus bi- cornis Wood, Polygraphus rufipennis (Kirby), Hylocurus flaglerensis Blackman, Micra- cisella opacicollis (LeConte), [ps avulsus (Eichhoff), Dryocoetes affaber (Mannerheim), D. autographus (Ratzburg), D. granicollis (LeConte), Lymantor decipiens (LeConte), Try- podendron betulae Swaine, T. lineatum (Olivier), T. retusum (LeConte), 7. scabricollis (LeConte), Ambrosiodmus obliquus (LeConte), Xyleborus planicollis Zimmermann, Xy- losandrus crassiusculus (Motschulsky), Cryphalus rubentis Hopkins, Pityoborus com- matus (Zimmermann), Pityophthorus balsameus Blackman, P. cariniceps LeConte, P. confusus Blandford, P. liguidambarus Blackman, P. opaculus LeConte, P. puberulus (LeConte), Pseudopityophthorus asperulus (LeConte), and Corthylus punctatissimus (Zimmermann). Of the 110 species reported in the state, 19 are not native to North America. Key Words: The geography of Maryland is excep- tionally diverse for its small size. Five physiographic provinces cover the State from east to west (Coastal Plain, Piedmont, Ridge and Valley, Blue Ridge, and Alle- gheny Plateau). With this diversity in land forms comes a diversity of forest types. In addition, due to its position in the Mid-At- lantic Region, forest types commonly found in the southeastern U.S. and the northeast- ern U.S. can be found in Maryland. The variety of forests in Maryland is re- flected in its forest insect fauna. This is es- pecially true of the bark and ambrosia bee- tles. On the Coastal Plain in southeastern Maryland, southern species, such as Den- droctonus frontalis (Zimmermann), are common, whereas in the mountainous, Scolytidae, distribution, Maryland western portion of the State, more northern species, such as Polygraphus rufipennis (Kirby) are found. Historically, many gen- eral collectors and scolytid specialists at the National Museum of Natural History, Smithsonian Institution (USNM), Washing- ton, DC and USDA in Beltsville, MD, have collected extensively in Maryland. The USNM collection and publications from the first half of the 20th century contains many bark beetle records from the state; however, recently there has been very little informa- tion on the scolytid fauna of Maryland. Since 1987, bark beetle collecting and surveys have been conducted as part of the routine forest pest survey activities by the Maryland Department of Agriculture. Ad- ditionally, since 1993, an extensive trapping 374 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Non-native scolytids found in Maryland. Species Origin Year Ambrosiodmus rubricollis (Eichhotf) Asia 1940's Dryoxylon onoharaensum (Murayama) Asia 1990's Euwallacea validus (Eichhoftt) Asia 1980's Hylastes opacus Erichson Europe 1990's Hylastinus obscurus (Marsham) Europe <1900 Hypothenemus californicus Hopkins Neotropics? 9202: Hypothenemus crudiae (Panzer) Neotropics? 19?? Scolytus mali (Bechstein) Europe 1950's Scolytus multistriatus (Marsham) Europe 1900's Scolytus rugulosus (Muller) Europe <1900 Tomicus piniperda (Linnaeus) Europe 1990's Xyleborinus saxeseni (Ratzeburg) Europe <1900 Xyleborus pelliculosus Eichhoft Asia 1980°s Xyleborus pfeili (Ratzeburg) Europe 1990's Xyleborus atratus Eichhoff Asia 1980's Xyleborus californicus Wood Asia 1990's Xyleborus dispar (Fabricius) Europe 1900's Xylosandrus germanus (Blandford) Asia 1930's Xylosandrus crassiusculus (Motschulsky) Asia 1980's program has been conducted as part of sev- eral USDA-APHIS programs to survey for exotic bark beetles that may have been in- troduced into the US. One hundred ten species are listed herein as occurring in Maryland. This is compa- rable to the numbers of species reported from some of the much larger western (Furniss et al. 1992, Furniss and Johnson 1987, Gast et al. 1989). Since 1987, 106 species have been collected in the above mentioned surveys, the remain- states ing four species are from historical records. Thirty species are reported as new to Mary- land. Most of these new records are for spe- cies previously reported in one or more neighboring states. The exceptions are Pi- tyoborus commatus (Zimmermann) and Hy- locurus flaglerensis Blackman with closest records from North Carolina and Florida, respectively. In a recent paper (Rabaglia and Williams 2002), two species of Hyle- sinus from western and midwestern North America were reported from Maryland. Sources of published records are Wood and Bright (1992) and Bright and Skidmore (1997). Species listed as ‘Newly Reported in Maryland” are those species not listed from Maryland in these publications or oth- er refereed publications since 1994. Species electronically reported as occurring in Maryland (1.e., Tomicus piniperda (L.), Hy- lastes opacus Erichson and Xylosandrus crassiusculus (Motschulsky)) are reported as new in this publication. Of the 110 species now reported from Maryland, 19 are not native to North Amer- ica (Table 1). Recent publications have dis- cussed the origins and the increasing oc- currence of these introductions (Atkinson et al. 1990 and Vandenberg et al. 2000). Since 1980, nine species of exotic scolytids have become established in Maryland, six of which are native to Asia. With increasing trade, an active shipping port and a diver- sity of forests, this number is likely to con- tinue to increase. Abbreviations for repositories of speci- mens new to Maryland are: MDAC— Maryland Department of Agriculture, An- napolis, MD; RJRC—Robert J. Rabaglia collection, Annapolis, MD and USNM— National Museum of Natural History, Smithsonian Institution, Washington, DC. Numbers of specimens appear in parenthe- sis with abbreviations. VOLUME 105, NUMBER 2 NEw MARYLAND RECORDS OF SCOLYTIDAE Species are arranged taxonomically as listed in Wood and Bright (1992). For Maryland localities, the first Maryland re- cord is given, followed by other counties from which the species has been collected. Subfamily Hylesininae Hylastes opacus Erichson Distribution.—Asia, Europe; USA: ME, NH, NY, VI, VA, WV, MD: Garrett Co, Keyser’s Ridge, 25 April 1996, Virginia pine log, MDA coll. (1, MDAC). Other counties: Allegany, Baltimore, Carroll, Cecil, Harford, Queen Anne’s, Washington. Comments: This exotic bark beetle is be- coming increasingly common in stumps of cut pine trees, and survey traps baited with a-pinene. Dendroctonus valens LeConte Distribution USA: AZ, CA, CO, DE, ID, IL, IN, KS, ME, MA, MI, MN, MT, NV, NH, NJ, NM, NY, NC, OH, OR, PA, SD, UT, VT, VA, WA, WV, WI, WY, MD: Anne Arundel Co., Annapolis, 6 April 1988, R.J. Rabaglia coll. (1, MDAC). Other counties: Allegany, Baltimore, Carroll, Cecil, Fred- erick, Garrett, Harford, Montgomery, Queen Anne’s, Washington. Comments: The red turpentine beetle is a common spe- cies across the state except for the extreme southeastern Coastal Plain where it is re- placed by Dendroctonus terebrans (Olivi- er). Tomicus piniperda (L.) Distribution.—Africa, Asia, Europe; USA: IL, IN, ME, MI, NH, NY, OH, PA, VT, WI, MD: Allegany Co., Oldtown, 3 April 1995, ex: a-pinene-baited trap, MDA coll. (1, USNM). Other counties: Frederick, Garrett, Washington. Comments: The pine shoot beetle has been found only in a-pi- nene baited funnel traps in the four western counties. It has not been collected in trees through 2001. 375 Phloeotribus dentifrons (Blackman) Distribution—USA: DC, FL, IL, KS, LA, MI, MS, NJ, ND, OH, PA, SC, TX, VA, WV, MD: Kent Co., Meliota, 7 May 1997, MDA coll. ex: Ipslure-baited trap (1, MDAC). Carphoborus bicornis Wood Distribution —USA: AL, DC, FL, PA, SC, MD: Talbot Co., Trappe, 2 February 1988, R.J. Rabaglia coll. (3, MDAC). Other counties: Baltimore, Calvert, Caroline, Queen Anne’s, Wicomico. Polygraphus rufipennis (Kirby) Distribution. USA: AK, AZ, CO, DC, ID, ME, MA, MI, MN, MT, NH, NJ, NM, NY, NC; ND; OR; PA; SD; ITN; UT, VT, VA, WA, WV, WY, MD: Garrett Co., Bit- tinger, 29 July 1988, R.J. Rabaglia coll. (1, MDAC). Subfamily Scolytinae Hylocurus flaglerensis Blackman Distribution. USA: DE, FL, MD: Kent Co., Sandy Bottom, 3 May 1994, R.J. Ra- baglia coll. (1, RJRC). Other counties: Bal- timore, Talbot. Comments: Prior to these collections, and collections in Delaware (Rabagha and Valenti 2003), this species was known only from the type series in Florida. Micracisella opacicollis (LeConte) Distribution —USA: DE, DC, IL, Ga, Ks, Ma, MI, Mn, Mo, Nj, Ny, Nc, Ou, Pa, Va, Wv, MD: Frederick Co., Ijamsville, 21 April, 1995, MDA coll. ex: Ipslure-baited trap (1, MDAC). Ips avulsus (Eichhoff) Distribution USA: AL, AR, DC, FL, GA, LA, MS, NJ, NC, PA, SC, TX, VA, WV, WI, MD: Washington Co., Ft Freder- ick State Park, 19 October 1993, R.J. Ra- bagha coll. ex: Ipslure-baited trap (3, MDAC). Other counties: Anne Arundel, 376 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Montgomery, Prince George’s, St. Mary’s, Talbot, Washington, Wicomico, Worcester. Dryocoetes affaber (Mannerheim) Distribution USA: AK, AZ, CO, ID, ME, MI, MN, MT, NH, NM, NY, NC, OR, PA, SD, TN, UT, WA, WV, WI, WY, MD: Garrett Co., Swallow Falls State Park, 27 June 1995, R.J. Rabaglia coll. (1, MDAC). Dryocoetes autographus (Ratzeburg) Distribution.—Africa, Asia, Europe; USA: AK, CA, CO, DE, ME, MI, MN, MT, NV, NH, NM, NY, NC, OR, PA, SD, TN, UT, VA, WA, WV, WI, WY, MD: Garrett Co., Swallow Falls State Park, 27 June 1995, R.J. Rabaglia coll. (2, MDAC). Dryocoetes granicollis (LeConte) Distribution—USA: AR, DE, DC, FL, GA, IL KY, LA, MO, NJ, NY, NC, OH, PA, TX, VA, MD: Baltimore Co., Glen Arm, 30 May 1987, W. Steiner coll. (2, USNM). Other counties: Anne Arundel, Baltimore, Cecil, Queen Anne’s, St. Mary’s. Lymantor decipiens (LeConte) Distribution. USA: DE, IA, KY, ME, MA, MI, MN, MS, NJ, NY, OH, PA, VA, WV, MD: Caroline Co., Tuckahoe State Park, 5 May 1994, R.J. Rabaglia coll. (1, MDAC). Other counties: Allegany, Balti- more, Caroline, Carroll, Cecil, Frederick, Harford, Kent. Trypodendron betulae Swaine Distribution —USA: ID, ME, MA, MO, MT, NH, NJ, NY, OR, SD, WI, MD: Garrett Co., Hutton, 10 April 1995, MDA coll. ex: a-pinene baited trap (1, MDAC). Trypodendron lineatum (Olivier) Distribution.—Africa, Asia, Europe; USAZ AK AZ; CA;:CO; Cli Dex MES MI, MO, MT, NV, NH, NM, NY, OR, PA, SD, TN, UT, WA, WV, WY, MD: Garrett Co., Accident, 5 May 1993, MDA coll. ex: ter- pene-baited trap (2, MDAC). Other coun- ties: Washington. Trypodendron retusum (LeConte) Distribution—USA: AZ, CA, CO, CT, ID, MI, MO, MT, NV, NH, NM, NY, OR, PA, SD, UT, VT, WA, WV, WI, WY, MD: Garrett Co., Accident, 12 April 1994, Joe Winters coll. ex: Ipslure-baited trap (1, MDAC). Trypodendron scabricollis (eConte) Distribution —USA: AR, ME, MO, MS, NJ, NC, PA, SC, VA, WV, MD: Allegany Co., Luke, 3 April 1995, MDA coll. a-pi- nene-baited trap (1, MDAC). Other coun- ties: Anne Arundel, Frederick, Garrett, Montgomery, Washington. Ambrosiodmus obliquus (LeConte) Distribution.—Africa, South America; USA: DE, DC, FL, GA, LA, MS, NC, VA, MD: Charles Co., Cedarville State Forest, 3 June 1988, R.J. Rabaglia coll. (1, RJRC). Other counties: Anne Arundel, Dorchester, Kent, Queen Anne’s, St. Mary’s, Talbot. Xyleborus planicollis Zimmermann Distribution.—USA: IL, IN, MO, NC, PA, MD: Garrett Co., Accident, 1 October 1999, MDA coll. ex: Allison-Pike trap (1, MDAC). Xylosandrus crassiusculus (Motschulsky) Distribution —Africa, Asia; USA: DE, FL, GA, LA, MS, NC, SC, TX, MD: Wash- ington Co., Big Pool, 29 July 1993, R.J. Rabaglia coll. ex: Frontalin-baited trap (1, MDAC). Other counties: Anne Arundel, Baltimore, Cecil, Harford, Kent, Prince George’s, Queen Anne’s, Worcester. Com- ments: Commonly known as the Asian am- brosia beetle, this species is becoming in- creasingly common in survey traps, and oc- casionally attacks landscape trees and nurs- ery stock. Cryphalus rubentis Hopkins Distribution USA: NC, PA, WV, MD: Garrett Co., Table Rock, 10 April 1995, MDA coll. ex: a-pinene-baited trap (3, MDAC). Other counties: Allegany. Com- VOLUME 105, NUMBER 2 ments: Hundreds of specimens have been collected in funnel traps baited with a-pi- nene in the western counties. Pityoborus commatus (Zimmermann) Distribution USA: AL, FL, GA, LA, MS, NC, SC, MD: Cecil Co., Elk Neck State Forest, 17 April 1997, MDA coll. ex: sulcatol-baited trap (5, MDAC). Com- ments: This is the first record of this species north of North Carolina. It has been found only in sulcatol-baited funnel traps. Pityophthorus balsameus Blackman Distribution —USA: ME, MI, MN, NH, NY, NC, WY, WI, MD: ‘Garrett ‘Co:.. Key- sers Ridge, 28 May 1997, MDA coll. ex: spruce (1, MDAC). Pityophthorus cariniceps LeConte Distribution USA: CT, DE, IN, KY, ME, MA, MI, MN, NH, NY, NC, OH, PA, SC, VA, WI, MD: Baltimore Co., Loch Ra- ven Resv., 20 April 1993, R.J. Rabaglia coll. ex: Ipslure-baited trap (1, MDAC). Other counties: Allegany, Cecil, Frederick, Garrett, Harford. Comments: This twig-in- festing species probably occurs state-wide, and is found often in Ipslure-baited funnel traps. Pityophthorus confusus Blandford Distribution USA: AL, AR, DE, FL, GA, LA, MS, NC, PA, SC, TX, VA, WV, MD: Talbot Co., Seth State Forest, 29 June 1988, R.J. Rabaglia coll. (1, MDAC). Other counties: Cecil, Dorchester, Prince George’s, St. Mary’s, Worcester. Pityophthorus liquidambarus Blackman Distribution USA: AR, CT, DE, DC, FIL; \GAgulls; -IN A, IEA, MS; NG; WY, MD: Somerset Co., Wellington, 19 May 1994, R.J. Rabaglia coll. (1, MDAC). Other counties: Charles. Pityophthorus opaculus LeConte Distribution USA: AK, AZ, CA, CO, DE, ID, ME, MI, MN, NV, NH, NM, NY, B77 PA; SD) UT, WYV,"W Ys “MD: 'Carroll Cos, Eldersburg, | June 1995, MDA coll. ex: Chalcoprax-baited trap (1, MDAC). Pityophthorus puberulus (eConte) Distribution USA: DE, DC, IN, KS, KY, ME, MA, MI, MN, NH, IA, NJ, NY, NC, OH, PA, RI, VA, WV, WI, MD: Cecil Co., Chesapeake City, 25 April 1994, R.J. Rabaglia coll. (1, MDAC). Other counties: Allegany, Calvert, Garrett, Howard, Kent, Prince George’s, St. Mary’s, Talbot, Wash- ington. Pseudopityophthorus asperulus (LeConte) Distribution USA: CT, DC, FL, GA, IL, LA, ME, MA, MS, MI, NJ, NY, NC, PA, TN, TX, WV, MD: St Mary’s Co., Hol- lywood, 12 June 1996, R.J. Rabaglia coll. (1, MDAC). Corthylus punctatissimus (Zimmermann) Distribution USA: AR, CO, CT, DC, FL, GA, IL, KY, MA, MI, MN, MS, NH, NJ, NY, NC, OH, PA, RI, TN, VT, VA, WV, MD: Montgomery Co., Brighton, 13 April 1995, MDA coll. ex: Ipslure-baited trap, (1, MDAC). Other Anne Arundel, Cecil, Kent. counties: LisT OF MARYLAND SCOLYTIDAE a = Species newly reported in Maryland. b = Exotic species found in Maryland. c = Species previously reported in Mary- land, but not collected in recent sur- veys. Hylesininae Hylastini Hylastes opacus Erichson—a,b Hylastes porculus Erichson Hylastes salebrosus Eichhoff Hylastes tenuis Eichhoff Hylurgops rugipennis pinifex (Fitch) Hylesinini Hylastinus obscurus (Marsham)—b Hylesinus aculeatus Say Hylesinus californicus (Swaine) 378 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hylesinus criddlei (Swaine) Hylesinus fasciatus LeConte Hylesinus pruinosus Eichhott Tomicini Dendroctonus frontalis (Zimmermann) Dendroctonus terebrans (Olivier) Dendroctonus valens LeConte—a Hylurgopinus rufipes (Eichhoff) Tomicus piniperda (L.)—a,b Bothrosternini Cnesinus strigicollis LeConte Phloeotribini Phloeotribus dentifrons (Blackman) Phloeotribus frontalis (Fabricius) Phloeotribus liminaris (Harris) a Phloeosinini Chramesus chapuisi LeConte—c Chramesus hicoriae LeConte Phloeosinus dentatus (Say) Polygraphini Carphoborus bicornis Wood—a Carphoborus bifurcus Eichhoff Polygraphus rufipennis (Kirby)—a Scolytinae Scolytini Scolytus mali (Bechstein)—b Scolytus multistriatus (Marsham)—b Scolytus muticus Say Scolytus quadrispinosus Say Scolytus rugulosus (Muller)—b Micracini Hylocurus flaglerensis Blackman—a Hylocurus langstoni Blackman—c Hylocurus rudis (LeConte) Micracis suturalis LeConte Micracis swainei Blackman Micracisella opacicollis (LeConte)—a Pseudothysanoes lecontei Blackman Thysanoes fimbricornis LeConte Ipini Ips avulsus (Eichhoff)—a Ips calligraphus (Germar) Ips grandicollis (Eichhoftf) Ips pini (Say) Orthotomicus caelatus (E1ichhoff) Pityogenes hopkinsi Swaine Pityogenes plagiatus (LeConte) Dryocoetini Dryocoetes affaber (Mannerheim)—a Dryocoetes autographus (Ratzburg)—a Dryocoetes betulae Hopkins Dryocoetes granicollis (LeConte)—a Dryoxylon onoharaensum (Murayama)—b Lymantor decipiens (LeConte)—a Crypturgini Crypturgus alutaceus Schwarz Xyloterini Trypodendron betulae Swaine—a Trypodendron lineatum (Olivier)—a Trypodendron retusum (LeConte)—a Trypodendron scbricollis (LeConte)—a Xyloterinus politus Say Xyleborini Ambrosiodmus obliquus (LeConte)—a Ambrosiodmus rubricollis (Eichhoff)—b Ambrosiodmus tachygraphus (Zimmer- mann) Euwallacea validus (Eichhoff)—b Xyleborinus saxeseni (Ratzburg)—b Xyleborus affinis Eichhoff Xyleborus atratus Eichhotf—b Xyleborus californicus Wood—b Xyleborus celsus Eichhotf Xyleborus dispar (Fabricius )—b Xyleborus ferrugineus (Fabricius) Xyleborus intrusus Blandford Xyleborus pelliculosus Eichhoff—b Xyleborus pfeili (Ratzeburg)—b Xyleborus planicollis Zimmermann Xyleborus pubescens Zimmermann Xyleborus sayi (Hopkins) Xyleborus viduus Eichhott Xyleborus xylographus (Say) Xylosandrus crassiusculus (Motschul- sky )—a,b Xylosandrus germanus (Blandford)—b a VOLUME 105, NUMBER 2 Cryphalini Cryphalus rubentis Hopkins—a Hypothenemus californicus Hopkins—b Hypothenemus crudiae (Panzer)—b Hypothenemus dissimilis (Zimmermann) Hypothenemus eruditus Westwood Hypothenemus interstitialis (Hopkins) Hypothenemus rotundicollis (Eichhoff) Trischidas atoma (Hopkins)—c Corthylini Conophthorus coniperda (Schwarz) Corthylus columbianus Hopkins Corthylus punctatissimus (Zimmermann)—a Gnathotrichus materiarius (Fitch) Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pityoborus commatus (Zimmermann)—a Pityophthorus balsameus Blackman—a Pityophthorus cariniceps LeConte—a Pityophthorus confusus Blandford—a Pityophthorus crinalis Blackman Pityophthorus dentifrons Blackman—c Pityophthorus lautus Eichhoff Pityophthorus liquidambarus Blackman—a Pityophthorus opaculus LeConte—a Pityophthorus puberulus (eConte)—a Pityophthorus pulchellus Eichhoff Pityophthorus pulicarius (Zimmermann) Pityophthorus pullus (Zimmermann) Pseudopityophthorus asperulus (LeConte) —a Pseudopityophthorus minutissimus (Zim- mermann) Pseudopityophthorus pruinosus (Eichhoff) Pseudopityophthorus pubescens Blackman ACKNOWLEDGMENTS Donald E. Bright identified or verified the identity of many of the specimens, his 379 help is greatly appreciated. I also thank the Maryland Department of Agriculture field staff for aiding in survey trap setup and col- lections. Also thanks to Gaye Williams for her critical reviews of earlier drafts of the manuscript. LITERATURE CITED Atkinson, T. H., R. J. Rabaglia, and D. E. Bright. 1990. Newly detected exotic species of Xyleborus (Co- leoptera: Scolytidae) with a revised key to species in eastern North America. Canadian Entomologist 122: 92-104. Bright, D. E. and R. E. Skidmore. 1997. A catalog of Scolytidae and Platypodidae (Coleoptera), Supple- ment | (1990-1994). National Research Council Research Press, Ottawa, Ontario. 368 pp. Furniss, M. M. and J. B. Johnson. 1987. List of Idaho Scolytidae (Coleoptera) and notes on new records. Great Basin Naturalist 47: 375-382. Furniss, M. M., J. B. Johnson, R. L. Wescott, and T. R. Torgersen. 1992. List of Oregon Scolytidae (Coleoptera) and notes on new records. Great Ba- sin Naturalist 52: 373-378. Gast, S. J.. M. M. Furniss, J. B. Johnson, and M. A. Ivie. 1989. List of Montana Scolytidae (Coleop- tera) and notes on new records. Great Basin Nat- uralist 49: 381-386. Rabaglia, R. J. and G. L. Williams. 2002. Two species of western North American Hylesinus Fabricius (Coleoptera: Scolytidae) new to the eastern United States. Proceedings of the Entomological Society ot Washington. 104: 1058-1060. Rabaglia, R. J. and M. A. Valenti. 2003. Annotated list of the bark and ambrosia beetles (Coleoptera: Scolytidae) of Delaware, with new distributional records. Proceedings Entomological Society of Washington. 105: 312-319. Vandenberg, N. J., R. J. Rabaglia, and D. E. Bright. 2000. New records of two Xyleborus (Coleoptera: Scolytidae) in North America. Proceedings of the Entomological Society of Washington. 102(1): 62-68. Wood, S. L. and D. E. Bright. 1992. A Catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxonomic Index. Great Basin Naturalist Memoirs No. 13. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 380-387 DESCRIPTION OF A NEW SPECIES OF BRACONIDAE, MONOCTONUS ALLISONI (HYMENOPTERA: BRACONIDAE: APHIDIINAE), AND DISCUSSION OF THE APHID-PARASITOID GUILD FOR NASONOVIA SPP. (HEMIPTERA: APHIDIDAE) IN NORTHWESTERN UNITED STATES KEITH S. PIKE, P. STARY, AND G. GRAF (KSP) Entomologist and (GG) Research Technician, Washington State University, lr rigated Agriculture Research and Extension Center, 24106 N Bunn Rd, Prosser, WA 99350, U.S.A. (e-mail: kpike @tricity.wsu.edu); (PS) Entomologist, Institute of Entomol- ogy, Academy of Sciences of the Czech Republic, Brani8ovska 31, 370 05 Ceské Budé- jovice, Czech Republic Abstract.—A new species of aphid parasitoid, Monoctonus allisoni Pike and Stary, is described from Washington. It is found in association with Nasonovia (Eokakimia) wah- inkae (Hottes) (Hemiptera: Aphididae) on pale larkspur, Delphinium glaucum Watson, in a montane forest setting. This is the first native Monoctonus in North America known to attack Nasonovia spp. A key to the parasitoid guild of Nasonovia is provided. Key Words: A new species of aphid parasitoid of the Braconid genus Monoctonus Haliday was found in Washington’s Blue Mountains at- tacking aphids of the species Nasonovia (Eokakimia) (Hottes) on pale larkspur, Delphinium glaucum Watson. This wahinkae unusual occurrence represents the first na- tive Monoctonus in North America known to attack Nasonovia. The parasitism was over a large colony, suggesting that the aphid was a preferred host of the parasitoid. The new species is described here. A key to the Nasonovia parasitoid guild for north- western United States is provided. MATERIAL AND METHODS All aphid parasitoids were reared from field-collected aphids. The key to the Na- sonovia parasitoid guild is based on rear- ings from aphid collections originating from a wide array of plants (see Pike et al. 2000), other original information of the au- thors, and from published records by Mack- aphid, parasitoid, Monoctonus, Nasonovia auer (1962a), Marsh (1979), and Stary and Remaudiere (1977). They are also the basis for establishing and confirming tritrophic (plant-aphid-parasitoid) associations. Addi- tionally, a broad framework of records of parasitoids of Nasonovia spp. from Europe [Stary (1966, 1976), Stary et al. (1971, 1973, 1977), Tizado (1992), Tizado and Nunez Perez (1991), and original files of P. Stary] was reviewed in reference to the present studies. Descriptive morphology characterizing Monoctonus allisoni follows that of Huber and Sharkey (1993), and Sharkey and Wharton (1997). Aphid nomenclature cor- responds to Remaudiere and Remaudiere (1997). DESCRIPTION Monoctonus allisoni Pike and Stary, new species (Figs. 1-7) Diagnosis.—The shape of the ovipositor sheath distinguishes the new species from VOLUME 105, NUMBER 2 stigma RI insertion point of effaced 2RS Figs. 1-7. 381 7 Monoctonus allisoni, female (illustrations not to same scale). 1-2, Forewings (in part), two draw- ings showing variation in venation. 3—4, Propodea (dorsal view, two drawings showing variation). 5, Petiole (dorsal view). 6, Genitalia with ovipositor sheath (lateral view). 7, Ovipositor sheath apex, close-up. Abbrevia- tions: Rl = distal abscissa of post-marginal vein [= metacarpus]: r, 2RS, 3RS = wing veins. the known Nearctic species of Monoctonus in which the ovipositor sheath is distinctly plough-share shaped; in M. allisoni, the sheath is only moderately widened ventral- ly. The latter character places M. allisoni close to Harkeria rufa (Cameron), but it is easily distinguished by antennal segment number: M. allisoni has 15-16 segments: H. rufa has 18-19 segments (see Discussion below). Etymology.—Named in recognition of David Allison, Washington State University Research Technologist, who has contributed significantly for many years to the research on aphid parasitoids in northwestern USA. Description.—Female: Head: Eye medi- um-sized, with sparse setae. Tentorio-ocular line shorter than half of inter-tentorial line. Malar space almost twice as long as tento- rio-ocular line (5:3). Maxillary palpus 4-, labial palpus 3-segmented. Antenna 15—16 segmented, filiform. Flagellomere | (F1) three times as long as wide, setae somewhat longer than half its width, without longitu- dinal placodes. Flagellomere 2 subequal to Fl, with one placode. Medial to apical fla- 382 gellomeres only 4 wider than F1, antenna not thickened toward apex. Mesosoma: Mesonotum smooth, with sparse long setae along sides. Notauli dis- tinct in ascendant portion only. Forewing (Figs. 1—2): Stigma about 5 times as long as broad. Metacarpus (R1) short, equal to about 4 stigma width. Radial abscissa (r) slightly longer than half of 3RS. 2RS sometimes effaced or rather colorless in basal portion. M+m-cu often colorless; stigma, metacarpus (R1), r, and 3RS well- pigmented and distinct. Propodeum (Figs. 3—4): Distinctly areo- lated, pentagonal areola complete in spite of some variation due to rugosities in upper portion. Metasoma: Petiole (Fig. 5) about twice as long as wide at apex; width at spiracles % shorter than segment length; spiracular tu- bercles prominent laterally, situated at end of basal third, surface feebly rugose, with indications of longitudinal carinae along sides and with 4—5 long setae along sides in middle and apical third. Genitalia: Ovipositor sheath (Fig. 6—7) slightly arcuate and only moderately wid- ened ventrally. Coloration: Generally bicolorous. Head brown, face and lower third yellow. Man- dible with brown apices. Palpi yellow. An- tenna brown, scape and pedicel light brown with yellow markings. Mesosoma with me- sonotum brown, sometimes scutellum and propodeum somewhat darkened, remainder yellow. Tegula brown. Wing venation light brown. Legs yellow. Metasoma with basal and apical areas yellow, brown in middle. Petiole basally yellow, remainder more or less brown. Ovipositor sheath yellow. Body length: 2.0—2.2 mm. Male. Antenna 17-18 segmented. Col- oration generally bicolorous to prevalently brown. Head prevalently brown with clyp- eus, lower part of gena, mandible and palpi light brown. Antenna brown, apex of ped- icel with yellow ring. Mesosoma brown with yellow markings, mesonotum brown. Wing venation light brown. Legs yellow to PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON largely yellow brown. Metasoma brown, except more or less yellow base. Material.—Holotype °, reared from Na- sonovia (Eokakimia) wahinkae (Hottes), USA, Washington, Columbia County, Blue- wood Ski Resort, 17-IX-2001, on Delphin- ium glaucum, sample coded A1G342, coll. G. Graf. Paratypes (same data as holotype), dry mounted 7 (1 2, 6 d) slide mounted 13 (8 2,5 3). Holotype deposited in Na- tional Museum of Natural History, Smith- sonian Institution (USNM), Washington, D.C. Paratypes deposited in USNM (2 @, 2 3d), Washington State University-Prosser (4 2,3 3), and collection of P. Stary in Ceské Budéjovice (3 2, 6 3d). PARASITOID GUILD OF NASONOVIA SPP. IN NORTHWESTERN UNITED STATES The following species-guilds were ana- lyzed: Nasonovia (Kakimia) alpina (Gillette & Palmer), N. (K.) aquilegiae (Essig), N. (Capitosiphon) crenicorna (Smith & Knowlton), N. (K.) cynosbati (Oestl.), N. (K.) houghtonensis (Troop), N. (K.) pole- monii (Gillette & Palmer), N. (Nasonovia) ribisnigri (Mosley), N. (Eokakimia) wah- inkae (Hottes), and unidentified Nasonovia spp. Most of the Nasonovia parasitoid guild members (Aphidius polygonaphis Fitch, Ephedrus californicus Baker, Lysiphlebus testaceipes (Cresson), Praon occidentale Baker, P. humulaphidis Ashmead, P. uni- cum (Smith) are more or less broadly oli- gophagous species across various aphid genera (Pike et al. 2000). Aphidius kaki- miaphidis Smith, is the most common par- asitoid of Nasonovia, although it does not attack all species in the genus. The new species, Monoctonus allisoni, is currently known to attack only a single aphid species, Nasonovia (Eokakimia) wahinkae. Nasonovia ribisnigri, a European aphid now widespread in North America, has a parasitoid guild of native broad oligophages (Ephedrus californicus, Praon humulaphi- dis, and P. unicum), and an introduced Eu- ropean parasitoid, Monoctonus crepidis (Haliday) (recognized in Canada and parts VOLUME 105, NUMBER 2 of the USA, see Mackauer 1962a, Marsh 1979; also see discussion below). Interest- ingly, this aphid is not attacked by the com- mon native Nasonovia-attacking parasitoid, Aphidius kakimiaphidis. KEY TO THE PARASITOIDS OF NASONOVIA IN NORTHWESTERN UNITED STATES l. Forewing with four enclosed cells along an- terior margin (R, M1, IR1, 2R1) (Fig. 8) einige hums udicwa read Ephedrus californicus (Baker) ~ Forewing with less than 4 enclosed cells alongyantenormareiny 2s. 23 2 es ie 2 2(1). Forewing with vein r perpendicular or al- most perpendicular to stigma (Figs. 1-2); Ovipositor sheath arcuate, curved downward (Figs. 6, 14) (Monoctonus) ........... 3 - Forewing with vein r not perpendicular to stigma: ovipositor sheath curved slightly upward"(Bigs 15-16)! 22. se. owes 4 3(2). Ovipositor sheath plough-share shaped (Fig. 14). Antenna 14-15 segmented ...... Bidito. coe ote Monoctonus crepidis (Haliday) - Ovipositor sheaths only gradually widened ventrally (Fig. 6). Antenna 15—16 segment- ..- Monoctonus allisoni Pike and Stary, n. sp. 4(2). Forewing with vein 2RS absent (Fig. 11) (CBM) Rates ccitee caress ahs, Stadt ee, Spenew ene 5) ~ Forewing with vein 2RS present (Figs. 9 NO) siepcireyericirat st asses lately eto 3, lp sheet aye panel! sp sugire ts 7 5(4). Flagellomere | dark. Mesosoma_ unicolo- TOUS; UNIfOrMYy Ganks cx 22s, grees chee ue kee) = 6 - Flagellomere | testaceous. Mesosoma dis- tinctly bicolorous, more or less lighter in lower part. Antenna 17—20 segmented Praon humulaphidis Ashmead . Antenna 17-18 segmented ........... PP ieatap ce seal<, cheapo Praon occidentale Baker - Antenna 14—16 segmented Ms. By cys Sg Re eeeuelis Praon unicum Smith . Forewing with vein M & m-cu incomplete (Fig. 9), Propodeum smooth (Fig. 12)... . hp eMs sR eataStie Lysiphlebus testaceipes (Cresson) ~ Forewing with vein M & m-cu complete (Fig. 10). Propodeum areolated (Fig. 13) (Aphidius) Antenna 13-14 sesmented™: <2 2... 2s Mes te Sis es Aphidius kakimiaphidis Smith ~ Antenna 17-19 segmented ........... Stata ane Sas 7 Aphidius polygonaphis (Fitch) 8(7). DISCUSSION Taxonomy of Monoctonus allisoni.—The combination of several key characters of M. 383 allisoni position it between Monoctonus and Harkeria Cameron (sensu van Achter- berg 1989). Harkeria rufa Cameron (spec- imens from Finland and from northwestern United States examined by P. Stary) have relatively distinct pentagonal areola on the propodeum, and a typical narrow petiole without prominent spiracular tubercles, whereas Paramonoctonus Stary (see Stary 1959) which was synonymized with Har- keria by van Achterberg (1989), lack the complete areola, i.e., have merely lower di- vergent carinae. This suggests a need for full re-evaluation and _ reclassification of Monoctonus, Harkeria and Paramonocton- us (possibly Boreogalba Mackauer, men- tioned below) In Monoctonus allisoni, the propodeum is distinctly areolated, in spite of some var- iation in size and sculpture (Figs. 3—4) due to less defined carinae among the rugae. Monoctonus allisoni has the maxillary palpi 4- and labial palpi 3-segmented (M. crepi- dis has the same characters), whereas 4- and 2 are defined for Harkeria and Monoctonus by van Achterberg (1989). The ovipositor sheath in M. allisoni is quite similar to H. rufa, opposite to the plough-share shaped characteristic of M. crepidis (Mackauer 1962a) and other species in northwestern United States (Pike et al. 2000). The fore- wing venation in M. allisoni is typical of Monoctonus, although some specimens have a strong reduction in vein pigmenta- tion with merely the radial abscissa present (Fig. 2). A similar variation is known for Harkeria (= Paramonoctonus) angustival- va (Stary) in Europe (Stary 1959). The combination of characters for M. allisoni of wing venation (variation, reduction), areo- lated propodeum, and shape of the petiole and ovipositor sheaths, point to some like- ness with the monotypic genus Boreogalba Mackauer (B. gladifer Mackauer, host un- known, see Mackauer 1962b, Mackauer and Stary 1967), but other characters differ widely, e.g., shape of the propodeum (B. gladifer squarish, M. allison anteriorly rounded) and number of antennal segments 384 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON S< —=2RS ) SS M & m- cu, fo << (9 oN aes A Figs. 8-16. Various features of parasitoids attacking Nasonovia aphids (illustrations not to same scale). 8— 11, Forewings. 8, Ephedrus californicus. 9, Lysiphlebus testaceipes. 10, Aphidius polygonaphis. 11, Praon occidentale. 12—13, Propodea. 12, L. testaceipes. 13, A. polygonaphis. 14—16, Genitalia. 14, Monoctonus cre- pidis. 15, L. testaceipes. 16, A. polygonaphis. VOLUME 105, NUMBER 2 (B. gladifer 11 segments; M. allisoni 15-16 segments). Monoctonus crepidis in North Ameri- ca.—Monoctonus crepidis (Haliday), a spe- cies broadly distributed in Europe, was de- tected on Nasonovia ribisnigri in Canada by Mackauer (1962a) (also reported by Marsh 1979). It was presumed introduced into North America from Europe via ship- transported plants with parasitized aphids. Evidence points to its establishment in North America in about 1953 in southeast- ern Québec. In 1977, Stary and Remaudiere (1977) further recognized its presence on N. ribisnigri in Québec and its outward spread in North America. Although not confirmed in northwestern United States, we expect it will become part of the Nasonovia parasit- oid guild; thus, it is included in the preced- ing key. Harkeria rufa and Nasonovia houghto- nensis.—This association was presented by Pike et al. (2000), but, after careful check- ing of the original labeled material, it was determined Macrosiphum osmaroniae was the correct host, instead of Nasonovia houghtonensis. Thus, Harkeria rufa is not recognized here as part of the Nasonovia- parasitoid guild. Nasonovia-parasitoid guilds in Europe and North America.—In Europe, the known Nasonovia species attacked by parasitoids are: Nasonovia (Kakimia) brachycyclica Holman, N. (Kakimia) dasyphylli Stroyan, N. (Nasonovia) nigra Hille Ris Lambers, N. (Nasonovia) pilosellae Boerner, N. (Naso- novia) ribisnigri (Mosley), N. (Kakimia) saxifragae Stroyan, and some unidentified species. The typical or common parasitoid guild for Nasonovia are Aphidius hiera- Stary, Harkeria (Stary), Monoctonus crepidis (Haliday), and Praon pubescens Stary. These are al- most all uniformly oligophagous on Naso- novia species. On rare occasion, the com- mon guild is supplemented by Aphidius ervi Haliday, A. picipes (Nees), Ephedrus cer- asicola Stary, E. persicae Froggatt, and E. plagiator (Nees). Also included in the guild clorum angustivalva 385 is Monoctonus hispanicus Tizado, which apparently attacks only species in the sub- genus Kakimia. Certain aphid-plant combinations may play a part in the parasitoid guild compo- sion for a given area. Aphidius hieracior- um and Praon pubescens parasitize Naso- novia on both its primary (Ribes) and sec- ondary (Hieracium) hosts. In contrast, Har- keria angustivalva and Monoctonus crepidis (Mackauer 1962a) attack Nasonov- ia only on Hieracium. Populations of Aphi- dius ervi, A. picipes, and Ephedrus plagia- tor, usually uncommon on Nasonovia, may increase depending upon proximity and type of other suitable aphids immediately present or nearby. In reviewing Nasonovia guilds in Europe and in North America, some basic phenom- ena are evident. (1) The European guild man- ifests more Nasonovia-specific oligophagous parasitoids than the North American guild, even though each continent has prevalently native aphid species. (2) The host range anal- ysis of the aphidiid guilds manifests four groups (Stary 1981): a) species-specific par- asitoids, which are rare (possibly Monoctonus hispanicus), b) oligophages specific to Na- sonovia (examples: Aphidius hieraciorum, A. kakimiaphidis, Praon pubescens); c) oligophages specific to Nasonovia and closely related groups (ex- amples: Monoctonus crepidis on Nasonovia, and Hyperomyzus [the latter genus is closely associated with Nasonovia, see Hille Ris Lambers 1949, Heie 1979, Foottit and Rich- ards 1993]):; and d) broadly oligophagous, or “sweeper species. Economic significance of Nasonovia.—A majority of the Nasonovia spp. (43 recog- nized, Remaudiere and Remaudiere 1997) reside in meadows and forest undergrowth from lowlands to high mountains, with some species adapted to urban parks and gardens where suitable hosts exist. Most of Harkeria angustivalva, and these are of littke or no economic impor- tance. Most are uncommon and the parasit- oid associations are unknown. There are, however, four species, Nasonovia brachy- 386 cyclica Holman, N. cynosbati (Oestlund), N. houghtonensis (Troop), and N. ribisnigri (Mosley), which at times are pestiferous on Ribes spp. (gooseberries and currants). The latter species also is recognized to feed on lettuce and other herbaceous plants. The aphids feeding on Ribes commonly cause retardation and a curling and twisting of ter- minal growth and leaves (Blackman and Eastop 1984). The distribution of the economic or po- tentially economic species is as follows: N. brachycyclica, Czech Republic (Holman 1972); N. cynosbati and N. houghtonensis: North America; and N. ribisnigri, Europe, east to Ukraine, and accidentally introduced in North America and South America (Blackman and Eastop 1984). ACKNOWLEDGMENTS This study was made possible in part with support from Washington State Uni- versity, Grant A6007105 (Grant Agency of the Academy of Sciences, Czech Republic), and the Entomology Institute Project Z5007907 (Academy of Sciences of the Czech Republic). LITERATURE CITED Achterberg, C. van. 1989. Revision of the subtribe Monoctonina Mackauer sensu stricto (Hymenop- tera: Braconidae: Aphidiinae). Zoologische Me- dedelingen 63: 1-22. Blackman, R. L. and V. FE Eastop. 1984. Aphids on the World’s Crops: An Identification and Infor- mation Guide. John Wiley & Sons, New York, 466 pp. Foottit, R. G. and W. R. Richards. 1993. The Genera of the Aphids of Canada. The Insects and Arach- nids of Canada, Part 22, Agriculture Canada, Pub- lication 1885, Canada Communications Group— Publishing, Ottawa, Canada, 766 pp. Heie, O. E. 1979. Revision of the Aphid Genus Na- sonovia Mordvilko, Including Kakimia Hottes & Frison, with Keys and Descriptions of the Species of the World (Homoptera: Aphididae). Entomo- logica Scandinavica, Supplement No. 9, 105 pp. Hille Ris Lambers, D. 1949. Contributions to a mono- graph of the Aphididae of Europe. IV. Temminck- ia 8: 182-232. Holman, J. 1972. Nasonovia (Kakimia) brachycyclica sp. n. on Ribes grossularia L. From Czechoslo- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON vakia. Acta Entomologica Bohemoslovaca 69: 317-323. Huber, J. T. and M. J. Sharkey. 1993. Structure, pp. 13-59. In Goulet, H. and J. T. Huber, eds. Hy- menoptera of the World: An Identification Guide to Families. Research Branch, Agriculture Cana- da, Ottawa, Ontario, Publication 1894/E, 688 pp. Mackauer, M. 1962a. Monoctonus crepidis (Haliday) (Hymenoptera: Aphidiidae) an aphid parasite new to North America. Canadian Entomologist 94: 1089-1093. . 1962b. A new genus of Aphidiidae (Hyme- noptera) from northern Canada. Canadian Ento- mologist 94: 1107-1109. Mackauer, M. and P. Stary. 1967. World Aphidiidae, pp. 1-167. Jn Delucchi V. and Remaudieére G., eds. Index of Entomophagous Insects. LeFrancois, Paris. Marsh, P. M. 1979, Aphidiidae, pp. 295-313. Jn Krom- bein, K. V.. P. D. Hurd, Jr, D. R. Smith, and B. D. Burks., eds. Catalog of Hymenoptera of Amer- ica North of Mexico, Vol. 1, Symphyta and Apo- crita (Parasitica). Smithsonian Institution Press, Washington, D.C. Pike K. S., P. Stary, T. Miller, G. Graf, D. Allison, L. Boydston, and R. Miller. 2000. Aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) of North- west USA. Proceedings of the Entomological So- ciety of Washington 102: 688-740. Remaudiere, G. and M. Remaudiere. 1997. Catalogue of the World’s Aphididae, Homoptera Aphidoidea. Institut National de la Recherche Agronomique, Paris, 473 pp. Sharkey, M. J. and R. A. Wharton. 1997. Morphology and terminology, pp. 20-37. /n Wharton, R. A., P.M. Marsh, and M. J. Sharkey, eds. Manual of the New World Genera of the Family Braconidae (Hymenoptera). Special Publication of the Inter- national Society of Hymenopterists, No. 1, 439 PP- Stary, P. 1959. A revision of the European species of the genus Monoctonus Haliday (Hymenoptera, Braconidae, Aphidiinae). Acta Societatis Ento- mologicae Cechosloveniae 56: 237-251. . 1966. Aphid Parasites of Czechoslovakia. Ac- ademia Publication House, Prague and Dr. W. Junk Publishers, The Hague, 242 pp. . 1976. Aphid parasites (Hym., Aphidiidae) of the Mediterranean area. Transactions of the Cze- choslovak Academy of Sciences, Series of math- ematical and natural sciences 86(2): 1-95, co-edi- tion with Dr. W. Junk Publishers, The Hague. . 1981. On the strategy, tactics and trends of host specificity evolution in aphid parasitoids (Hy- menoptera, Aphidiidae). Acta Entomologica Boh- emoslovaca 78: 65-75. Stary, P, G. Remaudiere, and F Leclant. 1971. Les Aphidiides (Hym.) de France et leurs hotes (Hom., VOLUME 105, NUMBER 2 Aphididae). Mémoire Hors Série, Entomophaga 5: 1-72. . 1973. Nouvelles données sur les Aphidiides de France (Hym.). Annales de la Societé Ento- mologique de France, N.S., 9: 309-329. . 1977. Nouveaux compléments sur les Aphi- diides (Hym.) de France et leurs hétes. Annales de la Societé Entomologique de France, N.S. 13: 165-181. Stary, P. and G. Remaudiere. 1977. Some aphid para- 387 sitoids (Hym. Aphidiidae) from Nearctic America. Annales de la Societé Entomologique de France, N.S. 13: 669-674. Tizado, E. J. 1992. Monoctonus hispanicus sp.n. (Hy- menoptera, Braconidae: Aphidiinae). Anales de Biologia 18: 53-57. Tizado, E. J. and E. Nunéz Pérez. 1991. Aportacion al conocimiento en Espana de los parasitoides de la subfamilia Aphidiinae (Hym., Braconidae). Bolle- tin de Sanidad Vegetal de las Plagas 17: 545-554. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 388-396 REDESCRIPTION OF BLASTOBASIS YUCCAECOLELLA DIETZ 1910 (LEPIDOPTERA: GELECHIOIDEA: COLEOPHORIDAE: BLASTOBASIND), WITH OBSERVATIONS ON ITS BIOLOGY DAvib ADAMSKI AND OLLE PELLMYR (DA) Department of Systematic Biology, Entomology Section, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (email: dadamski @sel.barc.usda.gov); (OP) Department of Biological Sciences, University of Ida- ho, P.O. Box 443051, Moscow, ID 83844-3051, U.S.A. (e-mail: pellmyr@uidaho.edu) Abstract.—A redescription of the adult of Blastobasis yuccaecolella Dietz 1910 is giv- en, and for the first time the larva and pupa are described. Observations on the biology of this species are discussed. Photographs of the habitat, host plant, and dried floral parts and seed pods infested by the larva are included. Key Words: Three blastobasine moths, Holcocera gt- gantella (Chambers 1876), H. paradoxa Powell 1976 and Blastobasis yuccaecolella Dietz 1910 are known to feed on Yucca (Agavaceae) in southwestern United States and México. Blastobasis yuccaecolella 1s known from Texas and from Nuevo Leon, México. Adults of this species were first reared from the seed pods of Yucca baccata Torrey in Texas (Dietz 1910). Holcocera gi- gantella and H. paradoxa are known to be sympatric in at least part of their ranges, but it is unknown whether they co-occur with Blastobasis yuccaecolella. Although the Blastobasinae (Coleophor- idae) are treated by Powell (1980) and Pow- ell et al. (1999) as scavengers, several spe- cies feed primarily on living plant tissue. Here we redescribe B. yuccaecolella and for the first time describe the larva and pupa. In addition, the host, habitat, larval habits, and distribution of B. yuccaecolella are dis- cussed. MATERIALS AND METHODS Larvae of Blastobasis yuccaecolella ranging from second to final instar were Blastobasinae, distribution, Yucca collected from the seed pods of Yucca tre- culeana Carriere in February, March, and November between 1995—2000 from La- guna Atascosa National Wildlife Refuge, Cameron County (26°15'N, 7°21’W), and Santa Ana National Wildlife Refuge, Hi- dalgo County, Texas (26°02'N, 98°05'W). A single specimen was reared from Yucca filifera Chabaud (Agavaceae) in Bano de San Ignacio, near Linares, Nuevo Leon, México (24°52'N, 99°34'W). In addition, a series of specimens used by Dietz (1910) for his original description was examined at the National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM). All of these latter specimens are labelled as having been collected in 1897 from Yucca baccata in Texas. For SEM study, larvae and pupae were cleaned in 10% alcohol with a camel’s hair brush, and dehydrated in increasing concen- trations 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 Cressing- VOLUME 105, NUMBER 2 389 Figs. 1-7. Habitat, host, pupa, and adult of Blastobasis yuccaecolella. 1, Yucca treculeana and habitat in Texas. 2, Dried fruits. 3, Dried seed pod. 4, Pupa. 5, Live adult. 6-7, Pinned specimens of reared adults ton sputter coater. The ultrastructure of the Morphological observations and mea- larva and pupa was studied with an Amray surements of the larva, pupa, and adult, 1810 scanning electron microscope at an were made using dissecting (reflected light) accelerating voltage of 10 kV. and compound microscopes (transmitted 390 light) with a calibrated micrometer. The Methuen Handbook of Colour (Kornerup and Wanscher 1978) was used as a color standard. Genitalia were dissected as de- scribed by Clarke (1941), except Mercuro- chrome and chlorazol black were used as staining agents. All pinned specimens of reared adults and lots of immature speci- mens were assigned voucher label and de- posited in USNM. RESULTS Blastobasis yuccaecolella Dietz 1910 (Figs. 1-23) Adult.—Head: Scales of vertex and frontoclypeus brown tipped with pale brown, or brownish gray tipped with pale brown; outer surface of labial palpus dark brown or brown intermixed with pale brown scales to near apical area of segment II and basal and apical areas of segment III, inner surface pale brown intermixed with few brown and dark brown scales along ventral and subventral surfaces; scape of antenna with brown scales tipped with pale brown, flagellum pale brownish gray; fla- gellomeres wider, with longer cilia in male than in female; first flagellomere excised, forming a deep notch in male; proboscis pale brown. Thorax: Tegula with scales brownish gray tipped with pale gray, or pale brown basally near white distally, or pale brown basally brownish gray distally; me- sonotum with scales brownish gray tipped with pale gray, or scales dark brownish gray tipped with pale gray basally, or brownish gray tipped with pale gray distally. Fore- wing (Figs. 6—7): Length 5.6—8.5 mm (n = 122): Brown scales tipped pale brown and white, intermixed with pale brown and dark brownish-gray scales; median fascia com- plete or incomplete; base with dark brown- ish-gray scales between costa and CuP absent; cell with three dark brownish-gray spots, | near middle and 2 near distal end; marginal spots distinct, faint, or absent; venation with M, and CuA, branched from distoposterior angle of cell; present or PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON CuA, distal from CuA,; undersurface brown. Hindwing: Translucent brownish gray; venation with cubitus 4-branched, each vein stalked in series and extending to posterior margin. Male genitalia (Fig. 9): Uncus wider ba- sally than long, gradually narrowed toward a rounded apex, sparsely setose dorsally, with 3 pairs of long lateral setae; gnathos a rounded ventral support for tuba analis, posterorventral margin medially produced into a small bidentate process; vinculum wide; juxta bandlike; ventral part of valva distally emarginate ventrally near abruptly rounded base; apically produced into a spinelike process with a flat inner margin: valva with dorsal part setose, elongate, dig- itate, widening basally and confluent with proximal flange; proximal flange elongate, apicoventral margin rounded, with dense stout setae to margin; microtrichiate mem- brane across posterior margin of tegumen and margins of proximal flange; tergal setae present; aedeagus nearly straight, slightly narrowed from base to apex; sclerite of ae- deagus singly twisted from midlength; anel- lus conical, with several stout setae. Female genitalia (Fig. 10): Ovipositor telescopic, with 4 membranous subdivi- sions; eighth sternum elongate, setose along posterior margin; eighth tergum with a nar- row, elongate, darkly pigmented pattern along median longitudinal axis; ostium bur- sae slightly posterior to seventh sternum, surrounded by sparse microspinules on sur- rounding membrane; seventh sternum with perpendicular notch on posterolateral mar- gin; inception of ductus seminalis slightly anterior to ostium bursae; ductus bursae long, membranous in about posterior ¥,, with rows of internal, imbricate platelets within anterior ¥,; corpus bursae slightly elongate, with lobe on posterior end and hornlike signum near midlength. Larva (Figs. 11—20).—Length 6.1—10.5 mm [n = 39]. Body pale violet on dorso- lateral surface, tonofibrillary platelets, areas between folds, and venter white; setae on small pinacula; prolegs with crochets on VOLUME 105, NUMBER 2 391 Figs. 8-9. mm. 9, Male genitalia. Scale = 0.5 mm. A3-A6 and A10. Head (Figs. 11-14, 19): Hypognathous; adfrontal sclerites slightly narrowed (Fig. 11), Pl about 4 times the length of P2, about even with apex of frons and slightly below F2; F2 above apex of frons, slightly longer than Fl; C3 at least twice the length of Fl; C2 in straight line with C3, and slightly above Cl; A2 in straight line with and above Al, Al and A3 at least twice length of A2; LI laterad to Wing venation and male genitalia of Blastobasis yuccaecolella. 8, Wing venation. Scale = 1.0 A3, about ¥, length of A3; SS3 at least twice the lengths of SS2 and SSI; SS] slightly laterad of mandibular condyle, SS2 posterad to stemma-5; S2 about twice the length of S1 and S3; S3 lateroposterad to SS2, S2 lateroposterior to stemma-1l, and S1 adjacent to stemma-3; labrum with 12 setae, 2 pair of subequal median setae in same horizontal plane and perpendicular with median longitudinal axis, 2 pair of 392 ae Fig. 10. Female genitalia of Blastobasis yuccae- colella. Scale = 1.0 mm. equal marginal setae ventrolaterad to me- dian setae, and 2 pair of subequal lateral setae along margin; mandible with two dis- tinct dentitions and 2 subequal setae on ventral surface (Fig. 19); apical sensilla on maxillary palpus (Fig. 13) similar to that of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Blastobasis graminea (Adamski 1999); an- tennal sensilla (Fig. 14) with 2 more sen- silla basiconica and | more sensillum tri- chodeum than B. graminea (Adamski 1999): submental pit oblong. Prothorax (Fig. 17): Shield with SD1 slightly longer than XDI and XD2; D2 about as long as SD1, about equidistant to XD2 and XD1, and in straight line with D1; SD2 as long as or slightly longer than D1, about equi- distant to SD1 and XD2, and anterirad to D2 and D1; L-group on pinaculum ven- troanterirad to spiracle; L1 about twice the length of L2 and L3; SV1 slightly longer than SV2; V1’s close, about ¥,—/, distance as V1’s on T2—T3. Mesothorax and meta- thorax (Fig. 17): D2 about 2Y, times the length of Dl; D2 and D1 on same pinacu- lum; D2 slightly anterirad and more than twice the length of Dl; SDI and SD2 on same pinaculum; SDI slightly anterirad to SD2 and more than twice the length of SD2; L1 and L2 on same pinaculum, ven- troanterior to pinaculum bearing SD1 and SD2; L1 at least twice the lengths of L2 and L3; L3 dorsoposterior of pinaculum bearing LI and L2; SV1 about as long as Ll, in straight line with L3. Abdomen (Figs. 15-16, 18, 20): Al—A2 (Fig. 18) with D2 at least 2% times length of D1, SD1 above spiracle; SD2 microscopic, on same pinaculum as SDI (not shown); L2 and L1 on same pinaculum, beneath and slightly posteriad spiracle; L2 about 2% times length of L1; L3 posteroventrad of pina- culum bearing L1—L2, and in line with D2; SV-group trisetose, SV1 about 2% times length of SV2 and SV3; crochets (Fig. 15) uniordinal, in circle, reduced or absent lat- erally; A8 (Fig. 20) with SV1 in line with L3; A9 (Fig. 20) with pinaculum bearing D1—D2 orientated about 45—90° from pin- aculum bearing LI—L2; SD1 closer to dor- sal pinaculum than to lateral pinaculum; L3 proximal to SV-group setae; SV1I and SV2 on separate pinacula, SV2 posteroventrad to SV 1; V-group setae slightly farther apart on A10 than on A8—A9; A10 (Figs. 16, 20): VOLUME 105, NUMBER 2 Figs. 11-16. 393 Scanning electron micrographs of larva of Blastobasis yuccaecolella. 11-12, Head capsule. Scale = 100 p. 13, Sensilla on apex of maxillary palpus. Scale = 10 p. A2 = sensillum styloconicum; Al, A3, M1, M2, L1, L2, and L3 = sensilla basiconica. 14, Sensilla on apical portion of antenna. Scale = 10 p. | sensilla basiconica, 2 = sensilla chaetica, 3 sensillum styloconicum, 4 = sensilla trichodeum. 15, Proleg. Scale 100 2. 16, Anal plate. Scale = 100 p. anal plate with SD2 and D2 equal in length, SD1 longest, D1 the shortest. Pupa (Figs. 21—23)—Ovate, brownish yellow to yellowish brown; length 4.0—4.8 mm (n = 20); mostly smooth with shallow wrinkles throughout; frontoclypeal suture present; epicranial suture distinct; caudal part of antennae converging, separating at distal end exposing metathoracic legs; max- illary palpus present; labial palpus absent; prothoracic leg exposed laterad to maxillary palpus. Biology.—The larvae of Blastobasis yuc- caecolella feed on dried remains of flowers (Fig. 2) and dried seed pods (Fig. 3) that accumulate in the leaf rosette. Old fruits oc- casionally remain on the infructescence for one or a few years, or become wedged in the narrow groove of rigid Yucca leaves created by their upwardly curved edges. 394 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Sv3 sv1 20 Figs. 17-20. Mandible and setae maps of larva of Blastobasis yuccaecolella. 17, Mandible. 18, Prothorax, meso- and metathorax. 19, Abdominal segments 1—2. 20, Abdominal segments 7—10. Larvae are found concealed underneath and inside old fruits, where they feed on the dried fleshy portions of the fruit and to a lesser extent on seeds. The feeding results in diagnostic frass beneath the fruits. Lar- vae also forage freely in larger aggregates of old flowers and fruits that may increase up for years and form strongly decomposed humuslike material. Pupation takes place in a thin cocoon lined externally with frass (Fig. 4), typically located beneath old fruits or other plant parts. Adults are known to emerge between December and March or later. Most emergences take place late in the afternoon. The emerging adults are cryptic on the plant matter (Fig. 5), and tend to hide VOLUME 105, NUMBER 2 Figs. 21- 1 mm. rather than to fly when disturbed. Adults have been observed at night in flowers. DISCUSSION Yucca baccata is a readily identified spe- cies, primarily distributed from New Mex- California (Reveal ico to easternmost 1977). The easternmost published record of Yucca baccata is from Crockett County, Texas, southwest of Big Lake 23. Pupa of Blastobasis yuccaecolella. 21, Lateral view. 22, Dorsal view (Webber Scale 23, Ventral view 1953), and all other reports refer to sites west of Marathon (McKelvey 1938). The species has been observed farther east than these records 43 km W of Eldorado in west- ernmost Schleicher County, Texas, in the transition zone from Chihuahuan desert to the Edwards Plateau ecoregion (Pellmyr, unpublished observation). Thus, it is likely that the population of Blastobasis yuccae- colella from which this species was first 396 collected and described originated from a site within the Chihuahuan desert, in the westernmost quarter of Texas. Elevation ranges from sea level for the easternmost records, 250 m for the Mexican site, and 750 m or more for Yucca baccata sites in western Texas. The characteristic habitats of Yucca tre- culeana in eastern Texas (Fig. 1) and Nuevo Leon are Tamaulipan grasslands, often in- terspersed by dense thornscrub (Pellmyr 1999). Yucca baccata occurs in several communities in westernmost Texas; so, the habitat of Blastobasis yuccaecolella in that area cannot be determined from available locality data. Powell and Mackie (1966) and Powell (1984) reared Holcocera gigantella and H. paradoxa trom both green and dried seed pods of Yucca whipplei (Torrey) Trelease. In addition, H. gigantella was reared by Co- quillett and Riley from seed pods of Y. brevifolia Engelmann, and Powell reared the moth from flowers of Agave orcuttiana Trelease (Agavaceae) (Powell 1984). Blas- tobasis yuccaecolella has not yet been reared from green seed pods of either Yucca baccata or Y. treculeana. In general, it appears that Blastobasis yuccaecolella and the above mentioned hol- cocerine moth species that utilize Yucca for their larval host feed within dried seed pods, and to a lesser extent, on the stems and floral parts of the plant. The number of generations per year, timing of oviposition, and ovipositional sites are not known for B. yuccaecolella. ACKNOWLEDGMENTS We thank Scott D. Whittaker, Director, Scanning Electron Microscopy Laboratory, Smithsonian Institution, Washington, DC, for his assistance with ultrastructural stud- ies; John S. Steiner and James FE Di Loreto, Office of Imaging, and Photographic Ser- vices, Smithsonian Institution, for the pho- tographs of the imagos and production of digital plates; and Kuniko Arakawa for the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON illustrations of the setal maps of the larva. Field work for this study was funded to Olle Pellmyr by the National Science Foun- dation and the National Geographic Socie- ty. LITERATURE CITED Adamski, D. 1999. Blastobasis graminea, new species (Lepidoptera: Gelechioidea: Coleophoridae: Blas- tobasinae), a stem borer of sugar cane in Colom- bia and Venezuela. Proceedings of the Entomo- logical Society of Washington 101(1): L64—174. Chambers, V. T. 1876. Tineina. Canadian Entomologist 8: 217-220. Clarke, J. E G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bulletin of the Brooklyn Entomological Society 36: 149-161. Dietz, W. G. 1910. On Pigritia Clem. Transactions of the American Entomological Society 27: 1O00— 120. Kornerup, A., and J. H. Wanscher. 1978. Methuen Handbook of Colour. 2nd ed. Methuen and Co., Ltd., London. 243 pp. McKelvey, S. D. 1938. Yuccas of the Southwestern United States, Vol. 1. Arnold Arboretum, Jamaica Plain, MA. Pellmyr, O. 1999. A systematic revision of the yucca moths in the Tegeticula yuccasella complex north of Mexico. Systematic Entomology 24: 248-271. Powell, J. A. 1976. The giant blastobasid moths of Yucca (Gelechioidea). Journal of the Lepidopter- ists’ Society 30: 219-229. . 1980. Evolution of larval food preferences in microlepidoptera. Annual Review of Entomology 25: 133-159. . 1984. Biological Interrelationships of Moths and Yucca schottii. University of California Con- tributions in Entomology 100. University of Cal- ifornia Press Ltd., Berkeley, California. vii + 93 pp- Powell, J. A.. and R. A. Mackie. 1966. Biological In- terrelationships of Moths of Yucca whipplei. Uni- versity of California Contributions in Entomology 42. University of California Press Ltd., Berkeley, California. 59 pp. Powell, J. A., C. Mitter, and B. Farrell. 1999. Evolu- tion of larval food preferences in Lepidoptera, pp. 404—422. In Kristensen, N. P., ed. Handbook of Zoology. Lepidoptera, Moths and Butterflies, Vol. 1. Walter de Gruyter, New York. x + 491 pp. Reveal, J. L. 1977. Yucca, pp. 527-536. In Cronquist, A., A. H. Holmgren, N. H. Holmgren, J. L. Re- veal, and P. K. Holmgren, eds. Intermountain Flo- ra, Vol. 6. Columbia University Press, New York. Webber, J. M. 1953. Yuccas of the Southwest. United States Department of Agriculture. Monograph No. 17. Washington, D.C. PROC. ENTOMOL. SOC. WASH 105(2), 2003, pp. 397-406 LUGOIOPS MAYA, A NEW GENUS AND SPECIES OF EPHEMEROPTERA (BAETIDAE) FROM CENTRAL AMERICA W. P. MCCAFFERTY AND D. E. BAUMGARDNER (WPM) Department of Entomology, Purdue University, West Lafayette, IN 47907, U.S.A. (e-mail: pat-:meccafferty@entm.purdue.edu); (DEB) Department of Entomology, Texas A & M University, College Station, TX 77843, U.S.A. (e-mail: dbaumgardner@ tamu.edu) Abstract.—Small minnow mayfly (Ephemeroptera: Baetidae) larvae taken from a moun- tain rivulet in Baja Verapaz, Guatemala are described as Lugotops maya, new genus and new species. The new genus is a member of the Baetodes complex of genera, being more advanced than Moribaetis or Mayobaetis, and sharing numerous synapomorphies with both Prebaetodes and Baetodes, and additional synapomorphies with Baetodes. The pre- cise cladistic relationships deduced for the new genus are discussed and a new key to genera of the Baetodes complex is provided. Larvae of L. maya are striking because of their relatively large size, unique structural characteristics, such as absence of gills 1, and their torrential habitat. Key Words: With the recent discovery of the genus Prebaetodes Lugo-Ortiz and McCafferty in Colombia and Ecuador (Lugo-Ortiz and McCafferty 1996), an essential phylogenet- ic link was found that demonstrated the re- lationship of the highly apotypic Western Hemisphere genus Baetodes Needham and Murphy with more typical baetines, vis-a- vis the Western Hemisphere genera Mori- baetis Waltz and McCafferty and Mayobae- tis Waltz and McCafferty. As a result, the phylogenetic origin of what had been an anomalous genus was hypothesized for the first time, and a distinctive monophyletic group of genera, the Baetodes complex, was delineated in the Western Hemisphere. McCafferty (1998) applied the cladistic re- lationships within the Baetodes complex to biogeographic analysis to show that the or- igin of the Pan-American genus Baetodes was Clearly Neotropical, as was that of oth- er members of the complex. The biogeog- Lugoiops, new genus, new species, Baetidae, Guatemala raphy of this complex was further elucidat- ed by McCafferty (1999). Recently, larvae of relatively large and unusual small minnow mayflies collected from a mountain rivulet in Guatemala in July of 2001 were sent to WPM indepen- dently by W. D. Shepard (Sacramento, Cal- ifornia) and DEB for examination. These larvae proved to be an unknown member of the Baetodes complex of Baetidae, sharing for example, the apicolateral found on the denticulate claws throughout the complex, a thickened labrum also com- mon throughout much of the complex, and absence of the villopore and other charac- teristics associated with other complexes of Baetidae. Study of the character state dis- tribution and cladistic relationships within the Baetodes complex revealed that the spe- cies should not be subsumed by any other genus in the complex and that a new genus was justified within the context of a strictly setal row 398 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON phylogenetic classification (see below). At the same time, the newly discovered lineage provided additional cladistic characteriza- tion that further demonstrated the morpho- logical transition from the more stereotypic minnowlike baetid larval form to the highly apotypic and unusual Baetodes form. More- over, its phenetic distinctiveness makes rec- ognition at the generic level practical. Be- low, we describe the new genus and spe- cies, and provide associated cladistic, di- agnostic, and habitat data. We are honored to name the genus after Carlos Lugo-Ortiz (Cidra, Puerto Rico), who in recent years has contributed the pri- mary thrust in the discovery and elabora- tion of the biodiversity of the family Bae- tidae in the Neotropics as well as the South- ern Hemisphere in general, but especially Africa and Madagascar. The work of Dr. Lugo-Ortiz has provided the basis for fur- ther research on the Baetidae by establish- ing standards of description and analysis, diligently seeking to bring the taxonomy of Baetidae in line with modern systematics that recognizes taxa as discreet inter-related phylogenetic lineages. The gender of the genus is masculine. Lugoiops McCafferty and Baumgardner, new genus (Figs. 1-11) Larva.—Head: Orientation (Figs. 1—2) extremely hypognathous. Antenna (Figs. |— 2) approximately one and one-half times length of head capsule length; scape and pedicel subcylindrical in cross-section, with tufted row of fine, simple setae dorsolater- ally. Lateral margins of frons broadly con- necting to clypeus (Fig. 2). Labrum (Figs. 2-3) basally somewhat constricted, with lateral margins rounded, relatively short, not expanded to nearly round overall shape; dorsal surface with slightly bulbous prom- inence basomedially and two such laterally. Mandible (Figs. 4—5) not continuously con- vex laterally; denticles not fan shaped. La- bial palp (Fig. 8) three-segmented, with segment | and 2 subequal in length and with short, somewhat dome-shaped seg- ment 3. Thorax: Legs (Figs. 1, 9) elongate, outspread from body; tibiae slightly twist- ed. Hindwingpad very small. Abdomen: terga lacking tubercles or patches of setae. Gills present on segments 2-7 (Fig. 1), dor- solaterally oriented but overlapping abdom- inal pleura. Paraprocts (Fig. 11) without marginal denticulation. Median caudal fil- ament reduced to unsegmented conical ves- tige (Fig. 1). Cercus (Fig. 1) bare, lacking medial setae and lateral spination. Type species.—Lugoiops maya, n. sp. Discussion.—Lugoiops larvae are dis- tinctive in many ways, but especially be- cause of their large size along with their elongate, outspread legs, the vestigial me- dian caudal filament, and the absence of gills 1. Edmunds et al. (1976) gave the body length dimensions of Baetodes larvae as 3.0 to 8.0 mm. This size range is also applicable to known Prebaetodes (Lugo- Ortiz and McCafferty 1996). Moribaetis and Mayobaetis larvae, which have gener- ally been considered to be rather large bae- tids have lengths of up to about 10.0 mm. Known Lugoiops larvae by these standards, however, are even larger, being around 12.0-13.0 mm when mature. A modified key to the larval stage of genera of the Bae- todes complex that includes Lugoiops and additional comparative characterization within the complex is included below. With application of the cladistic analysis of Lugo-Ortiz and McCafferty (1996) and character state polarity and distribution pre- sented by them, along with new character assessment and re-analysis herein, we have been able to place Lugoiops within the hy- pothesized cladogram. Demonstrative of its placement in the Baetodes complex, Lu- goiops shares a primary synapomorphy de- fining that complex, i.e., the possession of apicolateral setation on the denticulate claws. Lugoiops also possesses a basome- dial prominence dorsally on the labrum, which Lugo-Ortiz and McCafferty (1996) had originally indicated was an apomorphy VOLUME 105, NUMBER 2 399 Fig. 1. Larval habitus of Lugoiops maya. 400 found only in Moribaetis. However, we have found such a labrum not only in Lu- goiops, but also to a lesser extent in May- obaetis and also in Baetodes, where it can be highly developed in some species, e.g., B. inermis Cohen and Allen. Therefore, that characteristic should no longer be consid- ered a restricted autapomorphy within the Baetodes complex, but likely may very well have been present ancestrally in the com- plex and as such an apomorphy that further helps define this monophyletic grouping of genera. Within the Baetodes complex, Lugoiops shares all of the synapomorphies defining the clade made up of Mayobaetis, Prebae- and Baetodes, i.e., the lack of mar- ginal denticulation on the paraprocts, the dorsolateral setation at the base of the an- tennae, and reduced median caudal _fila- ment. Lugoiops does not share the single apomorphy, involving the unique attenuated frons, that is associated with the relatively plesiotypic genus Moribaetis within the complex. todes, Lugoiops does not possess any of the au- tapomorphies that are identified with the in- dividual Mayobaetis lineage, i.e., lateral spination on the cerci and dorsoventrally flattened antennal scapes and pedicels. On the other hand, it does share synapomor- phies previously shown to be common to Prebaetodes and Baetodes, including an even shorter median caudal filament, elon- gate legs, and the slightly twisted tibiae. Lugoiops does not share any of the apo- morphies that Lugo-Ortiz and McCafferty (1996) used to distinguish the individual Prebaetodes lineage, having neither the lat- erally convex mandibles nor expanded and nearly round labrum that are autapomorphic in Prebaetodes. Also, based on the cladistic analysis of Lugo-Ortiz and McCafferty (1996), except for an even shorter median caudal filament, Lugoiops does not share apomorphies that distinguish the individual Baetodes lineage, i.e., the apomorphic ab- dominal gill arrangement (1—5) or the dor- sal abdominal armature. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON A strong indication of the sister relation- ship of Lugoiops within this apotypic Lu- goliops-Prebaetodes-Baetodes clade, how- ever, is demonstrated by at least one com- pelling character state that we are able to add to the analysis, that is the complete loss of medial setae (swimming hairs) on the cercl of both Lugoiops and Baetodes. The loss of gills 1 in Lugoiops is an au- tapomorphy within the complex. Another characteristic possibly unique to the Lu- goiops lineage includes the reduction of the median caudal filament to a minute, unseg- mented, conical vestige. However, we can- not be absolutely sure such a middle tail vestige 1s unique to Lugoiops, because al- though considerable material of Baetodes that we have studied consistently have quite short (always less than one-fifth the cercus length) but segmented middle tails, Ed- munds et al. (1976) stated that the middle tails of Baetodes were reduced to stubs, and thus that might indicate that they had seen specimens with unsegmented vestiges as we have observed in Lugoiops. Regardless of the fine degree of reduc- tion of the median caudal filament, Prebae- todes and Baetodes could also be consid- ered two-tailed in the vernacular along with Lugoiops. Certain two-tailed, rheophilic baetid larvae throughout the world also have developed tergal tubercles or setal patches (e.g., Lugo-Ortiz and McCatferty 1999, Lugo-Ortiz et al. 2001, Jacobus and McCafferty 2001). Although this is also the case in Baetodes, there is no indication of such development in Prebaetodes or Lu- golops. In Moribaetis, Mayobaetis, and Prebae- todes, the gills are held outspread from the abdomen, or if appressed are oriented dor- sal of the abdominal pleura. In Lugoiops and many Baetodes (e.g., see abdominal il- lustrations in Cohen and Allen [1978]), the gills are held appressed to the abdomen more or less over the pleura. Also in Bae- todes, the gills are often held out from the abdomen ventral of the abdominal pleura, thus leading to the standard descriptor of VOLUME 105, NUMBER 2 401 rf Figs. 2-8. Lugoiops maya, larva. 2, Head (frontal). 3, Labrum (dorsal). 4, Angulate mandible. 5, Planate mandible. 6, Hypopharynx. 7, Maxilla. 8, Labium. 402 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9-11. Baetodes larvae as having ventral gills. From these observations, it appears that lat- erally appressed gills, especially as seen in the more posterior gills of Lugoiops, may have been the condition in the most im- mediate common ancestor to the Lugoiops- Baetodes clade. This orientation developed into an extreme ventral orientation within Baetodes. Hindwings are not present in Baetodes alates; however, hindwingpads can be either vestigial or entirely lacking in the larvae of Baetodes, which indeed are also intergener- ically variable with respect to the degree of labral thickening and degree of ventral ori- entation of the gills, as mentioned above. The hindwingpad variable conditions are not without precedence among other bae- tids, even within the same species, as Is il- lustrated by the North American Acentrella Lugoiops maya, larva. 9, Foreleg (anterior face). 10, Claw. 11, Paraproct. turbida (McDunnough), where in some lar- vae there are vestigial hindwingpads and in others there are none (see McCafferty et al. 1994). It is not all together clear from the very small size of the hindwingpads wheth- er the alates of Lugoiops will lack hind- wings or have only minute hindwings. In either case, this is an unusual instance among Ephemeroptera, where a large may- fly would have this amount of reduction in hindwings, because such reduction has usu- ally been regarded to be associated with body size reductionism in general (Mc- Cafferty and Waltz 1990). Another atypical example of such hindwing reduction in a large alate mayfly is represented by the very small hindwings found in the large Dipteromimus McLachlan mayflies (Pisci- forma: Dipteromimidae) in Japan. If the hindwings are indeed present in the alates VOLUME 105, NUMBER 2 1 Fig. 12. 403 Moribaetis Mayobaetis Prebaetodes Lugotops Baetodes Derived cladogram of the Baetodes complex. Evidential apomorphies (see text also): 1—frons attenuate; 2—loss of paraproct marginal denticulation, basal antennal setation developed, some shortening of middle tail; 3 reduction of middle tail setation, elongation of legs, slight twist of tibiae; 5 lateral cerci spines present, scape and pedicel depressed; 4 additional shortening of middle tail, laterally convex mandibles, ex- panded labrum; 6—cerci bare, middle tail more shortened, gill not outspread or held dorsal to abdominal pleura; 7—gills 1 lost, middle tail unsegmented; 8 armature present on middle abdominal terga. of Lugoiops, then the complete loss of the alate hindwings in Prebaetodes and Bae- todes, could be seen as a convergence re- lated to size reduction in those two lineag- es, because other members of the complex are all relatively good sized baetids. Such hindwing subject to homoplasy throughout many baetid clades. Lugo-Ortiz and McCafferty (1996) noted only antennal pedicel setation in the cladis- tic character state distributions among the Baetodes complex, and, as indicated above, its commonality in Mayobaetis, Prebaeto- des, and Baetodes (to which can now can be added Lugoiops). In actuality, Prebae- todes and Baetodes have antennal base se- tation essentially confined to the pedicel, whereas Mayobaetis and Lugoiops have such setation also well developed on the scape. Possible explanations of this, given the preponderance of cladistic evidence stated here, is that the common ancestor of the Mayobaetis-Prebaetodes-Lugoiops- loss 1s gills 6 and 7 lost, gills can be held ventral of abdominal pleura, Baetodes clade possessed setation on both the pedicel and scape and it became inde- pendently reduced on the scapes of Prebae- todes and Baetodes, or alternatively, that it developed independently on the scapes of Mayobaetis and Lugoiops. Because there is no knowledge of any function related to the various arrangements, we are left with no clue as to which alternative may be more probable. It is certainly possible that further review of the speciose genus Baetodes may reveal some variation of these two arrange- ments within that genus also. From the above review and analysis of cladistics, Lugoiops falls as a sister genus of Baetodes, which together are opposite Prebaetodes within this advanced clade of the Baetodes complex. Our entire hypoth- esized cladogram can be seen in Fig. 12. Sequencing conventions allow each of the sequenced lineages as described above to be recognized at the same taxonomic lev- el—in this case the genus level. 404 KEY TO THE LARVAE OF GENERA OF THE WESTERN HEMISPHERE BAETODES COMPLEX 1. Tarsal claws with 1—5 or more setae on api- colateral aspect of denticulate claws (Fig. 10; figs. 9, 15, 32, 39 [Lugo-Ortiz and McCafferty 1996]); villopore absent; labrum often raised over dorsal surface into three broad mounds or at least thickened basomedially (Fig. 2: figs. 2, 5 [Lugo-Ortiz and McCafferty 1996]) ... . Ru ate 30.2 Fs 20-2 3, a: evap suteat Baetodes complex, 2 — Tarsal claws without setae as above; villopore present or absent; labrum not thickened as ADON Gg ar vace other Western Hemisphere Baetidae . Median caudal filament over one-half length of cerci (fig. 10 [Lugo-Ortiz and McCafferty 1996]); antennal pedicels lacking rows of tufts of setae in dorsal or lateral aspect (fig. 7 [Lugo- Ortiz and McCafferty 1996]; figs. 2-5 [Waltz and McCafferty 1985]); hindwingpads relative- ly well developed ............... Moribaetis — Median caudal filament less than one-half length of cerci (Fig. 1; figs. 16, 37, 41 [Lugo- Ortiz and McCafferty 1996]); antennal pedicels with row or tuft of setae dorsolaterally (Fig. 2; figs. 12, 25, 38 [Lugo-Ortiz and McCafferty 1996]); hindwingpads variable or absent. ... 3 . Gills present on abdominal segments |—5 (figs. 9-24 [Cohen and Allen 1978]: fig. 40 [Lugo- Ortiz and McCafferty 1996]), ventral or ap- pressed laterally; middle abdominal terga with tubercles! ‘or'setallpatches:*t-yesy-pess =... = Baetodes — Gills present on abdominal segments 1—7 or 2— 7 (Fig. 1; fig. 12 [Waltz and McCafferty 1985]; fig. 33 [Lugo-Ortiz and McCafferty 1996]), outspread, dorsal, dorsolaterally or laterally ap- te tad pressed; abdominal terga without tubercles or setal patches 4. Cerci without medial setae (Fig. 1); gills pre- sent on abdominal segments 2—7 (Fig. 1); me- dian caudal filament reduced to unsegmented Vestiger (Rigs) kiss srremsren eck: MUM et aie Lugoiops — Cerci with medial setae (figs. 16, 37 [Lugo- Ortiz and McCafferty 1996]): gills present on abdominal segments 1—7 (gills 1 may be very small) (fig. 12 [Waltz and McCafferty 1985]: fig. 33 [Lugo-Ortiz and McCafferty 1996]): median caudal filament multisegmented short tail (figs. 16, 37 [Lugo-Ortiz and McCafferty VSG) fete, cs 5. Sect ay cays vansyel oy swans. ecet tops Cameeeute ns 5. Legs slender-elongate (fig. 31 [Lugo-Ortiz and McCafferty 1996]): hindwingpads vestigial: la- brum large and near round (fig. 25 [Lugo-Ortiz and McCafferty 1996]); median caudal fila- ment with lateral setae reduced (fig. 37 [Lugo- Ortiz and McCafferty 1996]) ..... Prebaetodes — Legs robust (fig. 14 [Lugo-Ortiz and Mc- Cafferty 1996]); hindwingpads relatively well developed; labrum shorter, somewhat quadrate in PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (fig. 12 [Lugo-Ortiz and McCafferty 1996]): median caudal filament with lateral setae de- veloped (fig. 16 [Lugo-Ortiz and McCafferty OO GI) Woes ncnscecueue en, omens tore Mayobaetis Lugoiops maya McCafferty and Baumgardner, new species (Figs. 1-11) Larva.—Mature body (Fig. 1) length 12.0 mm; cercus length 14.5 mm; antenna length 3.0 mm. Head: Color brown; frons (Fig. 2) with somewhat diffuse white spot centrally; white areas also between each compound eye and lateral ocellus; lateral branches of frontal suture below but not contacting lateral ocelli. Antennal scape and pedicel (Fig. 2) brown, with white mem- branous sub-base exposed. Short labrum (Figs. 2—3) slightly emarginate, with thick brush of short marginal setae extending more sparsely along rounded lateral mar- gins. Mandibles shaped as in Figs. 4 and 5, with denticles extremely fused; angulate mandible (Fig. 4+) with mola nearly perpen- dicular to long axis of body of mandible, and with prostheca with only slight branch- ing; prostheca of planate mandible (Fig. 5) long, double and with branching setules in distal two-thirds. Hypopharynx (Fig. 6) se- tate distally, with lingua narrowly rounded distally, and with superlinguae overlapping bilobular distally. Maxillary palp (Fig. 7) two-segmented, short, extending about as far as nearly conical galealacinae. Labium (Fig. 8) with glossa reduced; paraglossa well developed, with strong apical tuft of brushing setae; labial palp with segment 2 with minute medioapical thumb, and seg- ment 3 with short marginal setae and sparse, small submarginal spines. Thorax: Nota (Fig. 1) brown with lighter brown stri- ations and patches; pleura brown with ex- tensive white inter-sclerite areas; sterna white. Legs (Figs. 1, 9) light brown with some markings anteriorly and cream con- colorous posteriorly; anterior face of fem- ora (Fig. 9) with whitish swordlike mark extending medially from base to somewhat over one-half length of femora, and with VOLUME 105, NUMBER 2 darker diffuse elongate marking (near dor- sal edge and dorsad of swordlike mark) ex- tending in middle two-thirds of length of femora; ventral tip of femora overlapping strongly recurved dorsal tip (Fig. 9); thick row of medium sized setae extending along dorsal edge of femora and extending along edge of tibiae and tarsi (Fig. 9): tarsi also with scattered short, stout setae and longer subapical bristle on surface opposite inward bend of tarsal claws, and with tufts of fine, hairlike setae apically and along on outer face (Fig. 10). Tarsal claws (Fig. 10) with five to eight denticles, most developed api- cally, and with five lateroapical, blond setae at base of curved claw tip. Abdomen: Ab- dominal terga light brown with dark brown markings as in Fig. 1, only terga 9 and 10 concolorous medium brown (lighter in younger larvae); terga 3 most darkly marked and tergum 5 least darkly marked. Sterna cream to yellow-brown, without dis- tinctive markings. Paraproct (Fig. 11) with irregular submarginal denticulation. Material examined.—Holotype: Mature larva, Guatemala, Baja Verapaz, unnamed stream, 0.3 km south of La Cumbre, 1,274 m elevation, 15/00/34N 90/13/S51W, VI-12- 2001, W. D. Shepard, fluid preserved in Purdue Entomological Research Collection (PERC), West Lafayette, Indiana. Para- types: One nearly mature larva, same data and deposition as holotype; one nearly ma- ture larva, same data, except collected by DEB and deposited at Texas A & M Uni- versity (TAMU), College Station, TX. Oth- er material: four larvae with some parts slide-mounted, same data, at PERC; two larvae, same data, at TAMU; and two lar- vae at Laboratorio Entomologia Systemati- ca, Univerisdad del Valle de Guatemala, Guatemala City. Etymology.—The specific epithet is a noun in apposition, after the Maya Native Americans, which lived in southern Mexico and northern Central America, including the Guatemalan region. Discussion.—Lugoiops maya was col- lected from a torrential mountain rivulet of 405 only a few centimeters in width and depth that was coursing down a deep and narrow- sided crevice at about a 30° grade. Larvae were clinging to rocky substrate. Other mayfly species collected at this site were also members of the Baetodes complex, in- cluding B. noventus Cohen and Allen, B. deludens Lugo-Ortiz and McCafferty, and Mayobaetis ellenae (Mayo). The former two were also found clinging in the rapid current, but M. ellenae was taken in the splash zone along the side of the stream. Lugo-Ortiz and McCafferty (1996) indicat- ed that the more plesiotypic members of the Baetodes complex were often found in moderate current and splash zone environ- ments in the Neotropics. Observations of splash-zone habitation had been made pre- viously by WPM in Costa Rica for Mori- baetis maculipennis (Flowers). Lugo-Ortiz and McCafferty (1996) also noted that Bae- todes and Prebaetodes were, however, known as clingers in current. To this latter adaptive grouping, can now be added Lu- goiops. Lugo-Ortiz and McCafferty (1996) showed correlation between habitat and morphological transitions within the Bae- todes complex. Based on this, the new data we present for Lugoiops would have been to a large extent predictable. ACKNOWLEDGMENTS We thank Arwin Provonsha (West Lafay- ette, IN) for the illustrations used herein, Pat Randolph and Lu Sun (West Lafayette, IN) for technical assistance, and Bill Shep- ard (Sacramento, CA) for contributing specimens. Mike Meyer and Pat Randolph (West Lafayette, IN) commented on early drafts of the manuscript. Jack Schuster and Enio Cano (Guatemala City, Guatemala) helped with collecting permits. LITERATURE CITED Cohen, S. D. and R. K. Allen. 1978. Generic revisions of mayfly nymphs III. Baetodes in North and Cen- tral America (Baetidae). Journal of the Kansas En- tomological Society 51: 253-269. Edmunds, G. EF, Jr, S. L. Jensen, and L. Berner. 1976. 406 Mayflies of North and Central America. Univer- sity Minnesota Press, Minneapolis. 330 pp. Jacobus, L. M. and W. P. McCafferty. 2001. Contri- bution to the systematics of Afrobaetodes Demou- lin (Ephemeroptera: Baetidae). African Entomol- ogy 9: 97-103. Lugo-Ortiz, C. R. and W. P. McCafferty. 1996. Phy- logeny and classification of the Baetodes complex (Ephemeroptera: Baetidae), with description of a new genus. Journal of the North American Ben- thological Society 15: 367-380. . 1999. A new genus of small minnow mayflies from New Guinea and New Britain. Annales de Limnologie 35: 57—70. Lugo-Ortiz, C. R., H. M. Barber-James, W. P. Mc- Cafferty, and R. C. deMoor. 2001. A non-para- phyletic classification of the Afrotropical genus Acanthiops Waltz and McCafferty (Ephemerop- tera: Baetidae). African Entomology 9: I-15. McCafferty. W. P. 1998. Ephemeroptera and the great PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON American interchange. Journal of the North Amer- ican Benthological Society 17: 1—20. . 1999. Biodiversity and biogeography: exam- ples from global studies of Ephemeroptera. Pro- ceedings of the Symposium on Nature Conserva- tion and Entomology in the 21 Century, Ento- mological Society of Korea, Chonan, Korea 1999: 3-22. McCafferty, W. P. and R. D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera) of North and Middle America. Transactions of the American Entomological Society 116: 769-799. McCafferty, W. P., M. J. Wigle, and R. D. Waltz. 1994. Systematics and biology of Acentrella turbida (Ephemeroptera: Baetidae). Pan-Pacific Entomol- ogist 70: 301-308. Waltz, R. D. and W. P. McCafferty. 1985. Moribaetis: a new genus of Neotropical Baetidae (Ephemer- optera). Proceedings of the Entomological Society of Washington 87: 239-251. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 407—424 A REVIEW OF THE NEOTROPICAL SPECIES OF BRACHYDEUTERA LOEW (DIPTERA: EPHYDRIDAE) WAYNE N. MATHIS AND ISAAC S. WINKLER (WNM) Department of Systematic Biology, Entomology Section, National Museum of Natural History, Smithsonian Institution, PO. Box 37012, MRC-169, Washington, DC 20013-7012, U.S.A. (e-mail: mathis.wayne@nmnh.si.edu); (ISW) Department of Ento- mology, University of Maryland, College Park, MD 20742-4454, U.S.A. (e-mail: isw @ wam.umd.edu) Abstract.—The Neotropical species of the genus Brachydeutera Loew are reviewed, including description of a new species, B. dentata (Paraguay: Parque Nacional Ybyqui [26°05'S, 56°48'W]). The extraordinary vagility of B. longipes Hendel is documented, having recently been discovered in South America as far south as Bolivia. Another pri- marily Neotropical species, B. neotropica Wirth, demonstrates slight but consistent vari- ation that is described and illustrated. Tools for identification (keys, illustrations, short diagnoses) as well as maps are provided. Key Words: cies, Neotropical Region Specimens of Brachydeutera Loew are most commonly associated with lentic aquatic habitats, and adults are frequently encountered on the surfaces of small, stag- nant pools or in ditches where the water is largely undisturbed (Williams 1938, Foote 1995, Keiper and Walton 2000). The habi- tats may be ephemeral and often they are slightly to highly polluted (Venkatesh 1976). Some species are associated with coastal marine habitats, including mangrove swamps (Foote 1995). Adults of Brachy- deutera are easily recognized among shore flies because of their conspicuously pecti- nate arista, projected and carinate face, la- teroclinate fronto-orbital setae, and the shortened costal vein (extending to vein R,,;) and vein R,,;. In addition, the thorax is often essentially two toned with the me- sonotum being darker, usually brown, and the pleural areas whitish gray to gray. Spec- imens are comparatively large for shore Review, Diptera, Ephydridae, Dagini, Brachydeutera, B. dentata new spe- flies, with body lengths of approximately 2-5 mm. So far as we know, the larvae are highly polyphagous on microbial, algal, and detrital materials (Foote 1995, Keiper and Walton 2000). Various throughout most temperate and tropical re- gions (Mathis and Zatwarnicki 1995), and worldwide there are 15 species with greater diversity in the Old World tropics, espe- cially the Afrotropical Region. The purpose of this paper is to review the Neotropical fauna, including description of a new species, B. dentata, from Argentina and Paraguay. Counting this new species, there are now five species known from the neotropics and 16 species worldwide (Mathis and Zatwarnicki 1995). One of these species, B. longipes Hendel, is adven- tive to the New World from Asia, and we document herein the tremendous vagility of this species, which was first reported from the East Coast of North America (Mathis species occur 408 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and Steiner 1986) and which has now been found throughout the Caribbean and Cen- tral America and in South America from Venezuela south to Bolivia. Brachydeutera had previously been clas- sified in: (1) the subfamily Napaeinae Cres- son (Sturtevant and Wheeler 1954), (2) the tribe Hyadinini Philips et al. of the subfam- ily Parydrinae Wirth and Stone (Wirth 1965, 1968, Cogan and Wirth 1977, Cogan 1980), (3) the tribe Parydrini of the subfam- ily Parydrinae (Cogan 1984, Mathis 1989), and (4) most recently in the subfamily Ephydrinae Zetterstedt, tribe Dagini Mathis (Zatwarnicki 1992, Mathis and Zatwarnicki 1995). Here we continue to advocate place- ment of Brachydeutera in the tribe Dagini of the subfamily Ephydrinae. Keiper and Walton (2000) provided evidence from feeding habits to support placement of Brachydeutera in Ephydrinae but suggested a separate tribe for the genus, given its unique and opportunistic feeding strategy as a generalist on a variety of algal and mi- crobial taxa. We suggest, however, that its generalized polyphagy is probably a ple- siomorphic condition and thus prefer to maintain the genus in Dagini, perhaps as a lineage near the base of this tribe. Like oth- er tribes and genera of Ephydrinae, includ- ing Dagini, the fronto-orbital setae are la- teroclinate. In accordance with this classi- fication and to facilitate better recognition of Brachydeutera, we have provided a key to New World genera of Dagini. We also recharacterize Brachydeutera within the context of this tribe and subfamily. METHODS AND MATERIALS The descriptive terminology, with the ex- ceptions noted in Mathis (1986) and Mathis and Zatwarnicki (1990a), follows that pub- lished in the Manual of Nearctic Diptera (McAlpine 1981). Because specimens are small, frequently than 3.5 mm in length, study and illustration of the male terminalia required use of a compound mi- less croscope. We have followed the terminol- ogy for most structures of the male termin- alia that other workers in Ephydridae have used (see references in Mathis 1986, and Mathis and Zatwarnicki 1990a, b), such as surstylus. Zatwarnicki (1996) suggested that the pre- and postsurstylus correspond with the pre- and postgonostylus and that the subepandrial plate is the same as the medandrium. The terminology for struc- tures of the male terminalia is provided di- rectly on Figs. 3-5. The new species de- scription is composite and not based solely on the holotype. One head and two vena- tional ratios that are used in the descriptions are defined below. All ratios are based on three specimens, representing the largest, smallest, and one intermediate. Gena-to-eye ratio is the genal height measured at the maximum eye height divided by the eye height. Costal vein ratio is the straight line distance between the apices of R,,, and R,,; divided by the distance between the apices of R, and R,,;. M vein ratio is the straight line distance along vein M between crossveins (dm-cu and rm) divided by the distance apicad of dm-cu. Although many specimens for this study are in the National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. (USNM), we also borrowed and stud- ied numerous specimens that are deposited in the following museums: ANSP Academy of Natural Sciences of Philadelphia, Pennsylvania, USA. American Museum of Natural His- tory, New York, USA. The Natural History Museum (for- merly the British Museum (Natural History)), London, England, Unit- ed Kingdom. Canadian National Collection, Ot- tawa, Canada. Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA. Naturhistorisches Museum, Wien, AMNH BMNH CNC MCZ NMW Austria. VOLUME 105, NUMBER 2 SYSTEMATICS Tribe Dagini Mathis Dagini Mathis 1982: 3. Type genus: Dagus Cresson 1935. Key TO THE NEW WoRLD GENERA AND SPECIES GROUPS OF DAGINI 1. Costal vein short, extending only to vein R,.<: face with distinct, mediovertical tended from ptilinal suture to oral margin; aris- carina, ex- ta distinctly pectinate, longest rays subequal to length of Ist flagellomere ... Brachydeutera Loew — Costal vein long, extending to vein M; face evenly transversely arched, not carinate; arista essentially bare to subpectinate with longest rays not more than 2 length of Ist flagellomere ... 2 Ventral 2—*4 of face distinctly projected ante- tv riorly, arched transversely and vertically (best seen in profile); fronto-orbital setae 3, anterior seta smaller; pulvilli lacking: postpronotum with | to a few setulae Dagus Cresson — Face shieldlike, broadly but more shallowly projected over most of height; usually 1—2 fronto-orbital setae: pulvilli present, conspicu- ous; postpronotumiibares. 22% ...-.2 520+. 3 3. Gena high, equal to or greater than eye height; genal seta well developed and conspicuous: prescutellar acrostichal setae well developed; propleuron setulose; Sth tarsomere with dor- soapical process extended beyond base of tar- Saliclawsy <2 4.2 ...4 Diedrops Mathis and Wirth — Gena short, usually not more than % eye height; genal seta, if present, weakly developed and inconspicuous; prescutellar acrostichal se- tae not evident; propleuron lacking setulae; Sth tarsomere not as above (Physemops) 4. Halter capitellum black; ocellar setae lacking; arista long, over twice combined length of first 3 antennal segments; vein CuA, along poste- rior margin of discal cell bowed posteriorly epude) late het chs ilete,.e.:6%,0) eet sie cio qe the Nemorosus group — Halter capitellum pale, usually yellowish; ocel- lar setae present, conspicuous; arista shorter, rarely not over twice combined length of first 3 antennal segments; vein CuA, along poste- rior margin of discal cell straight ....... Biba otras) te car eee eT fy ees OLS PRT the panops group Genus Brachydeutera Loew Brachydeutera Loew 1862: 162. Type spe- cies: Brachydeutera dimidiata Loew 1862 (= Notiphila argentata Walker 1853), monotypy.—Williston 1908: 309 [key].—Hennig 1952: 349 [immature stages, key].—Wirth 1964: 3-12 [revi- 409 sion]; 1968: 20 [Neotropical catalog].— Mathis 1983: 177-180 [review Nearctic fauna].— Mathis and Steiner 1986: 56— 61 [key to Nearctic species with addition of B. longipes Hendel].—Lizarralde de Grosso 1989: 46—47 [review, Argenti- na].—Mathis and Zatwarnicki 1995: 229-232 [world catalog]. Description.—Small to large shore flies, body length 1.45—5.35 mm. Head: Frons much wider than long, mi- crotomentose, appearing dull, mesofrons not distinguished from remainder of frons; lateroclinate fronto-orbital setae 2—3; both inner and outer vertical setae present; | pair of cruciate, interfrontal setae inserted in front and laterad of anterior ocellus; ocelli arranged in isosceles triangle, distance be- tween anterior ocellus and posterior ocelli less than that between posterior ocelli; ocel- lar setae well developed, proclinate and di- vergent; pedicel lacking dorsoapical, prom- inent seta; arista conspicuously pectinate, bearing 6—12 dorsally branching rays, lon- ger rays subequal to length of Ist flagello- mere. Face generally bare except for 3—12 setulae laterally; face with prominent ver- tical, median carina between antennal grooves, extending from ptilinal suture to epistome; epistomal margin of face broadly emarginate medially, emargination shallow- ly rounded: clypeus conspicuous within ep- istomal emargination as a wide, transverse band. Eye bare, large, prominent, slightly oval vertically. Genal seta lacking or much reduced. Thorax: Chaetotaxy generally weakly developed; postpronotal, presutural, supra- alar, and prescutellar acrostichal setae re- duced but usually evident; posterolateral dorsocentral and postalar setae better de- veloped; otherwise macrochaetotaxy of scutum reduced; acrostichal and dorsocen- tral tracks represented by reduced setae: scutellum setose dorsally, with a basal and apical seta, these not arising from tubercles: notopleural setae usually 2, anterior seta weakly developed; anepisternum with | 410 seta; katepisternum usually with | seta, fre- quently pale, inconspicuous, less well de- veloped than anepisternal seta; prosternum bare of setulae. Costal vein short, extending to vein R,,;; vein R,,, short, not extending beyond level of crossvein dm-cu; vein M apicad of crossvein dm-cu evanescent. Legs generally lacking prominent setae; forebas- itarsomere short, subequal in length to other tarsomeres; mid- and hindbasitarsomeres longer, about twice length of other tarso- meres; claws with curvature more abrupt apically; pulvilli evident but poorly devel- oped. Abdomen: Flattened dorsoventrally in fe- male; convex dorsally in male; 5th tergite of male elongate, length subequal to ter- gite’s basal width. Male terminalia as fol- lows: epandrium attenuate dorsally, not ex- tended around cerci to form cercal cavity, extended ventromedially (perhaps with fused surstyli) to form a narrow, ventral plate posteroventral to cerci (the surstyli are ventral extensions of the epandrium and may not be homologous with the gonosty- li); postsurstylus and subepandrial sclerite lacking; hypandrium attached basally at basal % of surstyli; pre- and postgonites fused to each other and to hypandrium as digitiform, well-sclerotized processes; ae- deagus poorly sclerotized, as a conical or keel-shaped, semihyaline lobe or protuber- ance; ejaculatory apodeme apparently lack- ing. Discussion.—Although the phylogenetic relationships between Brachydeutera and other genera within the subfamily Ephydri- nae are not resolved, there is considerable evidence supporting the monophyly of the genus. Morphological evidence, mostly au- tapomorphies, is as follows: 1. arista con- spicuously pectinate. 2. face, especially at oral margin, projected. 3. face vertically carinate. 4. fronto-orbital setae lateroclina- te. 5. costal vein short, extending to vein R,.;. 6. vein R,,, short. 7. male epandrium attenuate dorsally, not extending dorsally around and above the cerci to form a cercal cavity. 8. epandrium extended ventromedi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ally (perhaps including fused surstyli) to form a narrow, ventral plate, posteroventral to cercl. Key TO NEOTROPICAL SPECIES OF BRACHYDEUTERA 1. Brown color of anepisternum gradually becom- ing paler ventrally, merging with pale gray . . B. longipes Hendel — Brown color of anepisternum continued ven- trally to about dorsal 4—', thereafter sharply delimited from pale gray coloration on ventral pleuraliiresionie.. ~ = ssc ci cia) a eek ee 2 2. Facial carina low, bluntly rounded, especially ventrally; male terminalia as in Figs. 17-18 Pee Se te RA B. sturtevanti Wirth — Facial carina high, sharply defined, especially ventrally 3. Gonite spatulate apically ... B. neotropica Wirth — Gonite digitiform or tapered toward apex ... 4 4. Apparent merger of fused surstyli with epan- drium in lateral view indicated by an angulate emargination; gonite tapered toward apex, bearing well developed, stout, spinelike setae apically (Fig. 3) B. dentata, new species — Apparent merger of fused surstyli with epan- drium in lateral view broadly rounded; gonite digitiform apically, narrowly rounded, some- times with stronger setae apically but not stout, spinelike; (Pigs [ere nyse B. argentata (Walker) Brachydeutera argentata (Walker) (Figs. 1-2) Notiphila argentata Walker 1853: 407 [USA; LT (sex ?; designated by Mathis 1983: 178), BMNH]. Brachydeutera argentata: Osten Sacken 1878: 203 [generic combination].— Crawford 1914: 37 [parasitoid: Cyrtogas- ter glasgowi Crawford (= C. trypherus (Walker), Pteromalidae)].—Sturtevant and Wheeler 1954: 204—205 [review, dis- tribution].—Wirth 1964: 5-6 [revi- sion].—Scheiring and Foote 1973: 163 {natural history].—Mathis 1983: 179 [figures of male terminalia]—Woodley and Hilburn 1994: 29 [list, Bermuda].— Mathis and Zatwarnicki 1995: 230 [world catalog]. Brachydeutera dimidiata Loew 1862: 163 [USA. District of Columbia: **Washing- ton”; LT 2 (designated by Mathis 1983: VOLUME 105, NUMBER 2 Structures of the male terminalia of Brachydeutera argentata. 1, Cerci, epandrium (+ fused surstyli ?), gonite, and aedeagus, lateral view. 2, Figs. 1-2. Same, posterior view. 178), MCZ].—Osten Sacken 1878: 203 [synonymy]. Diagnosis.—This species is distinguished from congeners by the following combina- tion of characters: facial carina high, sharp- ly defined, especially ventrally; brown color of anepisternum continued ventrally to about dorsal ¥.—/,, thereafter sharply delim- ited from pale gray coloration on ventral pleural region; apparent merger of fused surstyli with epandrium in lateral view broadly rounded (Fig. 1); gonite digitiform apically, narrowly rounded (Fig. 2). Specimens examined.—BERMUDA. Hamilton (swamp near), Dec, W. S. Brooks @iexe MCZ): Distribution.—Nearctic: Bermuda, Can- ada (QC), USA (FL, IA, MI, MA, ME, NC, NJ OH; TX): Remarks.—We have included this spe- cies in this review because of its occurrence on Bermuda, which is often omitted from other faunal reviews. Although having a 411 Neotropical component, the fauna of Ber- muda is mostly temperate and is especially influenced by the southeast Nearctic Region (Woodley and Hilburn 1994). Brachydeutera dentata Mathis and Winkler, new species (Figs. 3-5) Description.—Moderately small to me- dium-sized shore flies, body length 2.5—3.3 mm. Head: Frons uniformly brown except for olivaceous to greenish areas laterad of ocel- li; 3 lateroclinate fronto-orbital setae, anter- iormost seta weaker, /,—7; length of poste- rior setae. Antenna and ridge of facial ca- rina brown, concolorous with frons. Aristal rays 9-11. Facial carina prominent, narrow and pointed in holotype male, wider and more blunt in paratype females. Clypeus and face silvery white, except for ridge of carina and narrow margin along oral emar- gination. Face with 4—5 setulae laterally. Palpus pale, yellowish. Thorax: Mesonotal chaetotaxy poorly developed; setae barely evident, setae of main setal tracks small. Mesonotum mostly brown, with olivaceous stripes laterad of the acrostichal track, laterad of dorsocentral track, near dorsal margin of notopleuron, in supra-alar area, and at merger of scutum and scutellum. Scutellar ratio 0.71; distance between apical scutellar setae less than be- tween basolateral scutellar seta and apical seta. Anterior notopleural seta present, weaker than posterior seta; katepisternal seta present. Brownish coloration of meso- notum continued ventrally to dorsal , of anepisternum, thereafter abruptly delimited from grayish to silvery-white coloration of ventral portion of pleural areas; anepister- num with posterior margin silvery gray. Wing hyaline; vein R,,, slightly to moder- ately arched; vein R,,; noticeably arched; costal vein ratio 2.8—3.4; M vein ratio 0.58— 0.60. Femora mostly yellowish, with slight- ly darker patch apically on dorsal surface; tibiae yellowish basally, becoming gradu- ally darker apically; tarsomeres brown. Or. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cercus ) * HR epandrium — aedeagus gonite - / Figs. 3-5. Structures of the male terminalia of Brachydeutera dentata. 3, Cerci, epandrium (+ fused surstyli ?), gonite, and aedeagus, lateral view. 4, Left gonite, lateral view. 5, Same as 4, posterior view. Abdomen: Dorsum with posterolateral, grayish areas on tergites 2—5 of males and 2—6 of females. Terminalia of male are as follows (Figs. 3—5): dorsal surface of epan- drium in posterior view shallowly concave: epandrial width at dorsum only slightly wider than lateral margins of cerci, parallel sided; epandrium + surstyli (Fig. 3) elon- gate, narrow in ventral view, sides subpar- allel, widest in the middle, and rounded api- cally; in lateral view, apex of epandrium + surstyli bluntly rounded with several short setae present apically on dorso-anterior sur- face, parallel sided on distal /,, forming an- gulate concavity at base of fused surstyli; aedeagus in ventral view angulate, pointed distally, with rounded ventral lobes basally: gonite (Figs. 4—5) subrectangular basally, posteroventral angle drawn out into digiti- form process which is strongly procurved, narrowing apically, with 1—2 strong apical spurs and a prominent dorsal seta. Type material—Holotype ¢ is labeled “PARAGUAY. Parag.: Parg. Nac. Ybyquf (26°05’S, 56°48°W) 15 Mar. 1986/HOLO- TYPE <6 Brachydeutera dentata W. N. Mathis & Winkler USNM [red; species name, gender symbol, and “& Winkler” 3 handwritten].’"> The holotype is double mounted (glued to a paper triangle), is in good condition (wings flexed down), and is deposited in the USNM. Twelve paratypes (3 6, 9 2; USNM) bear the same locality data as the holotype. Other paratypes are from: ARGENTINA. Buenos Aires: Punta Lara (8 km N Villa Elisa), 4 Dec 1979, C. M. and O. S. Flint, Jr. (1 d, 2 2; USNM). Distribution (Fig. 6).—Neotropical: Ar- gentina (Buenos Aires), Paraguay (Para- guay). Etymology.—The specific epithet, den- tata, is of Latin derivation and refers to the toothlike setae at the apex of the gonite. Remarks.—Externally this species is nearly identical to three congeners: B. ar- gentata (eastern North America), B. ibari Ninomyia (Europe and northern Asia), and B. neotropica (Florida south to Argentina). Of these species, however, only the range of B. neotropica includes Argentina. Struc- tures of the male terminalia of B. dentata differ significantly from other Brachydeu- tera and easily distinguish it. The gonite is tapered gradually, more so apically, and has distinctive, stout, spurlike setae at the apex. In addition, the ventromedial extension of VOLUME 105, NUMBER 2 413 Fig. 6. Distribution map for Brachydeutera dentata (filled circles) and B. longipes (filled squares). 414 the epandrium (? and fused surstyli) is nar- rower and more elongate than in similar congeners. In contrast, B. neotropica has a spatulate gonite, and B. ibari and B. argen- tata have digitiform gonites with blunt api- ces (these may have thickened, spinelike se- tae apically, but these are not comparable to the stout, pointed, spurlike gonal setae of B. dentata). The shape of the elongate epan- drium (+ fused surstyli ?) of B. ibari, a Palearctic species, somewhat resembles that of B. dentata in having a relatively deep and angulate emargination along the ante- rior surface (best seen in lateral view), but the extended epandrium (+ fused surstyli) of B. ibari are more nearly parallel sided in posterior view (see illustrations in Mathis and Ghorpadé 1985: 13, and Wirth 1964: 10 [as B. argentata}). Brachydeutera longipes Hendel (Figs. 6-8) Brachydeutera longipes Hendel 1913: 99 [Taiwan. Kankau; LT d (designated by Mathis and Ghorpadé 1985: 15), NMW].—Wirth 1964: 7-8 [revision].— Venkatesh 1976: 345-346 [biology].— Venkatesh et al. 1977a: 60—61 [coexis- tence with Culex fagitans Wiedemann]; 1977b: 10-13 [behavior].—Mathis and Ghorpadé 1985: 15-18 [revision].— Mathis and Steiner 1986: 56—60 [revi- sion|.—Mathis and Zatwarnicki 1995: 231 [world catalog]. Diagnosis.—This species is distinguished from congeners by the following combina- tion of characters: facial carina low, less prominent; generally paler in color, espe- cially the dorsum of mesonotum and ab- domen; anterior notopleural seta present; brown color of anepisternum gradually be- coming paler ventrally, merging with pale gray; apparent merger of fused surstyli with epandrium in lateral view with a shallow, broad swelling along anterior margin; gon- ite (Fig. 7) narrow, sinuous, slightly spatu- late apically. Specimens examined.—BARBADOS. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-8. Structures of the male terminalia of Brachydeutera longipes. 7, Cerci, epandrium (+ fused surstyli ?), gonite, and aedeagus, lateral view. 8, Same, posterior view. St: Philip: Gemswick (13°05’N, 59°28.5'W), 31 Aug 1997, W. N. Mathis (1 3, 1 2; USNM). BOLIVIA. La Paz: Guanay (15°29.8’S, 67°52.7'W; 460 m), 13 Mar 2001, W. N. Mathis (1 ¢; USNM). CAYMAN ISLANDS. Grand Cayman: Hutland (1.5 km S; 19°20'N, 81°13’W), 18 Feb 1993, W. E. Steiner, J. M. Swearingen, FE J. Burton (1 6, 2 2; USNM). DOMINICAN REPUBLIC. Hato Mayor: Hato Mayor (5.5 km E; 18°46.4'N, 69°12.5'W), 26 May 1998, D. and W. N. Mathis (2 ¢; USNM). La Vega: Rio Camu (G:5* km: NW? ‘La Vega; 19°13t8oN; 70°35.2'W; 100 m), 18 May 1998, D. and W. N. Mathis (3 d6, 2 2; USNM). Peder- nales: Pedernales (18°01.8'N, 71°44.7’W), 19-20 Mar 1999, W. N. Mathis (8 <6; USNM). Puerto Plata: River Camu (14 km E Puerto Plata; 19°11.9’N, 70°37.4'W), 17 VOLUME 105, NUMBER 2 May 1995, W. N. Mathis (1 6, 2 2: USNM). GRENADA. St. George: Point Salines Airport (W end; 12°0.3’N, 61°47.7’W), 12 Sep 1997, W. N. Mathis (2 ¢6; USNM). GUYANA. Georgetown (6°48.6'N, 58°08.6'W), 20-29 Aug 1997, W.N. Mathis (14 6, 3 2; USNM). HONDURAS. Cortés: San Pedro Sula (8 km S; 15°25.7'’N, 88°1.4’'W), 25-26 Sep 1995, D. and W. N. Mathis (1 d; USNM). JAMAICA. Manchester: Alligator Pond (17°52.1'N, 77°33.9'W), 8 May 1996, D. and W. N. Mathis, H. Williams (4 &; USNM). St. Andrew: Mavis Bank (near coffee factory; 18°01.4'N, 76°39.7'W; wa- terfall), 21-23 Apr 2000, W. N. Mathis (10 36,6 2; USNM). St. Elizabeth: near Port Kaiser (17°52.3'N, 77°34.9'W), 8 May 1996, D. and W. N. Mathis, H. Williams (3 2; USNM). MEXICO. Veracruz: Ciudad Aleman (100 m), 3 May 1985, W. N. Mathis (7 2: USNM). PUERTO RICO. Jayuya (2 km E; Rio Saliente; 18°12.8'N, 66°33.9'W), 22 Sep 1995, D. and W. N. Mathis (13 6, 4 @; USNM). VENEZUELA. Guarico: Hato Masa- guaral (45 km S Calabozo; 8°57'N, 67°58'W; 75 m; uv light; gallery forest), 3— 5 Jun 1988, M. Epstein (11 6, 7 2; USNM). Zulia: Maracaibo, Lagunas de Ox- idaci6n-ICLAM, Ciudad Universitaria (tanque pequeno), 18 May 1994, M. Garcia (2.d,,1 2; USNM). Distribution (Fig. 6).—Nearctic: Canada (ON), USA (GA, MD, NC). Neotropical: Bolivia (La Paz), Guyana, Honduras, Mex- ico (VRC), Venezuela (Guarico, Zulia), West Indies (Barbados, Cayman Islands, Dominican Republic, Grenada, Jamaica, Puerto Rico). Oriental: Cambodia, China, India, Indonesia, Malaysia, Pakistan, Phil- ippines, Singapore, Sri Lanka, Taiwan, Thailand, Vietnam. Palearctic: Iraq, Japan. Remarks.—Over 15 years ago, Mathis and Steiner (1986) first reported this species from the New World, and all of the locality 415 records were then from the Nearctic Region (southern Canada south to Georgia). In their world catalog, Mathis and Zatwarnicki (1995) included Mexico and Venezuela to the known distribution in the New World, and here, we add several islands in the Ca- ribbean and other siltes as far south as Bo- livia in South America (Fig. 6). These rec- ords clearly indicate that this species is well adapted to habitats in the New World and that it apparently disperses with consider- able ease or is more synanthropic and is dispersed by human activity. This species is clearly the most distinc- tive and easily identified species in the New World, especially the brown color of the anepisternum, which gradually becomes paler ventrally, merging with the pale gray coloration ventrally. The distinctive mor- phological features provide additional evi- dence that this species is adventive in the New World. In the Old World, B. longipes is quite similar and closely related to B. pleuralis Malloch, which also overlaps in much of its distribution (Mathis and Ghor- padé 1985). Distinguishing between B. lon- gipes and B. pleuralis will probably require examination of characters of the male ter- minalia. Brachydeutera neotropica Wirth (Figs. 9-16) Brachydeutera argentata of authors, not Walker [misidentification]: Williston 1897: 4 [in part, Brazil].—Cresson 1918: 67-68 [Costa Rica]; 1930: 105 [in part, Argentina, Mexico]; 1938: 34 [in part, Brazil].—Hendel 1930: 134 [in part, Bo- livia]|—Souza Lopes 1941: 654 [list, Brazil].—Sturtevant and Wheeler 1954: 204-205 [in part]—Wirth 1956: 18 [in part, Bahamas]; 1964: 5 [in part, distri- bution]. Brachydeutera neotropica Wirth 1964: 8 [Mexico. Vera Cruz: La Gloria, Cardel; HT 36, USNM (66452)]; 1968: 20 [Neo- tropical catalog]|.—Mathis 1983: 178— 179 [key, illustrations of male termina- lia]—Lizarralde de Grosso 1973: 83 [bi- 416 Figs. 9-10. Structures of the male terminalia of Brachydeutera neotropica. 9, Cerci, epandrium (+ fused surstyli ?), gonite, and aedeagus, lateral view. 10, Same, posterior view. ology, immature stages, Argentina]; 1989: 47 [review, Argentina].—Mathis and Zatwarnicki 1995: 231 [world cata- log]. Diagnosis.—This species 1s distinguished from congeners by the following combina- tion of characters: facial carina high, sharp- ly defined, especially ventrally; brown color of anepisternum continued ventrally to about dorsal /—/,, thereafter sharply delim- ited from pale gray coloration on ventral pleural region; apparent merger of fused surstyli with epandrium in lateral view in- dicated by an angulate emargination; gonite spatulate apically. Specimens examined.—ARGENTINA. Buenos Aires: 27 Sep 1905 (1 2; USNM). Tucumdn: Lacavera, 22—23 Nov 1951, M. L. Aczel, R. Golbach (3 ¢; USNM); Tu- cuman (mercury light trap), Feb 1935, R. Golbach (1 ¢; USNM). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON BAHAMAS. South Bimini Island: May 1951, M. Cazier (1d; AMNH). BARBADOS. Christ Church: Graeme Hall Swamp (13°04.2’N, 59°34.7'W), 12 Sep 1996, W. N. Mathis (1 ¢; USNM). St. Joseph: Joes River (13°12.8'N, 59°32.3'W), 10 Sep 1996, W. N. Mathis (8 6, 2 @; USNM). St. Philip: Gemswick (13°05'N, 59°28.5'W), 31 Aug 1997, W. N. Mathis (9 3,4 2; USNM). BOLIVIA. Cochabamba: Cochabamba (i7e23:3'S 65°07 Wz 226110) mm), 25) Mar 2001, W. N. Mathis (9 6, 2 2; USNM). La Paz> iGuanaya (3° ‘kmisEx, gS5es02uSF 67°52.3'W; 500 m), 14 Mar 2001, W. N. Mathis (1 ¢6, 3 2; USNM); Guanay (22 km SE; 15°17.8'S, 68°15.6'W; 540 m), 17 Mar 2001, W. N. Mathis (1 2; USNM); Mapiri (Rio Mapiri; 15°18.6'S, 68°13'W; 720 m), 17 Mar 2001, W. N. Mathis (1 2; USNM). BRAZIL. Parand: Foz do Iguacu (25°30.1'S, 54°32.4'W), 26 Aug 2000, D. and W. N. Mathis (3 2; USNM). Rio de Janeiro: Wha da Marambaia (23°03.6'S, 43°59.1’'W), 4 Sep 2000, D. and W. N. Mathis (1 2; USNM). CAYMAN ISLANDS. Grand Cayman: Hutland (1.5 km S; 19°20'N, 81°13'W; black light; forest near mangrove), 18 Feb 1993, W. E. Steiner, J. M. Swearingen, E J. Burton (3 6, 3 2; USNM). COLOMBIA. Antioquia: El Retiro (7 km N), 26 Feb 1984, W. N. Mathis (1 &; USNM). COSTA RICA. Limon: Suerre (10°17'N, 83°24'W), 14 Jul 1923, A. Alfaro (1 6, 1 2; USNM). Puntarenas: Las Loras, FE Knab (1 2; USNM). San José: La Caja (8 km W San José), 1930, P. Schmidt (2 6; USNM); San José, J. E Tristan (1 6, 1 @; USNM). CUBA. Cienfuegos: Soledad, Jardin Bo- tanico (22°7.5'N, 80°19.2’W), 13 Dec 1994, W. N. Mathis (7 d; USNM); Topes de Col- lantes (5 km WNW; 21°56.5'N, 80°2.3'W; 600 m), 11 Dec 1994, W. N. Mathis (2 d, 2 2; USNM). Havana: Ojo de Aqua (23°54.6'N, 82°29.1'W), 8 Dec 1994, W.N. Mathis (1 6; USNM); Havana (beach; VOLUME 105. NUMBER 2 417 he) Figs. 11-15. Male terminalia of Brachydeutra neotropica. 11, Epandrium (+ fused surstyli ?) and gonite (Bolivia. Cochabamba: Cochabamba). 12, Epandrium (+ fused surstyli ?) and gonite (Bolivia. La Paz: Guanay). 13, Epandrium (+ fused surstyli ?) and gonite (Ecuador. Manabi: Flavia Alfonso (42.7 km E)). 14, Epandrium (+ fused surstyli ?) and gonite (Ecuador. Imbaburra: Laguna San Pablo). 15, Epandrium (+ fused surstyli ?) and gonite (Cuba. Pinar del Rio: Soroa). 23°5.8'N, 82°27.7'W), 2-14 Dec 1994, W. N. Mathis (5 6, 2 2; USNM). Pinar del Rio: Soroa (22°47.7'N, 83°W), 4-6 Dec 1994, W. N. Mathis (15 6, 1 2; USNM). Sancti Spiritus: TYopes de Collantes (21°55.2'N, 80°02’W; 350 m), 10 Dec 1994, W. N. Mathis (1 ¢d, 2 2; USNM); Topes de Collantes (21°54.4'N, 80°01.4’'W; 670 m), 9-11 Dec 1994, W. N. Mathis (2 &; USNM). DOMINICA. Cabrits, 22 Mar 1989, W. N. Mathis (1 2; USNM). DOMINICAN REPUBLIC. Barahona: Barahona (18°12'N, 71°05.3’W), 25 Mar 1999, W. N. Mathis (3 3d, 1 2; USNM); Cabral (canals E of Cabral; 18°15.2’N, 71°13.4'W), 16 May 1995, W. N. Mathis (4 é5 I 23 USNM); Ojeda (117°58.2'N, 71°10.6'W), 22 Mar 1999, W. N. Mathis (1 36; USNM); Paraiso (5 km N;: 18°0O1.5’N, 418 71°11.6'W; 150 m), 21 Mar 1999, W. N. Mathis (1 6, 2 2: USNM). Distrito Na- cional: Santo Domingo (Jardin Botanico; 18°29.9'N, 69°56.9’'W), 25 May 1998, D. and W. N. Mathis (1 2: USNM). El Seibo: El Seibo (5 km E; 18°44.73’N, 68°59.2'W; 120 m), 12 May 1995, W. N. Mathis (2 d, 3 2; USNM): Rincén (near; 18°45.3'N, 68°55.7'W), 12 May 1995, W. N. Mathis (1 3,1 2: USNM). Hato Mayor: Hato Mayor (5.5 km E; 18°46.4'N, 69°12.5'W), 26 May 1998, D. and W. N. Mathis (2 2; USNM). Independencia: Los Bolos (18°37.8'N, 71°39.2'W; 1,370 m), 24 Mar 1999, W.N. Mathis (2 6, 1 2; USNM); Puerto Escon- dido (18°19.6'N, 71°35'W; 1,370 m), 24 Mar 1999, W. N. Mathis (2 6, 2 2; USNM). La Romana: Isla Saona, Mano Juan (18°08.1'N, 68°44.5'W), 13 May 1995, W. N. Mathis (2 2; USNM). La Vega: El Rio @:5 km E; 19°0:9’N, 70°33.5'W; 980 m), 6-7 May 1995, W.N. Mathis (5 6,5 2; USNM); Jarabacoa (1-2 km S; 19°06.9'N, 70°37'W; 520 m), 8-21 May 1995, 1998, D. and W. N. Mathis (13 6, 8 2; USNM); Jarabacoa (5 km S; 19°05.8'N, 70°36.5’W; 640 m), 8-20 May 1995, W. N. Mathis (1 d, 1 2; USNM); Rio Camu (3.5 km NW La Vega; 19°13.8'N, 70°35.2’W; 100 m), 18 May 1998, D. and W. N. Mathis (6 6, 2 @; USNM): Salto de Jimenoa (19°06'N, 70°35.9'W; 575 m), 20 May 1995, W. N. Mathis (1 ¢:; USNM). Monserior Nouel: dam near Rodeo (18°53.1’N, 70°33.5’W), 22 May 1998, D. and W. N. Mathis (1 2 USNM); Pedernales: Pedernales (18°01.8'N, 71°44.7'W), 19-20 Mar 1999, W. N. Mathis (5 6, 2 2; USNM); Peder- nales (19 km N; 18°09.2'N, 71°44.8'W; 230 m), 20 Mar 1999, W.N. Mathis (2 d, 1 &; USNM). ECUADOR. Guayas: El Triunfo (60 km E Guayaquil), 11 Feb 1973, M. A. Deyrup (2 6; USNM). Imbabura: Laguna San Pab- lo (0°12.1'N, 78°14.1'W), 28 Aug 1999, W. N. Mathis (1 ¢; USNM). Manabi: Arabia (12 km W Chone;: 70 m), 8 Jan 1978, W. N. Mathis (2 2; USNM); Bandurria, Aug PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 19555) Je Re) Levi-GCastilloni(an 2 ae USNM). Orellana: Rio Tiputini (0°38.2’S, 76°8.9'W), 12-26 Aug 1999, W. N. Mathis, A. Baptista, M. Kotrba (2 3d, 1 2; USNM). GRENADA. St. George: Point Salines Airport (W end; 12°0.3'N, 61°47.7'W), 12 Sep 1997, W. N. Mathis 3 6, 3 Q; USNM),; True Blue Beach (11°59.9'N, 61°46.1'W), 15 Sep 1996, W. N. Mathis (1 2; USNM). St. John: Concord Falls (12°07.1'N, 61°43'W), 14-21 Sep 1996, W. N. Mathis (1 6; USNM); Concord Valley (12°06.9'N, 61°43.9’W), 14 Sep 1996, W. N. Mathis (1 ¢; USNM). St. Patrick: Bath- way Beach (12°12.6'N, 61°36.7’W), 12 Sep 1997, W. N. Mathis (1 ¢; USNM); Levera Bay (12°13.6'N, 61°36.6’W), 18 Sep 1996, W. N. Mathis (1 6; USNM). GUATEMALA. Cayuga, Jan 1915 (1 2; USNM). San Cristobal: Alta Vera Paz, 17 May 1926, J. M. Aldrich (dx 19: USNM). GUYANA. CEIBA (ca. 40 km S George- town; 6°29.9'N, 58°13.1’W), 13-21 Apr 1994, 1995, W. N. Mathis (1 ¢; USNM). Moco-Moco (30 km E Lethem, Kanuku Mountains; 3°18.2’N, 59°39.0'W), 29 Apr 1995, W. N. Mathis (1 6; USNM). JAMAICA. Clarendon: Grantham (18°09.3'N, 77°23.8'W; 340 m), 16 Apr 2000, W. N. Mathis (1 2; USNM); Portland Cottage (17°45.4'N, 77°11'W), 13 May 1996, D. and W. N. Mathis, H. Williams (5 6, 12s TUSNM);” Rest (Gio) skmrN: 17°54 N, 77-211 WwW); 9) May 19965: and W. N. Mathis, H. Williams (1 2; USNM); Rest (3 km N; 17°54.3'N, 77°21.4’'W), 15 Apr 2000, W. N. Mathis (1 3,2 2; USNM); Rocky Point (2 km S, near Jackson Bay Cave; black light), 10 Dev 1975, G. F Hevel (1 ¢, 7 2; USNM). Man- chester: Alligator Pond (17°52.1'N, 77°33.9'W), 8 May 1996, D. and W. N. Mathis, H. Williams (1 6, 1 2; USNM). Portland: Crystal Springs (18°12.5'N, 76°37.9'W), 18 May 1996, D. and W. N. Mathis, H. Williams (1 6; USNM); Long Bay (2.3 km W; 18°06.5'N, 76°20’W), 24 Apr 2000, W. N. Mathis (1 6, 2 &; VOLUME 105, NUMBER 2 USNM); Reach (4 km N; 18°03.6’N, 76°20.4'W), 15 May 1996, D. and W. N. Mathis, H. Williams (1 ¢6; USNM). St. An- drew: Cinchona (18°04.4'N, 76°39.3'W; 1400 m), 29 Apr 2000, W. N. Mathis (5 2, 3 2: USNM):; Mavis Bank (1.5 km W;: 18°01.4'N, 76°39.9'W), 22 Apr 2000, W. N. Mathis (9 ¢, 3 2; USNM); Mavis Bank (1.7 km E; 18°02.4’N, 77°39.5’W; 575 m), Yallahs River, 21-22 Apr—1 May 2000, W. N. Mathis (1 2; USNM); Mavis Bank (4.3 km SE; 18°01.4'N, 76°38.1'W; 480 m); Yal- lahs River, 22—23 Apr 2000, W. N. Mathis (1 9%; USNM). St. Elizabeth: Elim (18°07.1'N, 77°40.6'W), 10 May 1996, D. and W. N. Mathis, H. Williams (2 ¢, 1 @; USNM); Elim (18°07.1'N, 77°40.5’W), 10 Apr 2000, W. N. Mathis (1 6, 1 2: USNM); near Port Kaiser (17°52.3’N, 77°34.9'W), 8 May 1996, D. and W. N. Mathis, H. Williams (6 6, 2 2; USNM). MEXICO. Chiapas: Finca Prusia (33 km S Jaltenango; 1,000 m), 12 May 1985, W. N. Mathis (4 ¢, 1 2; USNM). Jalisco: Tla- quepaque, Sep 1965, N. L. H. Krauss (1 ¢; USNM). Veracruz: Ciudad Aleman (100 m), 3 May 1985, W. N. Mathis (1 6; USNM); Fortin de las Flores (952 m), 2— 31 May 1981, 1985, C. M. and O. S. Flint, Jr., W. N. Mathis (6 6, 2 2; USNM); La Gloria Cardel, 1938, J. Camelo (2 3, 16 @; USNM). PANAMA. Canal Zone: Fort Kobbe, 17 Jun 1952, EF S. Blanton (3 2; USNM); Pe- dro Miguel, R. C. Shannon (1 ¢d; USNM). Darien: Punta Patino, 21 Jun 1952, FE S. Blanton (1 2; USNM). PERU. Junin: Perene (10°58'S, 75°13'W), R. C. Shannon (2 2; USNM). Lima: Lima, Laguna de Villa, 30 Aug 1988, W. N. Mathis (1 ¢6; USNM). Loreto: Itaya (ca. 25 km S Iquitos), 22 Feb 1984, W. N. Mathis (1 2; USNM). Madre de Dios: Manu, Rio Manu, Cocha Salvador (240 m), 14 Sep 1988, W. N. Mathis (1 ¢6; USNM). PUERTO RICO. Caribbean National Forest, Toro Negro Unit (18°8'N, 66°30'W: 419 Rd. 564 K.5.7.; 878 m), Dona Juana Forest, 17 Aug 1965, S. M. Gaud (1 ¢; USNM). Jayuya (2 km E; Rio Saliente; 18°12.8’N, 66°33.9'W), 22 Sep 1995, D. and W. N. Mathis (3 6, 1 2; USNM). Mayaquez, Sep—Nov 1960, M. M. Beauchamp (1 d: USNM). Salinas (black light; sugar cane field), 19 Feb 1964, S. M. Gaud, G. Rivera (7 6,8 2; USNM). ST. LUCIA. Fond St. Jacques (13°50'N, 61°02'W), 13-14 Jun 1991, D. and W. N. Mathis (1 gd; USNM). Soufriere Botanical Garden (13°51'N, 61°04’W), 12 Jun 1991, D. and W. N. Mathis (6 6, 3 2; USNM). ST. VINCENT. Charlotte: Montreal (13°12'N, 61°11’W), 9 Jun 1991, D. and W. N. Mathis (1 6; USNM); Peruvian Vale (13°10.7'N, 61°08.7'W), 6-8 Sep 1997, W. N. Mathis (2 6, 2 2; USNM). St. George: Yambou Head, 27 Mar 1989, W. N. Mathis (4 56,4 2; USNM). SURINAM. Paramaribo, 4 Sep 1943, D. G. Hall (4 5,6 2; USNM). TOBAGO. St. Patrick: Pigeon Point (beach; 11°9.7'N, 60°50'’W), 19 Apr 1994, D. and W. N. Mathis (1 2; USNM). TRINIDAD. Victoria: Basse Terre (7 km E; 10°07’N, 61°14’W), 27 Jun 1993, W.N. Mathis (2 2; USNM). VENEZUELA. Aragua: Estacion Exper- imental Cataurito, Villa de Cura (32 km E; 1,100\m), 1 Feb 1983, O. S. Flint, Ir. (S. d; 10 2). Guarico: Fundo Masaguaral, Lago Guacimas, 19 Jan 1983, O. S. Flint, Jr. (1 2; USNM): Hato Masaguaral (45 km S Calabozo; 8°57'N, 67°58'W), 3—5 Jun 1988, M. Epstein (13 d, 33 2; USNM). Distribution (Fig. 16).—Nearctic: USA (FL, LA, NC, TX). Neotropical: Argentina (Buenos Aires, Tucuman), Bahamas, Boliv- ia (Cochabamba, La Paz), Brazil (Parana, Rio de Janeiro), Costa Rica, Ecuador, Gua- temala, Guyana, Mexico (CHI, JAL, VRC), Panama, Peru (Junin, Lima, Loreto, Madre de Dios), Surinam, Trinidad and Tobago, Venezuela (Aragua, Guarico), West Indies (Barbados, Cuba, Dominica, Dominican 420 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 16. Distribution map for Brachydeutera neotropica. Republic, Cayman Islands, Grenada, Ja- (1973) described and figured the immature maica, Puerto Rico, St. Lucia, St. Vincent). stages of this species, which she discovered Natural history.—Lizarralde de Grosso in a variety of habitats from coastal low- VOLUME 105, NUMBER 2 lands to elevations of 1000 m in the moun- tains. Like other species of Brachydeutera, the puparium of B. neotropica has com- pletely everted anterior spiracles. Remarks.—This paper was prompted when we noted variation in the shape of the gonites and in the wing venation of Boli- vian specimens from two sites (La Paz: Guanay (Fig. 12); and Cochabamba: Co- chabamba (Fig. 11)). We also noted slight variation in the ventromedial extension of the epandrium (+ fused surstyli) in other specimens. After studying numerous spec- imens from throughout the neotropics, in- cluding dissections of male terminalia (Figs. 9-15), we have concluded that there is a single species with slight variation from population to population. More detailed studies, however, may reveal two and per- haps more species, especially as the varia- tion noted was primarily between localities with almost no variation among specimens from a single site. Brachydeutera sturtevanti Wirth (Figs. 17-18) Brachydeutera sturtevanti Wirth 1964: 11 [USA. California: San Diego County, San Diego; HT 3d, USNM (66453)].— Mathis 1983: 179 [figures of male ter- minalia]|.—Mathis and Steiner 1986: 57 [key].—Mathis and Zatwarnicki 1995: 232 [world catalog].—Keiper and Walton 2000: 468—475 [natural history, imma- ture stages]. Diagnosis.—This species is distinguished from congeners by the following combina- tion of characters: facial carina low, bluntly rounded, especially ventrally; brown color of anepisternum continued ventrally to about dorsal /,—/,, thereafter sharply delim- ited from pale gray coloration on ventral pleural region; apparent merger of fused surstyli with epandrium in lateral view in- dicated by a rounded emargination; gonite (Fig. 17) tapered and moderately pointed Figs. 17-18. — Structures of the male terminalia of Brachydeutera sturtevanti. 17, Cerci, epandrium (+ fused surstyli ?), gonite, and aedeagus, lateral view. 18, Same, posterior view. apically, apex lacking a_ stout, toothlike seta. Specimens examined.—MEXICO. Méx- ico: Nevado de Toluca, 7 Aug 1972, R. Gangan, T. P. Sluss (1 6, 6 2; USNM); Mexico City, 17 Sep, J. Muller (1 2; USNM); Toluca (32 km E; 3,048 m), 12 Aug 1954, P. R. Dreisbach (1 2; USNM). Morelos: Cuernavaca, Jul 1965, N. L. H. Krauss (2 2; USNM). Distribution.—Nearctic: USA (AZ, CA, NM, TX). Neotropical: Mexico (MEX, MOR). Natural history.—Keiper and Walton (2000) published an excellent study on the immature stages and biology of this species and included a key to the third-instar larvae of all available species from the New World. They discovered that larvae of this species are hyponeustic (are suspended from the water’s surface by hydrofuge hairs on the posterior spiracles) and exhibit great versatility in feeding. The larvae collected 422 or scraped algae and detritus from solid substrates or brought their mouthparts to the water’s surface and created a vortex to initiate filter feeding. This species is able to colonize and complete larval development in habitats where the food quality varied considerably. ACKNOWLEDGMENTS We gratefully acknowledge the assistance and cooperation of many organizations and individuals who contributed to the field work and production of this paper. To Da- vid A. Grimaldi (AMNH), Jon K. Gelhaus and Donald E Azuma (ANSP), John E. Chainey (BMNH), J. Richard Vockeroth (CNC), and their institutions, who loaned specimens, We express our sincere thanks. The illustrations were carefully inked by Mr. Young T. Sohn. Hollis B. Williams pro- vided technical support. We thank Robert K. Robbins for making the digitized maps available to us. For reviewing a draft of this paper, we thank Terry Wheeler, Joe B. Kei- per, Tadeusz Zatwarnicki, and Stephen D. Gaimari. We are also grateful to David Challinor, former Assistant Secretary for Research, and to Anna K. Bernensmeyer and David L. Pawson, former Associate Di- rectors for Science, National Museum of Natural History, Smithsonian Institution, for financial support to conduct research at The Natural History Museum (BMNH), London, England, through grants from the Research Opportunity Fund. In 1995, 1996, and 1998, field work on the West Indies was funded in large measure by grants from the Biodiversity Program (Biotic Surveys and Inventories, BSI), National Museum of Natural History, Smithsonian Institution (Lynne R. Parenti, former chair, George R. Zug, chair). Field work on the West Indies was greatly expedited through the able and pleasant assistance of N. Dianne Mathis, Hollis B. Williams, Kelvin Guerrero, Dan- iel E. Pérez-Gelabert, and Oliver S. Flint, Jr. Field work in Bolivia was supported by PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON a grant from the Biodiversity Program (Bi- otic Surveys and Inventories, BSI), Nation- al Museum of Natural History, Smithsonian Institution (George R. Zug, chair). Field work in Guyana was supported by the Smithsonian Institution’s Biological Diver- sity of the Guianas Program (publication number 64; Vicki A. Funk, Director; Carol L. Kelloff, Coordinator). LITERATURE CITED Cogan, B. H. 1980. 71. Family Ephydridae, pp. 655— 669. In Crosskey, R. W., ed. Catalogue of the Dip- tera of the Afrotropical Region. British Museum (Natural History), London. 1437 pp. . 1984. Family Ephydridae, pp. 126-176. In Sods, A. and L. Papp, eds. Catalogue of Palaearc- tic Diptera. Elsevier Science Publishers, Amster- dam, and Akadémiai Kiad6, Budapest. 10: 402 pp. Cogan, B. H. and W. W. Wirth. 1977. Family Ephy- dridae, pp. 321-339. Jn Delfinado, M. D. and D. E. Hardy, eds. A Catalogue of the Diptera of the Oriental Region, Vol. If. Suborder Cyclorrhapha (excluding division Aschiza). University Press of Hawaii, Honolulu. 451+ vii pp. Crawford, J. C. 1914. Three new Hymenoptera (Chal- cidoidea). Insecutor Inscitiae Menstruus 2(3): 36— 38. Cresson, E. T., Jr. 1918. Costa Rican Diptera Collected by Philip P. Calvert, Ph.D., 1909-1910. Paper 3. A report on the Ephydridae. Transactions of the American Entomological Society 44: 39-68. . 1930. Studies in the dipterous Family Ephy- dridae. Paper II. Transactions of the American Entomological Society 56: 93-131. . 1938. Notes on, and descriptions of, some Neotropical Ephydridae (Dipt.). Revista de Ento- mologia 8(1—2): 24—40. Foote, B. A. 1995. Biology of shore flies. Annual Re- view of Entomology 40: 417-442. Hendel, E 1913. Acalyptrate Musciden (Dipt.) II. Jn H. Sauter’s Formosa-Ausbeute. Supplementa En- tomologica 2: 77-112. . 1930. Die Ausbeute der deutschen Chaco-Ex- pedition 1925/26. Diptera. XIX. Ephydridae. Ko- nowia 9(2): 127-155. Hennig, W. 1952. Die Larvenformen der Dipteren. 3: 1-628 + vii. Akademie-Verlag, Berlin. Keiper, J. B. and W. E. Walton. 2000. Biology and immature stages of Brachydeutera sturtevanti (Diptera: Ephydridae), a hyponeustic generalist. Annals of the Entomological Society of America 93: 468-475. Lizarralde de Grosso, M. S. 1973 [*1972”]. Notas so- VOLUME 105, NUMBER 2 bre Ephydridae Argentinos. I. (Diptera) Descrip- cion de las larvas y pupas de Scatella notabilis Creson y Brachydeutera neotropica Wirth. Revis- ta de la Sociedad Entomologica de Argentina 34(1-2): 79-84. . 1989. Ephydridae de la Republica Argentina (Insecta—Diptera). Serie Monografica y Didactica No. 3. Facultad de Ciencias Naturales e Instituto Miguel Lillo Universidad Nacional de Tucuman. 93 pp. Loew, H. 1862. Monographs of the Diptera of North America. Part 1. Smithsonian Institution, Smith- sonian Miscellaneous Collections 6(141): 1-221. Mathis, W. N. 1982. Studies of Ephydrinae (Diptera: Ephydridae), VI: Review of the Tribe Dagini. Smithsonian Contributions to Zoology 345: 30 + iv pp. . 1983. Notes on Brachydeutera Loew (Dip- tera: Ephydridae) from North America. Entomo- logical News 94(5): 177-180. . 1986. Studies of Psilopinae (Diptera: Ephy- dridae), I: A revision of the shore fly genus Pla- copsidella Kertész. Smithsonian Contributions to Zoology 430: iv+30 pp. . 1989. 66. Family Ephydridae, pp. 639-649. In Evenhuis, N. L., ed. Catalog of the Diptera of the Australasian and Oceanian Regions. B. P. Bishop Museum special publication 86 and E. J. Brill. Honolulu and Leiden. 1155 pp. Mathis, W. N. and K. D. Ghorpadé. 1985. Studies of Parydrinae (Diptera: Ephydridae), I: A review of the genus Brachydeutera Loew from the Oriental, Australian, and Oceanian Regions. Smithsonian Contributions to Zoology 406: iv + 25 pp. Mathis, W. N. and W. E. Steiner, Jr. 1986. An adven- tive species of Brachydeutera Loew in North America (Diptera: Ephydridae). Journal of the New York Entomological Society 94(1): 56-61. Mathis, W. N. and T. Zatwarnicki. 1990a. A revision of the western Palearctic species of Athyroglossa (Diptera: Ephydridae). Transactions of the Amer- ican Entomological Society 116(1): 103-133. . 1990b. Taxonomic notes on Ephydridae (Dip- tera). Proceedings of the Biological Society of Washington 103(4): 891-906. . 1995. A world catalog of the shore flies (Dip- tera: Ephydridae). Memoirs on Entomology, In- ternational 4: vi + 423 pp. McAlpine, J. EF 1981. Morphology and Terminology- Adults, pp. 9-63. In McAlpine, J. F. et al., eds. Manual of Nearctic Diptera. Ottawa. 1: 674 pp. Osten Sacken, C. R. 1878. Catalogue of the described Diptera of North America. Smithsonian Institu- tion, Smithsonian Miscellaneous Collections, 16(2 [= publication 270]): 1-276. Scheiring, J. F and B. A. Foote. 1973. Habitat distri- 423 bution of the shore flies of northeastern Ohio (Diptera: Ephydridae). Ohio Journal of Science 73(3): 152-166. Souza Lopes, H. l6gico capturado, pp. 641-660. /n Travassos, L., 1941. Relagao do material entomo- ed. Relatorio da terceira excursao a zona da Es- trada de Ferro Noroeste do Brasil realizada in fev- ereiro e marco de 1940. Memorias do Instituto Oswaldo Cruz 35(3)[1940]: 641—660. Sturtevant, A. H. and M. R. Wheeler. 1954. Synopses of Nearctic Ephydridae (Diptera). Transactions of the American Entomological Society 79: 151— 2511. Venkatesh, M. G. 1976. Some observations on the bi- ology of Brachydeutera longipes Hendel (Diptera: Ephydridae) from West Bengal. Journal of the Bombay Natural History Society 57: 345-346. Venkatesh, M. G., G. P. Channabasavanna, and M. D. Parthasarathy. 1977a. Co-existence of the shore- fly Brachydeutera longipes Hendel and the larvae of Culex fagitans Wiedemann in Natural Habitats. Second Oriental Entomology, Symposium, pp. 60-61. Venkatesh, M. G., G. P. Channabasavanna, and M. D. Parthasarathy. 1977b. Food and feeding behaviour of the shore-fly maggot, Brachydeutera longipes Hendel (Diptera: Ephydridae). Indian Journal of Behavior 1(3): 10-13. Walker, F 1853. Diptera. /n Saunders, W. W., ed. In- secta Saundersiana: or Characters of undescribed insects in the collection of William Wilson Saun- ders, Esq., EF R. S., EL. S., &c. Van Voorst, Lon- don. 1(4): 253-414. Williams, EF X. 1938. Biological studies in Hawaiian water-loving insects. Part HI. Diptera or flies. A, Ephydridae and Anthomyiidae. Proceedings of the Hawaiian Entomological Society 10(1): 85-119. Williston, S. W. 1897. Diptera Brasiliana. Part IV. Kan- sas University Quarterly series A, 6: 1-12. 1908. Manual of North American Diptera. New Haven: James T. Hathaway, 3rd ed., 405 pp. Wirth, W. W. 1956. The Ephydridae (Diptera) of the Museum Bahama Islands. American Novitates 1817: 1-20. . 1964. A revision of the shore flies of the ge- nus Brachydeutera Loew (Diptera: Ephydridae). Annals of the Entomological Society of America 57(1): 3-12. . 1965. Ephydridae, pp. 734—759. Jn Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson, eds. A Catalog of the Diptera of America north of Mexico. United States Depart- ment of Agriculture, Agriculture Handbook No. 276, 1696 pp. . 1968. 77. Family Ephydridae. /n Papavero, 424 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON N., ed. A Catalogue of the Diptera of the Ameri- | Zatwarnicki, T. 1992. A new classification of Ephy- cas south of the United States, 43 pp. Departa- dridae based on phylogenetic reconstruction (Dip- mento de Zoologia, Secretaria da Agricultura. Sao tera: Cyclorrhapha). Genus 3(2): 65-119. Paulo. . 1996. A new reconstruction of the origin of Woodley, N. E. and D. J. Hilburn. 1994. The Diptera Eremoneuran hypopygium and its implications for of Bermuda. Contributions of the American En- classification (Insecta: Diptera). Genus 7(1): 103— tomological Institute 28(2): 1—64. 175. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 425—440 MIRIDAE (HETEROPTERA) RECORDED FROM CHINA SINCE THE 1995 WORLD CATALOG BY R. T. SCHUH BAOYING QI, CARL W. SCHAEFER, NONNAIZAB BAI, AND ZHEMIN ZHENG (BQ, ZZ) Institute of Zoology, College of Biological Science, Shaanxi Normal Uni- versity, Xian, 710062, People’s Republic of China; (CWS) Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT 06269-3043, U.S.A.; (BQ, NB) Department of Biology, Inner Mongolia Normal University, Huhe-hot, 010022, Peo- ple’s Republic of China Abstract.—A list of described and recorded species of Miridae from China since the 1995 world catalog of plant bugs by R. T. Schuh is presented. Information about descrip- tions, figures, distributions, type specimens, host plants, and references are listed in order to guide further studies on these insects. Key Words: Miridae, the largest family in hemipteran suborder Heteroptera, about 10,000 species, belonging to eight subfam- ilies, about 30 tribes, and more than 1,000 genera (Schuh 1995, Schuh and_ Slater 1995). In 1995, Schuh published his fine world catalog of mirids, which followed Carvalho’s world catalog (1952-1960). In 1995, Zheng published a list of Chinese mirids described since Carvalho’s catalog. Because the mirid fauna of China is large, contains very interesting, and not wholly known, de- scriptions of new mirids have continued to be published since Schuh (1995) and Zheng (995). We have gathered these post 1995 de- scriptions, checked them, and present here a list of 215 species described or recorded from China since 1994. The list includes 4 genera, 125 species, | subspecies, and 44 records new for China, and 45 combina- tional or nomenclatorial changes. Abbreviations used: BML: The Natural History Museum, London; DBNU: Depart- ment of Biology, Tiejin Nankai University; DPPBAN: Department of Plant Protection, list, Heteroptera, Miridae, China Beying Agricultural University; HNHM: Hungarian Natural History Museum, Bu- dapest, Hungary; HUESJ: Hokkaido Uni- versity of Education, Sapporo, Japan; DBIMNU: Department of Biology, Inner Mongolia Normal University, China; IZAS: Institute of Zoology, Academia Sinica, Bei- jing; NSMT: National Science Museum (National History), Tokyo, Japan; NWAU: Northwestern Agricultural University, Yan- gling, Shaanxi Province, China; TNHM: Tianjin Natural History Museum, Tianjin; USNM: National Museum of Natural His- tory, Smithsonian Institution, Washington D.C.; ZIRASP: Zoological Institute, Rus- sian Academy of Sciences, St. Petersburg; BJ: Beying; GD: Guangdong Province; GS: Gansu Province; GX: Guangxi Province; FJ: Fujian Province; HEB: Hebei Province; HEN: Henan Province; HLJ: Heilongjiang Province; HN: Hainan Province; HUB: Hu- bei Province; HUN: Hunan Province; IM: Inner Mongolia Autonomous Region; JL: Jijin Province; LN: Liaoning Province; NX: Ningxia Province; QH: Qinghai Province; SAX: Shaanxi Province; TW: Taiwan Prov- 426 ince; XJ: Xinjiang Autonomous Region; XZ: Xizang Automonous Region; YN: Yunnan Province; ZJ: Zhejiang Province. Miridae Hahn 1833 Orthotylinae Van Duzee 1916 (1865) Halticini A. Costa 1853 Coridromius Signoret 1862 Coridromius chinensis Liu and Zhao 1999: 57-58, figs. 3-8 (n. sp. description; dis- tribution; dorsal view of body; male gen- italia. Holotype and 32 paratypes, GX; | paratype, TW; 57 paratypes, FJ. All types deposited in DBNU). Host plant un- known. Coridromius testaceous Liu and Zhao 1999: 56-57, figs. 1-2 (n. sp. description; distribution; dorsal view of body, hind leg. Holotype, HN. Deposited in DBNU). Host plant unknown. Orthotylini Van Duzee 1916 (1865) Cyllecoris Hahn 1834 Cyllecoris badius Liu and Zheng 1999a: 102, 103-104, figs. 22-1-1, 22-2 (n. sp. description; key; distribution; dorsal view of head and pronotum, male genitalia. Holotype, IM. Deposited in DBNU). Host plant unknown. Cyllecoris nakanishii Miyamoto 1969; Liu and Zheng 1999a: 102, 105 (new record; key; distribution; 2 specimens examined, JL. Deposited in DBNU). Host plant un- known. Cyllecoris opacicollis Kerzhner 1988; Liu and Zheng 1999a: 102, 105-106, figs. 22-4 (new record; key; distribution; male genitalia; 2 specimens examined, NX. Deposited in DBNU). Host plant un- known. Cyllecoris rectus Liu and Zheng 1999a: 102, 106-107, figs. 22-5 (n. sp. descrip- tion; key; distribution; dorsal view of head-pronotum; male genitalia. Holotype, HEN. Deposited in DBNU). Host: un- known. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Labopidea Uhler 1877 Labopidea algens (Vinokurov 1982) (Or- thotylus (Labopidea)); Tian and Bai 1998: 125, 126, figs. 31-33 (new record; key; brief description; distribution; male genitalia; IM). Host plant: Atriplex sp. Orthotylus Fieber 1858 Orthotylus (Melanotrichus) minutus Jakoy- lev 1877; Tian and Bai 1998: 125, 126, figs. 22-24 (new record; key; brief de- scription; distribution; male genitalia; IM). Host plant: Chenopodiaceae. Orthotylus (Orthotylus) riparius Kulik 1973; Tian and Bai 1998: 125, 126, figs. 7-9 (new record; key; brief description; distribution; male genitalia; IM). Host plant unknown. Orthotylus (Pinocapsus) alashanensis Tian and Bai 1999: 66-67, 69, fig. 1 (n. sp. description; distribution; host; dorsal view of body; male genitalia. Holotype, allotype and 7 paratypes, IM. Deposited in DBIMNU). Host plant: Sabina vulgar- is Antoine. Orthotylus (Pinocapsus) sabinae Tian and Bai 1999: 67-69, fig. 2 (n. sp. descrip- tion; distribution; host; dorsal view of body; male genitalia. Holotype, allotype and 7 paratypes, IM. Deposited in DBIM- NU). Host plant: Sabina vulgaris An- toine. Pseudoloxops Kirkaldy 1905 Pseudoloxops pilosus Liu and Zheng 1994: 119-121, figs. 1-3 (n. sp. description; list; key, distribution; male genitalia in different views. Holotype, GS. Deposited in DBNU). Host plant unknown. Pseudoloxops punctulatus Liu and Zheng 1994: 121-122, figs. 4-5 (n. sp. descrip- tion; list; key, distribution; male genitalia in different views. Holotype, GS. Depos- ited in DBNU). Host plant unknown. Pseudoloxops robustus Liu 1999: 51-53, 57, figs. 1-7 (n. sp. description; list; key, distribution; male genitalia in different views. Holotype and | paratype, YN. De- posited in DBNU). Host plant unknown. VOLUME 105, NUMBER 2 Zanchius Distant 1904 Zanchius innotatus Liu and Zheng 1999b: 388-389, fig. | (n. sp. description; distri- bution; dorsal view of body. Holotype and 3 paratypes, HN. Deposited in DBNU). Host plant unknown. Zanchius shaanxiensis Liu and Zheng 1999b: 389-390, figs. 2—6 (n. sp. de- scription; distribution; dorsal view of body; male genitalia. Holotype and 5 paratypes, SAX. Deposited in DBNU). Host plant unknown. Zanchius tibetanus Liu and Zheng 1999b: 390-391, figs. 7-10 (n. sp. description; distribution; dorsal view of body; male genitalia. Holotype, XZ. Deposited in DPPBAU). Host plant unknown. Phylinae Douglas & Scott 1865 Phylini Douglas & Scott 1865 Acrotelus Reuter 1885 Acrotelus pilosicornis ginghaiensis Qi and Bai 1995a: 144—145, figs. 1-5 (n. subsp. description; distribution; hind leg. Holo- type and 8 paratypes, QH. Deposited in DBIMNU). Host plant unknown. Atomophora Reuter 1879 Atomophora flavidus Bai and Yang 1994: 18-19, figs. 6-11 (n. sp. description; dis- tribution; dorsal view of body; front view of head; male genitalia. Holotype and 2 paratypes, IM. Deposited in DBIMNU). Host plant unknown. Atomophora punctulatus Bai and Yang 1994: 19-20, figs. 12-17 (n. sp. descrip- tion; distribution; dorsal view of body; front view of head; male genitalia. Ho- lotype and 7 paratypes, IM. Deposited in DBIMNU). Host plant unknown. Campylomma Reuter 1878 Campylomma annulicornis (Signoret 1865) (Litocoris); Qi and Bai 1996a: 50, 52, fig. 4 (new record; list, distribution, brief de- scription, male phallus and paramere; IM). Host plant unknown. Campylomma verbasci (Meyer-Duer 1843) 427 (Capsus); Qi and Bai 1996a: 50, 52-53, fig. 5 (new record; list, distribution, brief description, male phallus and paramere: IM). Host plant unknown. Compsidolon Reuter 1899 Compsidolon absinthii (Scott 1870) (Agal- liastes); Qi and Bai 1995b: 226 (new re- cord; distributions; QH). Host plant un- known. Compsidolon kerzhneri Kulik 1973; Bai and Yang 1994: 17 (new record; distri- bution: IM). Host plant unknown. Compsidolon pumilum (Jakovlev, 1876) (Agalliastes); Bai and Yang 1994: 17 (new record; distribution; IM). Host plant unknown. Compsidolon punctulatus Qi and Bai 1995b: 225-226, figs. 1—6 (n. sp. descrip- tion; distribution; dorsal view of body: male genitalia. Holotype and 9 paratypes, XJ. Deposited in DBIMNU). Host plant unknown. Dacota Uhler 1872 Dacota hesperia Uhler 1872; Bai and Chen 1996: 185 (new record; distribution; IM). Host plant unknown. Leucopterum Reuter 1879 Leucopterum candidatum Reuter 1879; Qi and Bai 1997: 11-12, 14, figs. 8-16 (new record; redescription; distribution; dorsal view of male body; claw; male genitalia; XJ). Host plant: Artemisia sp. Monochroica Qi & Bai 1996 Type species: Monochroica alashanensis Qi and Bai 1996 Monochroica Qi and Bai 1996b: 298—299, 302-303, figs. 1-9 (new genus; descrip- tion of general characters and the type species; identification). Monochroica alashanensis Qi and Bai 1996b: 299-301, 303-305, figs. 1-9 (n. sp. description; distribution; front view of head; lateral view of head-pronotum; tib- ia and tarsus of hind leg; male genitalia. Holotype, allotype and 8 paratypes, IM. 428 Deposited in DBIMNU). Host plant: Sa- bina vulgaris Antoine. Oncotylus Fieber 1858 Oncotylus Fieber 1858; Qi and Bai 1997: 12 (new record). Oncotylus vitticeps Reuter 1879: Qi and Bai 1997: 13-14, figs. 17—22 (new record; re- description; distribution; dorsal view of head and pronotum; male genitalia; XJ). Host plant: Artemisia sp. Phoenicocoris Reuter 1875 Phoenicocoris giliananus Zheng and Li 1996: 101-103, figs. 1-8 (n. sp. descrip- tion; distribution; front and lateral view of head; male genitalia. Holotype and 5 paratypes, GS. Deposited in DBNU). Host plant unknown. Placochilus Fieber 1858 Placochilus paraseladonicus Qi and Bai 1995a: 143-144, figs. 1-5 (n. sp. descrip- tion; distribution; male genitalia. Holo- type and 2 paratypes, XJ. Deposited in DBIMNU). Host plant unknown. Plagiognathus Fieber 1858 Plagiognathus muculosus Zhao and Li tion; distribution; male genitalia. Holo- type, allotype and 28 paratypes, GS. De- posited in DBNU). Host plant: Tamarix sp. Psallopsis Reuter 1901 Psallopsis halostachydis Putshkoy 1975; Bai and Yang 1994: 18 (new record; dis- tribution; IM). Host plant unknown. Psallopsis kirgisicus (Becker 1864) (Cap- sus); Bai and Yang 1994: 18 (new record; distribution; IM). Host plant unknown. Psallus Fieber 1858 Psallus (Psallus) fallenii Reuter 1883; Qi and Bai i994b: 342 (new record; distri- bution; IM). Host plant unknown. Psallus (Psallus) flavescens Kerzhner 1987; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Qi and Bai 1994b: 342 (new record; dis- tribution; IM). Host plant unknown. Psallus (Phylidea) innermongolicus (Qi and Bai 1995) (n. nom. for Psallus (Phylidea) kerzhneri, (a junior primary homonym of Psallus (Phylidea) kerzhneri Jositfov, 1992) by Qi and Bai 1995d: 256) (= Psallus (Phylidea) kerzhneri Qi and Bai 1994b: 341-344, figs. 3-7 (n. sp. descrip- tion; distribution; hind femora; male gen- italia. Holotype and allotype, IM. Depos- ited in DBIMNU. Host plant unknown)). (n. syn. of Psallus (Phylidea) ulmi Ker- zhner and Josifov 1966 by Kerzhner 1997: 245-248). Solenoxyphus Reuter 1875 Solenoxyphus flavicans Qi and Bai 1996c: 293-294, 299, figs. 1-7 (n. sp. descrip- tion; key; distribution; dorsal view of male body; male genitalia. Holotype and 116 paratypes, XJ. All deposited in DBIMNU). Host plant: Artemisia ssp. Solenoxyphus lepidus (Puton, 1874) (Ma- crocoleus) (Compsidolon gobicus Bai and Yang 1994: 17-18, figs. 1-5; n. syn. by Qi and Bai 1996c: 296-297, 299, figs. 14—20) (new record; description; key; distribution; male genitalia; XJ). Host plant: Artemisia ssp. Solenoxyphus viridulus Qi and Bai 1996c: 295-296, 299, figs. 8-13 (n. sp. descrip- tion; key; distribution; dorsal view of male body; male genitalia. Holotype and 116 paratypes, QH, XJ. All deposited in DBIMNU). Host plant: Artemisia ssp. Tuponia Reuter 1875 Tuponia (Tuponia) arcufera Reuter 1879; Qi and Bai 1997: 9, 14 (new record; dis- tribution; IM). Host plant: Tamarix sp. Tuponia (Tuponia) brevicula Qi and Bai 1997: 10-11, 14, figs. 1-7 (n. sp. descrip- tion; distribution; dorsal view of male body; male genitalia. Holotype and 8 paratypes, XJ. All deposited in DBIM- NU). Host plant: Tamarix sp. Tuponia (Tuponia) mongolica Drapolyuk 1980; Qi and Bai 1997: 10—11, 14 (new VOLUME 105, NUMBER 2 record; description; IM). Host plant un- known. Pilophorini Douglas & Scott 1876 Pilophorus Hahn 1826 Pilophorus clavatus (Linne 1767) (Cimex); Qi and Bai 1996a: 50, 51, fig. 2 (new record; list, distribution, brief description, male genitalia; IM). Host plant unknown. Pilophorus lucidus Linnavori 1962; Qi and Bai 1996a: 50, 51—S2, fig. 3 (new record; list, distribution, brief description, dorsal view of male body: IM). Host plant un- known. Hypseloecus Reuter 1891 Hypseloecus camplus (Kulik, 1968) (Pher- olepis), Qi and Bai 1996a: 50-51, fig. | (new record; list, distribution, brief de- scription, male genitalia; IM). Host plant unknown. Bryocorinae Baerensprung 1860 Bryocorini Baerensprung 1860 Bryocoris Fallén 1829 Bryocoris (Bryocoris) bui Hu and Zheng 2000: 244, 245-248, figs. 5, 10, 14, 26, 40, 41 (n. sp. description; key; distribu- tion; dorsal view of body; lateral view of head and pronotum; male genitalia. Ho- lotype and 4 paratypes, YN. Deposited in DBNU). Host plant unknown. Bryocoris (Bryocoris) concavus Hu and Zheng 2000: 244, 248-249, figs. 8, 30, 42 (n. sp. description; key; distribution; dorsal view of body; lateral view of head and pronotum; male genitalia. Holotype and 36 paratypes, YN, SC, XZ. Depos- ited in DBNU, BML, ZIRASP, HUESJ). Host plant unknown. Bryocoris (Bryocoris) insuetus Hu and Zheng 2000: 244, 249-251, figs. 13, 18, 24, 39, 44 (n. sp. description; key; distri- bution; dorsal view of body; lateral view of head and pronotum; male genitalia. Holotype and 5 paratypes, YN, SC. De- posited in DBNU). Host plant unknown. Bryocoris (Bryocoris) xiongi Hu and Zheng 429 2000: 244, 251-253, figs. 11, 12, 37, 38, 45 (n. sp. description; key; distribution, dorsal view of body; lateral view of head and pronotum; male genitalia. Holotype and 12 paratypes, YN. Deposited in DBNU). Host plant unknown. Bryocoris (Cobalorrhynchus) lit Hu and Zheng 2000: 244, 259-261, figs. 17, 23, 53 (n. sp. description; key; distribution; dorsal view of body; lateral view of head and pronotum; male genitalia. Holotype and 21 paratypes, XZ, YN. Deposited in DBNU). Host plant unknown. Bryocoris (Cobalorrhynchus) lobatus Hu and Zheng 2000: 244, 261—262, figs. 4, 31, 54 (n. sp. description; key; distribu- tion, dorsal view of body; lateral view of head and pronotum; male genitalia. Ho- lotype and 7 paratypes, YN. Deposited in DBNU). Host plant unknown. Bryocoris (Cobalorrhynchus) sichunanen- sis Hu and Zheng 2000: 244, 262-2064, figs. 2, 9, 28, 55 (n. sp. description; key; distribution, dorsal view of body; lateral view of head and pronotum; male geni- talia. Holotype and 26 paratypes, SC. De- posited in DBNU, IZAS, ZIRASP). Host plant unknown. Bryocoris (Cobalorrhynchus) vittatus Hu and Zheng 2000: 244, 264-265, figs. 6, 27, 56 (n. sp. description; key; distribu- tion, dorsal view of body; lateral view of head and pronotum; male genitalia. Ho- lotype, YN; 21 paratypes, YN, GX, SZ, TW. Deposited in DBNU, TNHM, BML). Host plant unknown. Dicyphini Reuter 1883 Mansoniella Poppius 1915 Mansoniella annulata Hu and Zheng 1999a: 159-161, figs. 3, 15 (n. sp. de- scription; dorsal view of body; lateral view of head. Holotype and 2 paratypes, SAX; | paratype, SC. Deposited in DBNU). Host plant unknown. Mansoniella cinnamomi (Zheng and Liu 1992) (Pachypeltis); Zheng and Liu 1992: 170, fig. 19 (n. comb.; distribution; 430 male paramere; HUN). Host plant un- known. Mansoniella cristata Hu and Zheng 1999a: 161-162, figs. 7, 12 (n. sp. description; distribution; dorsal view of body; lateral view of head-pronotum. Holotype, YN. Deposited in DBNU). Host plant un- known. Mansoniella flava Hu and Zheng 1999a: 164—165, figs. 5, 8 (n. sp. description; distribution; dorsal view of body; male genitalia. Holotype and 1 paratype, SAX. Deposited in DBNU). Host plant un- known. Mansoniella juglandis Hu and Zheng 1999a: 165-166, figs. 4, 13, 14, 18 (n. sp. description: distribution; dorsal view of body; lateral view of head-pronotum: male paramere. Holotype and 9 para- types, SC. Deposited in DBNU). Host plant unknown. Mansoniella rosacea Hu and Zheng 1999a: 166-167, figs. 2, 9, 17 (n. sp. description; distribution; dorsal view of body; lateral view of head-pronotum; male genitalia. Holotype, GS. Deposited in DBNU). Host plant unknown. Mansoniella rubida Hu and Zheng 1999a: 167-169, figs. 1, 10 (n. sp. description; distribution; dorsal view of body; lateral view of head-pronotum. Holotype and | paratype, SAX. Deposited in DBNU). Host plant unknown. Mansoniella sassafri (Zheng and Liu 1992) (Pachypeltis); Zheng and Liu 1992: 170, fig. 21 (n. comb.; distribution; left para- mere; HHN). Host plant unknown. Mansoniella wangi (Zheng and Li 1992) (Pachypeltis); Zheng and Li 1992: 170, fig. 20 (n. comb.; distribution; male par- amere; HN). Host plant unknown. Pachypeltis Signoret 1858 Pachypeltis biformis Hu and Zheng 1999b: 1-6, figs. 1-3, 5, 7 (m. sp. description; distribution; dorsal view of male body; lateral view of female head and prono- tum; male genitalia. Holotype and 6 para- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON types, GX; 6 paratypes, HN. Deposited in DBNU). Host plant unknown. Eccritotarsini Berg 1883 Sinevia Kerzhner 1988 Sinevia pallidipes (Zheng and Liu 1992) (Bryocoris); Zheng and Liu 2000: 265— 266, figs. 3, 49, 57 (n. comb.; distribu- tion; key: dorsal view of head; male gen- italia; HUN, GX, SAX. Material exam- ined deposited in DBNU, TNHM). Host plant unknown. Monaloniini Reuter 1892 Dimia Kerzhner 1988 Dimia Kerzhner 1988; Hu and Zheng 2001: 414—415 (new record, identification). Dimia inexpectata Kerzhner 1988; Hu and Zheng 2001: 415 (new record; distribu- tion; identification; HN). Host plant un- known. Eupachypeltis Poppius 1915 Eupachypeltis unicolor Hu and Zheng 2001: 416-417, fig. 1 (n. sp. description; dorsal and lateral view of head and pro- notum: front view of head. Holotype and | paratype, HN. Deposited in DBNU). Host plant unknown. Felisacus Distant, 1904 Felisacus amboinae Woodward 1954; Hu and Zheng 2001: 417 (new record; key; distribution; fig. 3, dorsal view of head; HN). Host plant unknown. Felisacus curvatus Hu and Zheng 2001: 417-419, fig. 2 (n. sp. description; dorsal view of body; paramere. Holotype and | paratype, HN. Deposited in DBNU). Host plant unknown. Felisacus insularis Miyamoto 1965; Hu and Zheng 2001: 419 (new record; key: identification; distribution; FJ, YN, GD, HN). Host plant unknown. Felisacus magnificus Distant 1904; Hu and Zheng 2001: 419 (new record; key; fig. 4, paramere; identification: distribution; FJ, YN, GD, HN). Host plant unknown. VOLUME 105, NUMBER 2 Parapachypeltis Hu & Zheng 2001 Type species: Parapachypeltis punctatus Hu and Zheng 2001 Parapachypeltis Hu and Zheng 2001: 422— 423 (new genus; description of general characters and the type species; identifi- cation). Parapachypeltis punctatus Hu and Zheng 2001: 423-424, fig. 5 (n. sp. description; dorsal view of body; dorsal and lateral view of pronotum; antennal segments; paramere. Holotype and 2 paratypes, GD. Deposited in DBNU). Host plant un- known. Ragwelellus Odhiambo 1962 Ragwelellus Odhiambo 1962; Hu and Zheng 2001: 424 (new record; identifi- cation). Ragwelellus rubrinus Hu and Zheng 2001: 424-245, fig. 6 (n. sp. description; dorsal view of body, paramere. Holotype and 9 paratypes, HN, YN, GD. Deposited in DBNU). Host plant unknown. Deraeocorinae Douglas & Scott 1865 Deraeocorini Douglas & Scott 1865 Alloeotomus Fieber 1858 Alloeotomus kerzhneri Qi and Bai 1994a: 458-459, figs. 1-5 (n. sp. description; distribution; male genitalia. Holotype, IM. Deposited in DBIMNU). Host plant unknown. Alloeotomus montanus Qi and Bai 1995c: 13, 15-16, figs. 1-8 (n. sp. description; key; distribution; hemelytra; male phal- lotheca, genitalia. Holotype and 7 para- types, IM. Deposited in DBIMNU). Host unknown. Bothynotus Fieber 1864 Bothynotus Fieber 1864; Qi and Bai 1994a: 460 (new record). Bothynotus pilosus (Boheman 1852); Qi and Bai 1994a: 460 (new record; distri- bution; IM). Host unknown. 431 Deraeocoris Kirschbaum 1856 Deraeocoris (Camptobrochis) brunneus Qi and Bai 1994a: 459-460, figs. 6—8 (n. sp. description; distribution; male genitalia. Holotype, IM. Deposited in DBIMNU). Host unknown. Deraeocoris (Camptobrochis) lutescens (Schilling 1837); Qi and Bai 1994a: 460 (new record; distribution; IM). Host un- known. Deraeocoris (Camptobrochis) onphoriensis Josifov 1992; Ma and Zheng 1997a: 3 (new record; distribution; GS, HEB, SX, JL). Host unknown. Deraeocoris (Camptobrochis) zoui Ma and Zheng 1997b: 20-23, figs. 1-3 (n. sp. de- scription; distribution; male genitalia. Holotype and | paratype, SC. Deposited in DBNU). Host unknown. Deraeocoris (Deraeocoris) guizhouensis Ma and Zheng 1997a: 1-3, figs. 1-7 (n. sp. description; distribution; male geni- talia. Holotype and 6 paratypes, GZ. De- posited in DBNU). Host unknown. Deraeocoris (Deraeocoris) morio (Bohe- man 1852); Qi and Bai 1994a: 460 (new record; distribution; IM). Host unknown. Deraeocoris (Deraeocoris) ventralis (Reu- ter 1904); Qi and Bai 1994a: 461 (new record; distribution; IM). Host unknown. Deraeocoris fujianensis Ma and Zheng 1998: 36-38, figs. 1-8 (n. sp. description; distribution; male genitalia. Holotype and 5 paratypes, FJ. Deposited in DBNU). Host unknown. Deraeocoris pallidicornis Josifoy 1983; Qi and Bai 1994a: 460 (new record; distri- bution; IM). Host unknown. Deraeocoris pseudokerzhneri Ma and Zheng 1998: 38—40, figs. 9-12 (n. sp. de- scription; distribution; male genitalia. Holotype and 18 paratypes, SC. Depos- ited in DBNU). Host unknown. Muirinae Hahn 1833 Mirini Hahn 1833 Allorhinocoris Reuter 1876 432 Allorhinocoris Reuter 1876; Lu 1994: 205 (new record). chinensis Lu 1994: 205— 208, figs. I—S (n. sp. description; distri- bution; key; male genitalia. Holotype and 6 paratypes, GS, SX, HEB. Deposited in DBNU). Host unknown. Allorhinocoris flavus Sahlberg 1878; Lu 1994: 205 (new record; distribution; key; XJ). Host plant unknown. Allorhinocoris Apolygus China 1941 Apolygus angustus (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 135 (Ly- gocoris (Apolygus)) (n. comb.; SC); Ker- zhner and Josifoy 1999: 62 (n. comb.). Host plant unknown. Apolygus sinicus (Kerzhner and Schuh 1995) (Lygocoris); Lu and Zheng 1996: 134-135 (Lygocoris (Apolygus) badius) (n. nom. and n. comb.); Kerzhner and Josifoy 1999: 67 (n. comb.). Host plant unknown. Apolygus castaneus (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 135 (Lygocoris (Apolygus)) (n. comb.; SC); Kerzhner and Josifov 1999: 62 (n. comb.). Host plant unknown. Apolygus concinnus (Wang and Zheng 1982) (Lygus); Lu and Zheng 1996: 135 (Lygocoris (Apolygus)) (n. comb.; FJ); Kerzhner and Josifov 1999: 63 (n. comb.). Host plant unknown. Apolygus curvipes (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 136 (Ly- gocoris (Apolygus)) (n. comb.; HB); Ker- zhner and Josifov 1999: 63 (n. comb.). Host plant unknown. Apolygus elegans (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 136 (Ly- gocoris (Apolygus)) (n. comb.; HN); Ker- zhner and Josifov 1999: 63 (n. comb.). Host plant unknown. Apolygus emeia (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 135 (Ly- gocoris (Apolygus)) (n. comb.; FJ, SC); Kerzhner and Josifov 1999: 63 (n. comb.). Host plant unknown. Apolygus eous (Poppius 1915) (Lygus); Lu PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and Zheng 1998c: 186 (n. comb.; brief description and note; TW). Host plant un- known. Apolygus evonymi (Zheng and Wang 1982) (Lygus); Lu and Zheng 1996: 135 (Ly- gocoris (Apolygus)) (n. comb.; SAX); Kerzhner and Josifov 1999: 63 (n. comb.). Host plant unknown. Apolygus fuhoshoensis (Poppius 1915) (Ly- gus); Lu and Zheng 1998c: 186 (n. comb.; brief description and note; TW). Host plant unknown. Apolygus fujianensis (Wang and Zheng 1982) (Lygus); Lu and Zheng 1996: 135 (Lygocoris (Apolygus)) (n. comb.; FJ); Kerzhner and Josifov 1999: 63 (n. comb.). Host plant unknown. Apolygus gleditsticola Lu and Zheng 1997a: 162-163, figs. 1—4 (n. sp. descrip- tion; distribution; male genitalia. Holo- type and 3 paratypes, HEN; | paratype, HB. Deposited in DBNU). Host plant un- known. Apolygus hainanensis (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 135 (Lygocoris (Apolygus)) (n. comb.; HN); Kerzhner and Josifov 1999: 64 (n. comb.). Host plant unknown. Apolygus kosempoensis (Poppius 1915) (Lygus); Lu and Zheng 1998c: 185-186, figs. 1-5 (n. comb.; brief description and note, male genitalia; TW). Host plant un- known. Apolygus liaoningensis (Lu and Zheng 1996) (Lygocoris (Apolygus)); Lu and Zheng 1996: 131-132, figs. 1-6 (n. sp. description; distribution; anterior and lat- eral view of head; hemelytra; male gen- italia. Holotype and 4 paratypes, LN. De- posited in DBNU. Host plant: Euvony- mus sp.); Kerzhner and Josifov 1999: 65 (n. comb.). Apolygus major (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 135 (Ly- gocoris (Apolygus)) (n. comb.; SC); Ker- zhner and Josifov 1999: 65 (n. comb.). Host plant unknown. Apolygus medionigritus Lu and Zheng 1997a: 163-165, figs. 5—8 (n. sp. descrip- VOLUME 105, NUMBER 2 tion; distribution; male genitalia. Holo- type and 58 paratypes, GS; 2 paratypes, SC. Deposited in DBNU). Host plant un- known. Apolygus mosaicus (Zheng and Wang 1982) (Lygus); Lu and Zheng 1996: 135 (Ly- gocoris (Apolygus)) (n. comb.; GD); Ker- zhner and Josifov 1999: 66 (n. comb.). Host plant unknown. Apolygus nigricans (Wang and Zheng 1982) (Lygocoris); Lu and Zheng 1996: 135 (Lygocoris (Apolygus)) (n. comb.; FJ); Kerzhner and Josifov 1999: 66 (n. comb.). Host plant unknown. Apolygus nigricostalis Lu and Zheng 1997a: 165-166, figs. 9-11 (n. sp. scription; distribution; male genitalia. Holotype and 6 paratypes, XZ. Deposited in IZAS). Host plant unknown. Apolygus ornatus (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 135 (Ly- gocoris (Apolygus)) (n. comb.; HUB); Kerzhner and Josifov 1999: 66 (n. comb.). Host plant unknown. de- Apolygus picturatus (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 136 (Lygocoris (Apolygus)) (n. comb.; GX); Kerzhner and 1999: (67 (nH: comb.). Host plant unknown. Apolygus triangulus (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 135 (Lygocoris (Apolygus)) (n. comb.; HUB); Kerzhner and Josifov 1999: 68 (n. comb.). Host plant unknown. Apolygus ulmi (Zheng and Wang 1983) (Ly- gus); Lu and Zheng 1996: 135 (Lygocoris (Apolygus) ulmicolus) (n. nom. and n. comb.; BJ); Kerzhner and Josifov 1999: 68. Host plant unknown. Apolygus wangi (Kerzhner and Schuh 1995) (Lygocoris), Lu and Zheng 1996: 135 (Lygocoris (Apolygus) signatus) (n. comb.; SC); Kerzhner and Josifov 1999: 68 (n. comb.). Host plant unknown. Apolygus yunnananus (Zheng and Wang 1983) (Lygus); Lu and Zheng 1996: 135 (Lygocoris (Apolygus)) (n. comb.; YN); Kerzhner and Josifov 1999: 68 (n. comb.). Host plant unknown. Josifov 433 Apolygus zhengianus Kerzhner and Schuh 1998: 171-172 (n. nom. for Lygus (Apol- ygus) marginatus Zheng and Wang 1983; SC); Kerzhner and Josifov 1999: 68 (n. comb.). Host plant unknown. Apolygus zizyphi Lu and Zheng 1997a: 166-168, figs. 12-15 (n. sp. description; distribution; male genitalia. Holotype and 4 paratypes, AH. Deposited in DBNU and TNHM respectively). Host plant: Z7- zyphus jujuba var. inermis (Bunge). Arbolygus Kerzhner 1979 Arbolygus ailaoensis Lu and Zheng 1998a: 82-83, figs. 1-5 (n. sp. description; key; list; distribution; male genitalia. Holo- type, YN. Deposited in DBNU). Host plant unknown. Arbolygus difficilis Lu and Zheng 1998a: 83-85, figs. 6-10 (n. sp. description; key; list; distribution; male genitalia. Holotype and 2 paratypes, SC; 9 paratypes, HUB; 2 paratypes, YN; 3 paratypes, XZ. De- posited in DBNU and IZAS respective- ly). Host plant unknown. Arbolygus glaber (Kerzhner 1987) (Lygo- corts (Arbolygus)); Miyamoto and Ya- sunaga 1989; Lu and Zheng 1998a: 80 (new record; list; distribution; SC. De- posited in DBNU). Host plant: Quercus dentate Thunberg. Arbolygus himalayicus Lu and Zheng 1998a: 80, 81, 86-88, figs. 16—19 (n. sp. description; key; list; distribution; male enitalia. Holotype and 7 paratypes. XZ; paratype, YN. Deposited in IZAS). Host plant unknown. Arbolygus longustus Lu and Zheng 1998a: 80, 81, 88-89, figs. 20—25 (n. sp. descrip- tion; key; list; distribution; male genita- lia. Holotype and 12 paratypes, HUB. Deposited in DBNU). Host plant un- known. Arbolygus picinus Lu and Zheng 1998a: 80, 81, 91-92, figs. 32—37 (n. sp. description; key; list; distribution; male genitalia. Ho- lotype and | paratype, YN. Deposited in DBNU). Host plant unknown. Arbolygus pronotalis (Lu and Zheng 1992) og r=) | 434 (Lygocoris (Arbolygus)); Lu and Zheng 1998a: 81 (n. comb.; list; key; distribu- tion; SC, HUN, FJ. Deposited in DBNU). Host plant unknown. Arbolygus renae Lu and Zheng 1998a: 80, 81, 89-91, figs. 26-31 (n. sp. description; key; list; distribution; male genitalia. Ho- lotype and | paratype, GS. Deposited in DBNU). Host plant unknown. Arbolygus tibialis Lu and Zheng 1998a: 80, 81, 93-94, figs. 38-43 (n. sp. description; key; list; distribution; male genitalia. Ho- lotype, SAX; 41 paratypes, SAX, NX, GS, HUB. Deposited in DBNU and HWAU respectively). Host plant un- known. Arbolygus wuzhiensis Lu and Zheng 1998a: 80, 81, 85—56, figs. 11-14 (n. sp. descrip- tion; key; list; distribution; male genita- lia. Holotype, HN. Deposited in DBNU). Host plant unknown. Arbolygus zhangi Lu and Zheng 1998a: 80, 81, 94-95, figs. 44-48 (n. sp. descrip- tion: key; list; distribution; male genita- lia. Holotype and 1 paratype, XZ. De- posited in IZAS). Host plant unknown. Castanopsides Yasunaga 1992 Castanopsides dasypterus (Reuter 1906) (Lygus); Lu and Zheng 1998: 80, 81 (Ar- bolygus) (n. comb.; list; distribution; HUB, HUN, SC, GZ, YN, XZ. Deposited in DBNU). Host plant: Hypericum sp. Castanopsides falkovitshi (Kerzhner 1979) (Lygocoris (Arbolygus)); Lu and Zheng 1998a: 80-81 (Arbolygus) (n. comb. and new record; list; key; distribution; HEB, FJ, SC. Deposited in DBNU). Host plant unknown. Castanopsides kerzhneri (Josifov 1985) (Lygocoris (Arbolygus)); Miyamoto and Yasunaga 1989; Lu and Zheng 1998a: 80 (new record; list; key; distribution; SC. Deposited in DBNU). Host plant: Quer- cus aliena Blume, Q. mongolicus Fischer. Castanopsides gotohi Yasunaga 1998a: 103, 106, figs. 11, 24-28 (n. sp. descrip- tion; key; distribution; male genitalia. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Holotype, TW. Deposited in HUES). Host plant unknown. Castanopsides montanus Yasunaga 1998a: 103; lid; figs: 145 33,34, 37; 39, 4\@: sp. description; key; distribution; male genitalia. Holotype, TW. Deposited in NSMT). Host plant unknown. Castanopsides taiwanus Yasunaga 1998a: 103, 111, figs. 43, 47, 48 (n. sp. descrip- tion; key; distribution; male genitalia. Holotype, TW. Deposited in NSMT). Host plant unknown. Castanopsides takaii Yasunaga 1998a: 103, 1055. figs: SOP 10) Ii lS e215 223i (esp: description; key; distribution; male geni- talia. Holotype, TW. Deposited in HUES). Host plant unknown. Cheilocapsidea Poppius 1915 Cheilocapsidea pura (Yasunaga 1995) (Carvalhopantilius); Yasunaga and Ker- zhner 1998: 88 (n. comb.; TW). Host plant unknown. Cheilocapsidea rufescens (Yasunaga 1995) (Carvalhopantilius); Yasunaga and Ker- zhner 1998: 88 (n. comb.; TW). Host plant unknown. Dichrooscytus Fieber 1858 Dichrooscytus helanensis Qi and Bai 1996b: 300-301, 303-305, figs. 10-14 (n. sp. description; distribution; male genitalia. Holotype and 42 paratypes, IM. Deposited in DBIMNU). Host plant: Sa- bina vulgaris Antoine. Heteropantilius Zheng & Liu 1992 Heteropantilius flavescens Yasunaga 1995a: 1 (n. sp. description; distribution. Holo- type, TW). Host plant unknown. Heteropantilius jinxiuensis Wang and Liu 2001: 320-322, figs. 1-3 (n. sp. descrip- tion; distribution; male genitalia. Holo- type and 3 paratypes, GX. Deposited in DBNU and IZAS). Host plant unknown. Eurystylopsis Poppius 1911 Eurystylopsis angustatus Zheng and Lu 1995: 7-8, figs. 1, 3, 5, 7, 10, 13 (um. sp. VOLUME 105, NUMBER 2 description; distribution; lateral view of head and pronotum; pubescence of anten- nal segment; male and female genitalia. Holotype and 4 paratypes, GS. Deposited in DBNU). Host plant unknown. Eurystylopsis hiripes Zheng and Lu 1995: 8-9, figs. 2, 4, 6, 9, 11, 14 (nm. sp. de- scription; distribution; lateral view of head and pronotum; pubescence of anten- nal segment; male and female genitalia. Holotype and 11 paratypes, GS. Depos- ited in DBNU). Host plant unknown. Parapantilius Reuter 1903 Parapantilius taiwanicus Yasunaga 1994: 688-691, fig. 4 (n. sp. description; distri- bution; key; vesica, posterior wall in an- terior view, and paramere. Holotype and 8 paratypes, TW. Deposited in HUESJ). Host plant unknown. Carvalhopantilius Yasunaga 1995 Type species: Carvalhopantilius purus Ya- sunaga 1995 Carvalhopantilius Yasunaga 1995b: 453-— 457, figs. 1-3 (new genus; description of general characters and the type species; identification). Carvalhopantilius purus Yasunaga 1995b: 452-455, figs. 1-2 (n. sp. description; distribution; dorsal view of body; lateral view of head-pronotum; male genitalia. Holotype, TW. Deposited in NSMT). Host plant unknown. Carvalhopantilius rufescens Yasunaga 1995b: 455—456, fig. 3 (n. sp. descrip- tion; distribution; sclerotized rings in ventral view, posterior wall of bursa co- pulatrix in anterior view. Holotype and | paratype, TW. Deposited in NSMT). Host plant unknown. Lygocorides Yasunaga 1991 Lygocorides affinis (Lu and Zheng 1997) (Lygocoris (Lygocorides)); Lu and Zheng, 1997b: 17-18, figs. 1-5 (n. sp. description; distribution; male genitalia. Holotype and | paratype, SAX. Depos- ited in DBNU). Host plant unknown. 435 Lygocorides rubricans Yasunaga 1996: 273 (n. sp. description; distribution; male genitalia. Holotype, TW. Deposited in HUES). Host plant unknown. Lygocoris Reuter 1875 Lygocoris (Lygocoris) calliger Lu and Zheng 2001: 123-125, fig. 1 (n. sp. de- scription; different views of male geni- talia. Holotype and 5 paratypes, YN. De- posited in DBNU). Host plant unknown. Lygocoris (Lygocoris) chengi Lu and Zheng 2001: 125-126, fig. 2 (n. sp. de- scription; different views of male geni- talia. Holotype and 25 paratypes, SC. De- posited in DBNU). Host plant unknown. Lygocoris (Lygocoris) diffusomaculatus Lu and Zheng 2001: 126-128, fig. 3 (n. sp. description; different views of male gen- italia. Holotype and 46 paratypes, GS, HUB. Deposited in DBNU, ZIRASP). Host plant unknown. Lygocoris (Lygocoris) dilutus Lu and Zheng 2001: 128-129, fig. 4 (n. sp. de- scription; different views of male geni- talia. Holotype and 23 paratypes, GS, HUB. Deposited in DBNU). Host plant unknown. Lygocoris (Lygocoris) ferrugineus Lu and Zheng 2001: 121-153, fig. 5 (n. sp. de- scription; different views of male and fe- male genitalia. Holotype and 35 para- types, YN. Deposited in DBNU). Host plant unknown. Lygocoris (Lygocoris) fuscoscutellatus (Reuter 1906) (Lygus striicornis var. fus- coscutellatus); Lu and Zheng 2001: 131— 132, fig. 6 (description; different views of male genitalia; YN). Host plant un- known. Lygocoris (Lygocoris) guangxiensis Lu and Zheng 2001: 132-134, fig. 7 (n. sp. de- scription; different views of male geni- talia. Holotype and 40 paratypes, GX. Deposited in DBNU, TNHM). Host plant unknown. Lygocoris (Lygocoris) idoneus (Linnavuori 1963) (Lygus); Lu and Zheng 2001: 134— 135, fig. 8 (new record; description and 436 distribution; different views of male gen- italia; GS, FJ, SC, YN). Host plant un- known. Lygocoris (Lygocoris) integricarinatus Lu and Zheng 2001: 136—137, fig. 9 (n. sp. description; different views of male gen- italia. Holotype and 3 paratypes, GS, SC. Deposited in DBNU). Host plant un- known. Lygocoris (Lygocoris) linnavuorti Lu and Zheng 2001: 137-138, fig. 10 (n. sp. de- scription; different views of male geni- talia. Holotype and 19 paratypes, YN. Deposited in DBNU). Host plant un- known. Lygocoris (Lygocoris) maculiscutellatus Lu and Zheng 2001: 139-141, fig. 11 (n. sp. description; different views of male and female genitalia. Holotype and 87 para- types, SC. Deposited in DBNU, TNHM, ZIRASP, USNM and HNHM respective- ly). Host plant unknown. Lygocoris (Lygocoris) rufiscutellatus Lu and Zheng 2001: 143-144, fig. 13 (n. sp. description; female genitalia. Holotype and | paratype, GS. Deposited in DBNU). Host plant unknown. Lygocoris (Lygocoris) rufomedialis Lu and Zheng 2001: 144-146, fig. 14 (n. sp. de- scription; different views of male geni- talia. Holotype, YN. Deposited in DBNU). Host plant unknown. Lygocoris (Lygocoris) sichuanicus Lu and Zheng 2001: 148-149, fig. 16 (n. sp. de- scription; different views of genitalia. Holotype and 5 paratypes, SC. Deposited in DBNU and TNHM respectively). Host plant unknown. Lygocoris (Neolygus) bimaculatus Lu and Zheng 1996: 134-135, figs. 12—16 (n. sp. description; distribution; male and female genitalia. Holotype and 22 paratypes, SC. Deposited in DBNU). Host plant un- known. Lygocoris (Neolygus) bipuncticollis (Pop- pius 1915) (Lygus); Lu and Zheng 1998c: 186-187, fig. 6 (n. comb.; brief descrip- tion and note; right paramere; TW). Host plant unknown. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Lygocoris (Neolygus) chinensis Lu and Ya- sunaga 1994: 100—102, figs. 5—10 (n. sp. description; distribution; male and female genitalia. Holotype and 17 paratypes, IM, HLJ. Deposited in DBNU and HUESJ). Host plant unknown. Lygocoris (Neolygus) disciger (Poppius 1915) (Lygus); Lu and Zheng 1998c: 187-188, figs. 7-11 (n. comb.; brief de- scription and note; male genitalia; TW). Host plant unknown. Lygocoris (Neolygus) elongatulus Lu and Zheng 1996: 205-206, 208-209, figs. 1— 6 (n. sp. description; distribution; male genitalia and paramere; female genitalia. Holotype and 3 paratypes, GS. Deposited in DBNU). Host plant unknown. Lygocoris (Neolygus) gansuensis Lu and Zheng 1996: 206-207, 209, figs. 7-10 (n. sp. description; distribution; male geni- talia. Holotype, GS. Deposited in DBNU). Host plant unknown. Lygocoris (Neolygus) lativerticis Lu and Wang 1997: 402—405, figs. 1-6 (n. sp. description; distribution; male genitalia. Holotype and 39 paratypes, GS. Depos- ited in DBNU). Host plant unknown. Lygocoris (Neolygus) rufilorum Lu and Zheng 1998b: 3, figs. 5—8 (n. sp. descrip- tion; distribution; male genitalia. Holo- type, FJ, GX, YN. Deposited in DBNU). Host plant unknown. Lygocoris (Neolygus) salicicola Lu and Zheng 1998b: 6, figs. 19-24 (n. sp. de- scription; distribution; male genitalia. Holotype, GS. Deposited in DBNU). Host plant unknown. Lygocoris (Neolygus) simillimus Lu and Zheng 1998b: 8, figs. 25-28 (n. sp. de- scription; distribution; male genitalia. Holotype, SC. Deposited in IZAS). Host plant unknown. Lygocoris (Neolygus) tilianus Lu and Zheng 1996: 132-134, figs. 7-11 (n. sp. description; distribution; male and female genitalia. Holotype and 5 paratypes, NX. Deposited in DBNU). Host plant: Tilia chinensis Maximowicz. Lygocoris (Neolygus) v-nigrum (Poppius VOLUME 105, NUMBER 2 1915) (Lygus); Lu and Zheng 1998c: 187 (n. comb.; brief description and note; TW). Host plant unknown. Lygocoris (Neolygus) wuyiensis Lu and Zheng 1998b: 2, figs. 1—4 (n. sp. descrip- tion; distribution; male genitalia. Holo- type, FJ, ZJ. Deposited in DBNU). Host plant unknown. Lygocoris (Neolygus) xizangensis Lu and Zheng 1998b: 9, figs. 19-32 (n. sp. de- scription; distribution; male genitalia. Holotype, XZ. Deposited in IZAS). Host plant unknown. Lygocoris (Neolygus) yulongensis Lu and Zheng 1998b: 5, figs. 13-18 (n. sp. de- scription; distribution; male genitalia. Holotype, YN. Deposited in DBNU). Host plant unknown. Lygocoris (Neolygus) zhengi Lu and Yasun- aga 1994: 99-100, figs. 1-4 (n. sp. de- scription; distribution; male genitalia. Holotype, SC. Deposited in DBNU). Host plant unknown. Macrolygus Yasunaga 1992 Macrolygus viridulus Yasunaga 1992; Lu and Zheng 1997c: 286 (new record; HUB). Host plant unknown. Metasequoiamiris Schwartz 1995 Type species: Metasequoiamiris carvalhot Schwartz 1995 Metasequoiamiris Schwartz 1995: 401— 408, figs. 1-19 (new genus; description of general characters and the type spe- cies; identification). Metasequoiamiris carvalhoi Schwartz 1995: 401—408 figs. 1-19 (n. sp. descrip- tion; distribution; male genitalia and par- amere. Holotype, HUB). Host plant: Ce- phalotaxus fortunet Hooker, Metasequoia glyptostroboides Hu and Cheng, Torreya sp. Orientomiris Yasunaga 1997 Orientomiris chinensis (Li and Zheng 1991) (Megacoelum), Yasunaga 1998b: 63-70 (n. comb.; FJ). Host plant unknown. Orientomiris pronotalis (Li and Zheng 437 1991) (Megacoelum); Yasunaga 1998b: 63-70 (n. comb.: ZJ). Host plant un- known. Orientomiris pseudopronotalis (Li and Zheng 1991) (Megacoelum),; Yasunaga 1998b: 63-70 (n. comb.; FJ). Host plant unknown. Orientomiris rubripedus (Li and Zheng 1991) (Megacoelum), Yasunaga 1998b: 63-70 (n. comb.; FJ). Host plant un- known. Orientomiris tenuicornis (Li and Zheng 1991) (Megacoelum); Yasunaga 1998b: 63-70 (n. comb.; GD). Host plant un- known. Orientomiris yunnananus (Li and Zheng 1991) (Megacoelum); Yasunaga 1998b: 63-70 (n. comb.; YN). Host plant un- known. Orientomiris zoui (Li and Zheng 1991) (Megacoelum), Yasunaga 1998b: 63-70 (n. comb.; YN). Host plant unknown. Phytocoris Fallén 1814 Phytocoris elongatulus (Bai and Chen 1996); Aukema 1999 (n. nom. for Phy- tocoris elongotus Bai and Chen 1996: 182, 185, figs. 1-3 (n. sp. description; distribution; male genitalia. Holotype, al- lotype and 5 paratypes, GS. Deposited in DBIMNU)). Host plant unknown. Phytocoris exohataensis Xu and Zheng 2001: 263-264, figs. 5, 32—40 (n. sp. de- scription; distribution; hemelytra; male genitalia. Holotype, YN. Deposited in DBNU and DPPBAN). Host plant un- known. Phytocoris macer Xu and Zheng 1997: 129-134 figs. 8-13 (n. sp. description; distribution; male genitalia. Holotype, NX, SAX. Deposited in DBNU). Host plant unknown. Phytocoris ningxiaensis Bai and Chen 1996: 182-183, 185, figs. 4—7 (n. sp. de- scription; distribution; male genitalia. Holotype and allotype, NX. Deposited in DBIMNU). Host plant unknown. Phytocoris nitrariae Xu and Zheng 1997: 129-134 figs. 1-7 (n. sp. description; dis- 438 tribution; male genitalia. Holotype, NX. Deposited in DBNU). Host plant un- known. Phytocoris wolongensis Xu and Zheng 2001: 257-259, figs. 2, 7-14 (n. sp. de- scription; distribution; hemelytra; male genitalia. Holotype, SC. Deposited in DBNU). Host plant unknown. Phytocoris wudingensis Xu and Zheng 2001: 260-263, figs. 4, 25-31 (n. sp. de- scription; distribution; hemelytra, prono- tum; male genitalia. Holotype, YN. De- posited in DBNU). Host plant unknown. Phytocoris yongpinganus Xu and Zheng 2001: 259-260, figs. 3, 15—24 (n. sp. de- scription; distribution; hemelytra, prono- tum; male genitalia. Holotype, YN. De- posited in DBNU). Host plant unknown. Poppiocapsidea Yasunaga 1998 Poppiocapsidea clypealis (Poppius 1915) (Megacoelum); Yasunaga 1998b: 63—70 (n. comb.; TW). Host plant unknown. Sabactus Distant 1910 Sabactus sauteri (Poppius 1912) (Lygus): Lu and Zheng 1998c: 188, figs. 12—16 (n. comb.; brief description and note; male genitalia; TW). Host plant unknown. Tinginotum Kirkaldy 1902 Tinginotum pini Kulik 1965; Lu and Zheng 1997c: 289 (new record; SC). Host plant unknown. Stenodemini China 1943 Stenodema Laporte 1833 Stenodema (Stenodema) deserta Bai and Chen 1994: 86-87, figs. 1-6 (n. sp. de- scription; distribution; dorsal view head and pronotum; antennae and leg; male genitalia. Holotype, allotype and 55 para- types, IM. Deposited in DBIMNU). Host plant unknown. Stenodema (Stenodema) crassipes Kirit- shenko 1931; Bai and Chen 1994: 86 (new record; IM). Host plant unknown. Stenodema (Stenodema) mongolica Bai and Chen 1994: 88-89, figs. 7-12 (n. sp. de- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON scription; distribution; dorsal view head and pronotum; antennae and leg; male genitalia. Holotype, allotype and 3 para- types, IM. Deposited in DBIMNU). Host plant unknown. Stenodema (Stenodema) sericanus Fieber 1861; Bai and Chen 1994: 86 (new re- cord; IM). Host plant unknown. ACKNOWLEDGMENTS We are very grateful to Prof. Zheng Ley1 and Dr. Liu Guoqing, Department of Biol- ogy, Tiejin Nankai University, and Prof. Ma Enbo, Department of Biology, Shanxi Uni- versity, for providing us with copies of their published papers. LITERATURE CITED Bai, N. and B. Y. Qi. 1999. pp. 56—64. Jn Insects of Inner Mongolia. Huhe-hot: Inner Mongolian Peo- ple’s Press. Bai, N. and J. Chen. 1994. A preliminary study of Stenodema Laporte (Hemiptera: Miridae) from In- ner Mongolia, China. Entomotaxonomia 16(2): 85-90. . 1996. Two new species and Chinese record of Miridae from Inner Mongolia, China (Hetreoptera: Miridae). Acta Entomologica Sinica 39(2): 182— 185. Bai, N. and Y. Q. Yang. 1994. Three new species and new Chinese records of Miridae from Inner Mon- golia, China (Hemiptera: Miridae). Zoological Re- search 15:17—22. Carvalho, J. C. M. 1952. On the major classification of the Miridae (Hemiptera) (with keys to the sub- families and tribes and a catalogue of the world genera). Anais da Academia Brasileira de Ciincias 24:31-110. . 1956. Heteroptera: Miridae. /n Insects of Mi- cronesia. Bernice P. Bishop Museum, Honolulu 7(1). 100 pp. . 1957. A catalogue of the Miridae of the world, Part I. Arquivos do Museu Nacional, Rio de Ja- neiro 44: 1-158. 1958a. A catalogue of the Miridae of the world, Part II. Arquivos do Museu Nacional, Rio de Janeiro 45: 1-216. . 1958b. A catalogue of the Miridae of the world, Part III. Arquivos do Museu Nacional, Rio de Janeiro 47: 1-161. . 1959. A catalogue of the Miridae of the world, Part ITV. Arquivos do Museu Nacional, Rio de Ja- neiro 48: 1-384. . 1960. A catalogue of the Miridae of the world, VOLUME 105, NUMBER 2 Hu, Part V. Arquivos do Museu Nacional, Rio de Ja- neiro 51: 1-191. Q. and L. Y. Zheng. 1999a. New species of genus Mansoniella Poppius from China (Hemiptera: Miridae: Bryocorinae). Acta Zootaxonomia Sinica 24(2): 159-170. . 1999b. A new species of genus Pschypeltis Signoret from China (Hemiptera: Miridae: Bry- ocorinae). Acta Scientiarum Naturalium Univer- sitatis Nankaiensis 32(1): 1-3. . 2000. A revision of Chinese species of Bry- ocoris Fallén (Hemiptera: Miridae). Acta Zootax- onomia Sinica 25(3): 241—167.?? —.. 2001. The Monaloniina from Mainland China (Hemiptera: Miridae: Bryocorinae). Acta Zootax- onomica Sinica 26(4): 414—430. Kerzhner, I. M. 1997. Notes on taxonomy and nomen- clature of palaearctic Miridae (Heteroptera). Zoos- ystematica Rossica 5(2): 245-248. Kerzhner, I. M. and M. Josifov. 1999. Miridae. Jn Auk- ma, B. and C. Rieger, eds. Catalogue of the Het- eroptera of the Palaearctic Region. 3. Cimicomor- pha II. Amsterdam. 577 pp. Kerzhner, I. M. and R. T. Schuh. 1998. Replacement Liu, Liu, names for junior homonyms in the family Miridae (Heteroptera). Zoosystematica Rossica 7: 171— 172. G. Q. 1999. Notes on Chinese species of genus Pseudoloxops (Heteroptera: Miridae). Acta Scien- tiarum Naturalium Universitatis 32(4): 51-53,57. G. Q. and L. Y. Zheng. 1994. Two new species of genus Pseudoloxops Westwood from China (Heteroptera: Miridae). Reichenbachia 30: 119— 122. Nankaiensis ———.. 1999a. On the Chinese Species of Cyllecoris Hahn (Hemiptera: Miridae: Orthotylinae), pp. 102-109. In Zhang Y. L., ed. Systematic and Fau- nistic Research on Chinese Insects. Proceeding of 5" National Congress of Insect Taxonomy, China Agriculture Press. ———.. 1999b. New species on the Zanchius Distant Liu, Tou; from China (Hemiptera: Miridae). Acta Zootax- onomia Sinica 24(4): 388-392. G. Q. and R. J. Zhao. 1999. New species of genus Coridromius Signoret from China (Heteroptera: Miridae). Acta Zootaxonomia Sinica 24(1): 55— 58. N. 1994. The genus Allorhinocoris Reuter from China (Hemiptera: Miridae). Entomolgia Sinica 1(3): 205-208. Lu, N. and H. J. Wang. 1996. Two new species of the genus Lygocoris Reuterfrom Gansu Province, China (Heteroptera: Miridae). Acta Zootaxonomia Sinica 21(2): 205-207. . 1997. A new species of the genus Lygocoris Reuter from Gansu, China (Heteroptera: Miridae). Acta Entomologica Sinica 40(4): 402-405. Lu, 439 N. and L. Y. Zheng. 1996. New species of genus Lygocoris Reuter from China (Insecta : Heterop- tera : Miridae). Reichenbachia 31(24): 131-137. . 1997a. Four new speceis of the genus Apol- ygus China from China (Insecta: Hemiptera: Mir- idae). Acta Zootaxonomia Sinica 22(2): 162-168. . 1997b. One new species of the subgenus Ly- gocorides Yasunaga of genus Lygocoris Reuter from China (Insecta: Heteroptera: Miridae). Rei- chenbachia 32(1—24):17-18. ———.. 1997c. Heteroptera: Miridae, pp. 272—290. In Insects of the Three Gorge Reservoir Area of Yangtze River, Science Press, Beijing. . 1998a. A taxonomic study on the genus Ar- bolygus (Heteroptera: Miridae) from China. En- tomotaxonomia 20(2): 79-96. . 1998b. New species of the genus Lygocoris (subgenus Neolygus) from China (Heteroptera: Miridae). Entomologische Berichten, Amsterdam 58: 1-10. . 1998c. Identity of some *Lygus” species de- scribed from Taiwan by B. Poppius (Heteroptera: Miridae). Tijdschrift voor Entomologie 140: 185— 190. . 2001. Revision of Chinese Species of Lygo- coris (subg. Lygocoris) Reuter (Hemiptera: Muiri- dae: Mirinae). Acta Zootaxonomica Sinica 26(2): 121-153. Lu, N. and T. Yasunaga. 1994. Two new species of the Ma, subgenus Neolygus Knight of the genus Lygocoris Reuter from China (Heteroptera: Miridae). Bulle- tin of the Biogeographyical Society of Japan 49(2): 99-103. C. J. and L. Y. Zheng. 1997a. A new species of the genus Deraeocoris Kirschhaum from China (Hemiptera: Miridae: Deraeocorinae). Acta Scien- tiarum Naturalium Universitatis Nankaiensis 30(4): 1-4. . 1997b. A new species of the genus Deraeo- coris Kirschhaum and the first description of the male of Deraeocoris alticallus Hsiao 1941 (Hem.: Miridae: Deraeocorinae). Wuyi Science 13: 20— 23: 1998. New species of genus Deraeocoris Kirschbaum from China (Hemiptera: Miridae: Deraeocorinae). Acta Zootaxonomia Sinica 23(1): 36-40. Qi, B. Y. and N. Bai. 1994a. New and newly recorded species of Deraeocorinae from Inner Mongolia, China (Hemiptera: Heteroptera: Miridae). Acta Zootaxonomica Sinica 19(4): 458—464. . 1994b. New species and new records of Psal- lus Fieber from China (Hemiptera: Miridae). Acta Entomologica Sinica 37(3): 341-343. . 1995a. A brief note on bugs of the genus A/- loeotomus Fieber from northern China, with de- scription of a new species (Insecta: Hemiptera: 440 Heteroptera: Miridae). Reichenbachia 31(3): 13— 16. . 1995b. A Preliminary study on genus Comp- sidolon Reuter 1899 from north China (Heterop- tera: Miridae: Phylinae). Entomologia Sinica 2(3): 225-227. . 1995c. Description of two new taxa of Miri- dae from northern China (Hemiptera: Heterop- tera). The Entomologist 114(3&4): 143-145. . 1995d. A declaration of changing name for Psallus (phylidea) kerzhneri Qi and Nonnaizab 1994. Acta Entomologica Sinica 38(2): 256. . 1996a. Chinese records of predacouse mirids from Inner Mongolia (Heteroptera: Miridae). Jour- nal of Inner Mongolia Normal University (Natural Science Edition.) 2: 49-53. . 1996b. A new genus and two new species of Miridae from China (Hemiptera: Miridae). Acta Entomologica Sinica 39(3): 298-305. . 1996c. New and little-known species of genus Solenoxyphus Reuter from northern China (He- miptera: Miiridae). Entomologia Sinica 3(4): 293— 299) 1997. New and little-known species of the tribe Phylini Douglas & Scott from China (Insec- ta: Hemiptera: Heteroptera: Miridae). Reichenba- chia 32(3): 9-15. Schuh, R. T. 1995. Plant Bugs of the world (Insecta : Heteroptera : Miridae), Systematic Catalog, Dis- tributions, Host List, and Bibliography. The New York Entomological Society, New York, 1,329 pp. Schuh, R. T. and J. A. Slater. 1995. True Bugs of the World (Hemiptera : heteroptera) Classification and Natural History, New York 169-181. Schwartz, M. D. 1995. Metasequoamiris carvalhoi, a new genus and species of conifer-inhabiting Mir- ini from China (Heteroptera: Miridae: Mirinae). Proceedings of the Entomological Society of Washington 97(2): 401—408. Tian, R. L. and N. Z. Bai. 1998. A preliminary study on genus Orthotylus Fieber from Inner Mongolia (Heteroptera: Miridae). Journal of Inner Mongolia Normal University (Natural Science Edition) 27(2): 123-128. . 1999. Two new species of Miridae from Mt. Helan of Inner Mongolia Autonomous Region, China. Acta Entomologica Sinica 42(1): 66-69. Wang, H. J. and G. Q. Liu. 2001. A new species of the Heteropantilius trom China (Heteroptera: Mir- idae: Mirinae). Acta Zootaxonomica Sinica 16(3): 320-322. Xu, B. H. and L. Y. Zheng. 1997. Two new species of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the genus Phyrocoris Fallén (Heteroptera: Miri- dae) from China. Entomologica Sinica 4: 129— 134. . 2001. Four new species of Phytocoris Fallén from China (Hemiptera: Miridae). Acta Zootax- onomica Sinica 26(3): 257-265. Yasunaga, T. 1994. Review of the Sino-Japanese plant bugs genus Parapantilius Reuter (Heteroptera, Miridae), with description of a new species from Taiwan. Japanese Journal of Entomology 62(4): 683-691. . 1995a. Discovery of the genus Heteropantil- ius Zheng et Liu from Taiwan, with description of a new species (Insecta: Heteroptera: Miridae). Journal of the Kokkaido University of Education Section II B 46(1): 1-5. . 1995b. A new genus of mirine plant bug, Car- valhopantilius, with two new species from Taiwan (Heteroptera, Miridae). Proceedings of the Ento- mological Society of Washington 97(2): 452—457. . 1996. Review of Lygocorides Yasunaga (Het- eroptera:Miridae). Tijdschrift voor Entomologie 139(2): 267-275. . 1998a. Revision of the mirine genus Castan- opsides Yasunaga from the eastern Asia (Heter- optera: Miridae). Entomologica Scandinavica 29(1): 99-119. . 1998b. Revision of the mirine genus Creon- tiades Distant and allies from Japan (Heteroptera: Miridae). Part Il. Neomegacoelum gen. n., and exotic new taxa, new synonymy and new combi- nations. Entomological Science 1: 63-70. Zhao, R. J. 1997. A record of four genera of Miridae in north-west part of China (Heteroptera: Miri- dae). Journal of Shanxi University (Natural Sci- ence Edition) 20(1): 99-100. Zhao, R. J. and C. A. Li. 1996. Faunistic structure of Miridae in northern-west part of China (Heterop- tera: Miridae). Journal of Shanxi University (Nat- ural Science Edition) 19(3): 352-354. Zheng, L. Y. 1995. A list of the Miridae (Heteroptera) recorded from China since J.C.M. Carvalho’s “World Catalogue.” Proceeding of Entomological Society of Washington 97(2): 458-473. Zheng, L. Y. and X. M. Li. 1996. A new species of the genus Phoenicocoris Reuter (Hemiptera: Mir- idae) from China. Entomotaxonomia 18(2): 103— 104. Zheng, L. Y. and S. Z. Ren. 1996. Hemiptera: Nabidae, Miridae, Lygaeidae, Rhopalidae, Pentatomidae, pp. 43-52. Jn Kala Kunlun Montains—lInsects from Kunlun Montains. Science Press, Beijing. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 441-446 FLEA BEETLES (COLEOPTERA: CHRYSOMELIDAE) OCCURRING ON AMARANTHUS RETROFLEXUS L. IN ERZURUM PROVINCE, TURKEY, AND THEIR POTENTIAL AS BIOLOGICAL CONTROL AGENTS IRFAN ASLAN, HIKMET OzBEK, AND ALEXANDER KONSTANTINOV (IA, HO) Atatiirk University, Agricultural Faculty, Department of Plant Protection, 25240 Erzurum, Turkey (e-mail: aslanir@hotmail.com); (AK) Systematic Entomology Laboratory, PSI, Agricultural Research Service U.S. Department of Agriculture, % Na- tional Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: akonstan @sel.barc.usda.gov) Abstract.—Species composition and abundance of flea beetles (Coleoptera, Chryso- melidae) associated with Amaranthus retroflexus L. (Amaranthaceae) were studied in Er- zurum province, Turkey, to find potential candidates for biological control of this weed. Insects were collected by sweeping net and aspirator on plant foliage at eight localities four times during the growing season. During three years of exploration, ten species of flea beetles were collected Chaetocnema breviuscula (Faldermann), Ch. concinna (Mar- sham), Ch. hortensis (Geoffroy), Ch. tibialis (liger), Longitarsus longipennis Kutschera, L. pellucidus Foudras, Phyllotreta atra (FE), Ph. cruciferae (Goeze), Ph. nigripes (FE), and Ph. vittula (Redtenbacher). In all localities Ch. tibialis was more abundant than other species. Simple feeding tests in petri dishes showed that only the Chaetocnema species were actually feeding on leaves of Amaranthus retroflexus. Analysis of flea beetle species associated with this plant shows that it is probably not native to Turkey. Key Words: Amaranthus retroflexus L. (Amarantha- ceae), or redroot pigweed, occurs through- out much of the world, including Europe, North and South America, Asia, Africa and Australia. In North America it is common from Canada to Mexico, and from the At- lantic to the Pacific coasts (Spencer 1957). It is a summer annual, commonly found in cultivated lands such as fields, gardens, and orchards; fallow land, stream valleys, beaches and streambanks, prairie ravines, roadsides, fence rows, and waste places (Roland and Smith 1969). Amaranthus re- troflexus is harmful to livestock because it affects the kidneys of swine when animals consume large quantities of fresh material (Wohlgemuth et al. 1987), and it is consid- flea beetles, biological control, Amaranthus retroflexus L., Chaetocnema ered an important exotic weed in North America (King 1966). However, the native range of this plant is unclear. Some authors believe it was introduced into the United States in the early 18th century (King 1966) and later to Eastern Europe and Russia (Ni- kitin 1983), but Auld and Medd (1987) con- sidered it to be native to North America. Different sources provide opposing ideas on the history of its distribution. For example, according to Haughton (1978) it is native to Spain, from which it was brought to the New World by conquistadors. Bermejo and Gonzalez (1994) suggest that the same con- quistadors transported it in the opposite di- rection, from South America to Spain. Three amaranth species occur in Turkey, 442 Amaranthus blitoides S. Wats, A. graeci- zans L., and A. retroflexus. However, only the latter species is common in Erzurum Province (Baytop 1994). Amaranthus retro- flexus grows from 15 cm to 100 cm tall and becomes particularly abundant in vegetable fields (Baytop 1994). For this reason it is considered one of the most serious weeds in Turkey (Baytop 1994) as well as in Eu- rope. Therefore it was chosen for biological control research within the framework of European Cooperation in the Field of Sci- entific and Technical Research (Miiller- Scharer 1993, Schroeder et al. 1993). Most of the biological control efforts re- garding Amaranthus retroflexus are concen- trated on the group of leaf beetles called flea beetles. It is the largest group within leaf beetles and is distributed throughout the world. The adults feed on the foliage of herbaceous plants, bushes and trees from a wide range of angiosperm families, as well as some gymnosperms. Larvae live in soil, plant tissues, or leaves. Many flea beetles cause severe damage to cultivated plants (Onat 1993; Naibo 1974; Kostromitin 1978: Aslan and Ozbek 1998, 2000), however many other species have been successfully used as biological control agents of noxious weeds (Konstantinov and Vandenberg 1996). Aslan et al. (1999) and Aslan and Warchalowski (2001) recorded 244 flea beetle species and subspecies belonging to 21 genera in Turkey. Of these species, 21 cause extensive damage to various weeds in the vicinity of Erzurum (Aslan and Ozbek 1999). Relatively few flea beetle species are recorded on Amaranthus retroflexus in Eu- rope (Doguet 1994, Cagan et al. 2000). The goal of our study was to collect flea beetles occurring on Amaranthus retroflex- us in Erzurum Province of Turkey, assess their abundance, and provide information on their distribution and host range. Pres- ence or absence of flea beetles limited in their host range to Amaranthus retroflexus would add valuable data to the ongoing dis- cussion on the native range of this weed. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MATERIALS AND METHODS Erzurum Province is located in the East- ern Anatolia region of Turkey at the aver- age altitude of 1,850 m. It is a montane country with a variety of climatic condi- tions. A continental climate with hot and relatively dry summer and cold snowy win- ter dominates the eastern and western parts of Erzurum province at the altitude of 1500-1900 m. Northeastern parts of the province with altitudes close to 700 m have a less continental climate. Field surveys of flea beetles associated with Amaranthus retroflexus were carried out in the first week of June, July, August and September at eight localities in differ- ent climatic regions of Erzurum province. Insects were collected by sweep net and by aspiration from plant foliage of 20 random- ly chosen plants. Collected insects were taken to the laboratory, where they were sorted and identified. At the same time, ten adults of each species were put in petri dishes with leaves of A. retroflexus. During the following ten days petri dishes were checked for leaf damage caused by feeding. RESULTS Adults of the following species were col- lected on Amaranthus retroflexus: Chaeto- cnema breviuscula (Faldermann), Ch. con- cinna (Marsham), Ch. hortensis (Geoffroy), Ch. tibialis (lliger), Longitarsus longipen- nis Kutschera, L. pellucidus Foudras, Phyl- lotreta atra (F.), Ph. cruciferae (Goeze), Ph. nigripes (F), and Ph. vittula (Redtenbach- er). Chaetocnema and Phyllotreta species were predominant in almost every locality. Chaetocnema species were found in all observed localities. The most common among them was Ch. tibialis accounting for 78.8% of all flea beetles. In the northeastern towns of the province, Tortum, Oltu, Olur and Renkaya, Ch. tibialis and Ch. concinna were more abundant than in the eastern and western towns of the province. Chaetocne- ma hortensis and Ch. were more abundant in the eastern and western breviuscula VOLUME 105, NUMBER 2 Table 1. Erzurum Province during 1998-2000. 443 Number of Chaetocnema breviuscula adults on Amaranthus retroflexus at different localities of 1998 1999 2000 Localities June July August Sept June July August Sept June July August Sept Arkale 1] 14 17 2 5 12 i19/ 1 15 12 11 2 Dumlu 2 1 7 l 2 b) 4 2 6 ) 7 0 Ilica Y) z 8 2 8 4 o 1 1] 6 12 4 Olur 2 (0) 5 0 3 | 3 ] | 3 2 0 Oltu 0 2 1 0) 1 | 2 0 3 2 2 l Pasinler 4 3 6 | 2 | 4 1 3 4 6 2 Renkaya 0 2 I 0) l 0 3 1 } 0 2 0) Tortum 3 1 4 | 2 l 2 0 2 1 3 0 localities than in the northeastern parts of Erzurum Province. The number of Chae- tocnema species collected in different lo- calities of Erzurum Province is shown in Tables 1—4. Phyllotreta species were sec- ond in abundance after Chaetocnema, with Phyllotreta atra being the most abundant species of this genus (with 38.81% of in- dividuals). Longitarsus species were least abundant with L. pellucidus being the most common species of this genus. The highest number of L. pellucidus was observed in Pasinler and Arkale. Very few specimens of L. longipennis were found in some locali- ties. Laboratory tests showed that only Chae- tocnema species (Ch. breviuscula (Falder- mann), Ch. concinna (Marsham), Ch. hor- tensis (Geoffroy), and Ch. tibialis (Mliger)) fed on A. retroflexus. The adults of other species did not feed on the leaves of A. re- troflexus. Table 2. zurum Province during 1998-2000. DISCUSSION Chaetocnema tibialis is known to feed on amaranthus plants in Europe (Nonveiller 1960, 1978) and in Turkey (Aslan 1997). Chaetocnema Ch. and Ch. tibialis are serious pests of sugar beet (Beta vulgaris L.) both in Turkey (As- lan and Ozbek 1998, 2000; Yildirim and Ozbek 1992) and in some European coun- tries (Gruev and Tomov 1986, Neves 1983, Slavchev 1984, Rimsa and Konecny 1983, Cooke 1992, Mostovaya 1994). However, our survey showed large numbers of them actively feeding on Amaranthus retroflexus causing significant damage to the plant. Chaetocnema hortensis commonly feeds on Gramineae species in Turkey (Aslan 1997) and some European countries (Mohr 1966, Gruev and Tomov 1986). Our survey con- firms that it also feeds on Amaranthus re- troflexus. It is important to note that all breviuscula, concinna Number of Chaetocnema concinna adults on Amaranthus retroflexus at differnet localities of Er- 1998 1999 2000 Localities June July August Sept June July August Sept June July August Sept Arkale 12 d ils) 2 1S 12 17 2 13 21 12 2 Dumlu il) 8 il9/ 1 16 19 25 5 11 9 27, 0 Thica 11 5 16 4 8 14 23 3 8 12 21 3 Olur 32 22 37 2 15 11 29 8 23 15 32 5 Oltu 28 22 38 5 24 16 37 4 18 13 18 3 Pasinler 16 14 16 qf 22 11 32 S) 13 17 26 5 Renkaya 25 18 32 I 24 33 28 5 22 15) 27 4 Tortum 23 14 31 | 12 15 29 2 24 19 32 0 444 Table 3. zurum Province during 1998—2000. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Number of Chaetocnema hortensis adults on Amaranthus retroflexus at different localities of Er- 1998 1999 2000 Localities June July August Sept June July August Sept June July August Sept. Arkale 10 7 25 3 12 8 17 2 11 6 8 1 Dumlu 4 3 7 1 5 3 7 3 8 4 17 5 Ilica 13 7 12 4 8 7 Y) 2; 5 3 1] 6 Olur 10) 2 >) | 2 1 4 3 6 8 2 Oltu 3 0) 2 0) 4 2 1 2 3 2 10) Pasinler 1] 5 16 7 12 6 14 J 1] ql 16 4 Renkaya 3 5) 11 l 5) 4 8 0 3 5 6 l Tortum 0 2 5) 10) 2 | 9 1 2 8 5 2 aforementioned Chaetocnema species are widely polyphagous, feeding on plants from many families (Lopatin 1977). They also have wide geographic ranges throughout al- most the entire Palearctic region (Konstan- tinov 1988). The most common host plant of Longi- tarsus longipennis and L. pellucidus is Con- volvulus arvensis (Gruev and Tomov 1986, Warchalowski 1996, Aslan 1997). Although commonly collected on A. retroflexus in Er- zurum Province, they did not feed on this plant under laboratory conditions. Their presence on A. retroflexus is temporary and accidental. It happened only because of close coexistence of Convolvulus arvensis and amaranth. Convolvulus arvensis often found climbing on A. retroflexus in many localities of Erzurum Province. All Phyllotreta species generally feed on Cruciferae in Turkey (Aslan and Ozbek 1988, 2000) and in Europe (Mohr 1966, Was Gruev and Tomoy 1986, Kostromitin 1978). Phyllotreta vittula is known as important pest of crucifers, as well as sugar beets, ce- reals (Naibo 1974), and maize (Szoeke 1997). All collected Phyllotreta species are widely oligophagous or polyphagous. In laboratory tests they did not feed on A. re- troflexus. None of the species collected on Ama- ranthus retroflexus in Erzurum seems suited for biocontrol of this weed, although they collectively cause significant damage to the plant. The same results were obtained by Cagan et al. (2000) tor Amaranthus species in Slovakia. One of the biological features of a plant native to any particular region is the pres- ence of phytophages (flea beetles in partic- ular) closely associated with the plant. The host ranges of these phytophages are nar- row (narrowly oligophagous or monopha- gous) and their distribution is often limited Table 4. Number of Chaetocnema tibialis adults on Amaranthus retroflexus at different localities of Erzurum Province during 1998-2000. 1998 1999 2000 Localities June July August Sept June July August Sept June July August Sept Arkale 20 17 45 9 45 32 27 8 17 21 25 2 Dumlu 20 8 27 6 16 31 36 9 21 29 17 5 Ilica 23 1] 20 a 18 24 29 7 15S 19 22 6 Olur 40 29 57 12 27 15 31 9 31 3g 36 2) Oltu 35 25 28 !) 14 21 27 1] 28 30 24 8 Pasinler 16 14 16 qf 22 11 32 ®) 13 17 26 i Renkaya 65 52 102 19 34 44 48 12 33 36 62 11 Tortum 30 2] 41 10 26 28 29 5 22 29 33 9 VOLUME 105, NUMBER 2 to or does not exceed the range of the host plant. Absence of such oligophages or mon- ophages on a given territory would suggest that it probably is not part of the native range of the plant. The fact that three years of exploration in Erzurum Province re- vealed no species of flea beetles specialized in feeding on Amaranthus retroflexus prob- ably suggests that Erzurum Province of Turkey is not part of the native range of Amaranthus retroflexus. ACKNOWLEDGMENTS We thank E. E. Grisell and A. L. Norr- bom (Systematic Entomology Laboratory, Washington, DC) and C. Staines (Depart- ment of Systematic Biology, Smithsonian Institution, Washington, DC) for reviewing this manuscript and providing valuable sug- gestions. LITERATURE CITED Aslan, I. 1997. Erzurum I Alticinae (Coleoptera, Chry- somelidae) Tiirleri Uzerinde Faunistik ve Siste- matik Bir Arastsrma. Doktora Tezi, Atatiirk Univ- ersitesi Fen Bilimleri Enstitiisi. 215 pp. Aslan, I. and H. Ozbek. 1998. Erzurum’da sebzelerde zarar yapan yaprak bécekleri (Coleoptera, Chry- somelidae). 2. Sebze Tarrm1 Sempozyumu, 28-30 September 1998, Tokat, pp. 335-340. 1999. Erzurum Ili’nde bazi yabanci otlarda beslenerek Gnemli derecede zarar veren yaprak bécekleri (Coleoptera, Chrysomelidae). Tiirkiye 4. Biyolojik Miicadele Kongresi Bildirileri, 26-29 January1999, Adana, pp. 75-86. 2000. Cruciferae Familyasi Igerisinde Yer Alan Bazi Sebzelerde Zararli Olan Phyllotreta (Coleoptera, Chrysomelidae), Alticinae) Turleri, Kisa Biyoloji ve Zararlari. II. Sebze Tarimi Sem- pozyumu, 11-13 September 2000, Isparta, pp. 203-207. Aslan, I. and A. Warchalowski. 2001. New records of the subfamily Alticinae (Coleoptera, Chrysomeli- dae) for the Turkish fauna. Journal of the Ento- mological Research Society 3(1—2): I—4. Aslan, [., B. Gruey, and H. Ozbek. 1999. A Prelimi- nary Review of the Subfamily Alticinae (Cole- optera, Chrysomelidae) in Turkey. Turkish Journal of Zoology 23(4): 373-414. Auld, B. A. and R. W. Medd. 1987. Weeds, an IHlus- trated Botanical Guide to the Weeds of Australia. Inkata Press. Melbourne Sidney. 255 pp. Baytop, T. 1994. Turkce Bitki Adlan S6zligu. Tiirk Dil Kurumu Yayinlari: 578. 508 pp. 445 Bermejo, J. E. H. and A. L. Gonzalez. 1994. Processes and causes of marginalization: the introduction of American flora to Spain. pp. 261-272. In Ber- mejo, J. E. H. and J. Leon, eds. Neglected Crops. 1492 from a Different Perspective. FAO Plant Production and Protection Series No 26. 341 pp. Cagan, L., M. Vrablova, and P. Toth. 2000. Flea beetles (Chrysomelidae: Alticinae) species occurring on Amaranthus spp. in Slovakia. http://www.agr.hr/ JCEA/Issues/jceal—1/fbsa-fulltext.htm Journal Central European Agriculture 1(1): 14—25. Cooke, D. A. 1992. Pests of sugar beet in the UK. Agricultural Zoology Reviews 5: 97-137. Doguet, S. Alticinae. Faune de France 80. Fédération Fran- caise des Sociétés naturelles. Paris. 697 pp. Gruey, B. and V. Tomoy, 1986. Fauna Bulgarica 13, Coleoptera, Chrysomelidae, Part I]. Chrysomeli- nae, Galerucinae, Alticinae, Hispinae, Cassidinae. In Aedibus Academia Scientiarum Bulgaricae. 388 pp. Haughton, C. S. 1978. Green Immigrants: Plants that Transformed America. Harcourt, Brace, Jovanov- ich, New York and London. 450 pp. King, L. J. 1966. Weeds of the World. Biological Con- trol. London. Leonard Hill. Interscience Publish- ers, Inc. New York. 526 pp. Konstantinov, A. S. 1988. Landscape and zonal fea- tures in the distribution of Chaetocnema species 1994. Coléoptéres Chrysomelidae, Vol. 2 (Coleoptera, Chrysomelidae) in the European part of the USSR and the Caucasus. Vestnik Zoologii 6: 44-47. Konstantinov, A. and N. Vandenberg. 1996. Handbook of Palearctic flea beetles (Coleoptera: Chrysome- lidae: Alticinae). Associated Publishers Gaines- ville. 1(3): 238-439. Kostromitin, V. B. 1978. Damage by crucifer flea bee- tle. Zashchita Rasteni 7: 3-7. Lopatin, I. 1977. Leat-beetles (Chrysomelidae) of Middle Asia and Kazakhstan. Nauka, Leningrad, 268 pp. Mohr, K. H. 1966. Chrysomelidae, pp. 95-299. In Freude, H., K. W. Harde, and G. A. Lohse (Goe- cke and Evers, eds.), Die Kafer Miteleuropas Kre- feld. Vol. 9. Mostovaya, R. N. 1994. Distribution by habitats of the chief pests in a rotation. Sakharnaya Svekla 9: 13— 14. Miller-Scharer, H. 1993. Biological control of weeds in crops: a proposal of new COST action, pp. 181-185. Jn Maitrise des adventises par voie non chimique. IFOAM Conference, Dijon. Naibo, B. 1974. Damage by the flea beetle Phyllotreta vittula Redt. on maize. Revue de Zoologie Agri- cole et de Pathologie Vegetale 73(2): 70-72. Neves, E. F 1983. On the insects fauna of sugar beet in Portugal. Boletim da Sociedade Portuguesa de Entomologia 2(37): 77—94. 446 Nikitin, V. V. 1983. Weedy plants of the flora of the USSR. Leningrad. Nauka. 451 pp. Nonveiler, G. 1960. Stetni buvaki kulturnog i drugog korisnog bilja Srbije (Halticinae, Fam. Chryso- melidae, Coleoptera). Institut za zastitu bilja, Be- agrad, pp. 37—43. . 1978. Les Altises de Serbie et Leurs Plan- teshotes (Chrysomelidae; Coleoptera). 1. Le gen- era Chaetocnema Steph. Recueil des travaux sur la faune d’insectes de la Serbie, t.2, de 1’ Academie Serbe des Sciences et des Arts, Beograd, pp. 91— DAs Onat, G. 1993. Tiirkiye’de pancar zararlisi olarak tespit edilen b6cekler. Reker Dergisi 87: 57-62. Popov, P. 1972. Insect pests of medicinal crops in Bul- garia, II] Coleoptera. Rastaniev’dini Nauki 9(5): 167-175. Rimsa, V. and I. Konecny. 1983. Sugar beet seedling pests and diseases: present and future control in Czecholovakia. 10th International Congress Brighton, England, 20-25 November 1983. Plant Protection for Human Welfare, p. 1208. Roland, A. E. and E. C. Smith. Nova Scotia. Halifax, NS. Nova Scotia Museum. 746 pp. 1969. The flora of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Schroeder, D., H. Miiller-Scharer, and C.S. A. Stinson, 1993. A European weed survey in 10 major crop systems to identify targets for biological control. Weed Research 33(6): 449-458. Slavchev, A. 1984. Protection of sugar beet against pests. Rastiteina Zaschita 32(2): 20-23. Spencer, E. R. 1957. All About Weeds. Dover Publi- cations, Inc., New York. 333 pp. Szoeke, K. 1997. Damage of millet flea beetle on spring barley and maize. Novenyvedelem 33(1): 33-34. Warchalowski, A. 1996. Ubersicht der westpalaark- tischen Arten der Gattung Longitarsus Berthold, 1827 (Coleoptera: Chrysomelidae: Halticinae). In- ternational Journal of Invertebrate Taxonomy (Supplement), Wroclaw, Poland, BS. 266 pp. Wohlgemuth, K., G. J. Schamber, A. R. Misek, and J. D. Crenshaw. 1987. Pigweed is toxic to pigs. North Dakota Agricultural Experiment Station. Bulletin 44(4): 21-22. Yildirim, E. and H. Ozbek. 1992. Erzurum Seker Fabrikasina bagh seker pancari Uretim alan- larindaki zararl ve yararl b6cek tiirleri. Tiirkiye II. Entomoloji Kongresi, 28-31 Ocak 1992, Ada- na, pp. 621-635. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 447-451 SOME NEW GENERIC NAMES IN THE CICADELLIDAE (HEMIPTERA: DELTOCEPHALINAE, SELENOCEPHALINAE) STUART H. MCKAMEY Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, “co National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0168, U.S.A. (email: smckamey @sel.barc.usda.gov) Abstract.—New replacement names are proposed for eleven genera of deltocephaline and selenocephaline leafhoppers: Mehiphanes tor Hemiphanes Emeljanov (subgenus of Euscelis), Ctenurellina tor Ctenurella Vilbaste, Mocolinna for Mocoa Linnavuori & DeLong, Saadevra for Saavedra Linnavuori & DeLong, Salsocolila for Salsolicola Ther- on, Suarezilinna for Suarezia Linnavuori & DeLong, Idyinna for Idyia Linnavuori (sub- genus of Cicadulina), Garusa tor Argusa Emeljanov (subgenus of Phlebiastes), Dwightla for Dwightia Linnavuorl & Al-Ne’amy, Bumka for Kumba Linnavuori & Al-Ne’amy, and Moluccazhangia for Moluccasia Zhang & Webb. The new generic names circumscribe 27 valid species, all of which, except three in the preoccupied subgenera, are here pro- posed in new combinations. The new tribe Dwightlini is proposed to replace Dwightiini. Key Words: Numerous generic homonyms were dis- covered while preparing a database of world leafhoppers that will be made avail- able separately. The following changes in Deltocephalinae and Selenocephalinae are meant to correct cicadellid with respect to the International Code of Zoological Nomenclature (ICZN; Interna- tional Commission on Zoological Nomen- clature 1999). Taxa are listed alphabetically within their higher taxa. It warrants mention that, if it again thought to constitute a separate ge- nus-group entity, a new replacement name would also be needed for /nsulanus Lin- navuorl, 1960 (described as a subgenus of Deltocephalus). The subgenus was placed in synonymy under Recilia Edwards, 1922, by Knight (1975a) and is preoccupied by a beetle genus described in 1930. Another preoccupied leafhopper genus- is Bubulcus Dlabola 1961b, classification is ever group name leafhoppers, Homoptera, Auchenorrhyncha, new genus, new tribe which was described as a subgenus of Paralimnus and is preoccupied by the ge- nus of the cattle egret, described in 1845. The type species of Bubulcus Dlabola is Paralimnus cingulatus Dlabola, 1960b, which was later designated as the type spe- cies of another monotypic genus, Paralim- nellus Emeljanov 1972d. Hence, as Asche and Webb (1994) noted, the valid combi- nation for the species is Paralimnellus cin- gulatus. No replacement name is needed. Note that Bubulcus Dlabola was treated as a full genus, separate from Paralimnus, in Hamilton’s (1975b) review of tribal classi- fication of Deltocephalinae. Letter designations for literature citations through 1985 are consistent with the bibli- ographies by Metcalf (1964a) and Oman et al. (1990). Subfamily Deltocephalinae Tribe Athysanini 448 Ctenurellina McKamey, new name Ctenurellina, nomen novum for Ctenurella Vilbaste 1968a: 140, preoccupied by Ctenurella Orvig 1960 (Pisces, fossil). Type species: Ctenurella paludosa Vil- baste. Ctenurellina paludosa (Vilbaste), new combination Ctenurella paludosa Vilbaste 1968a: 141 [n. sp.]. Euscelis (Mehiphanes) McKamey, new subgeneric name Mehiphanes, nomen novum for Euscelis (Hemiphanes) Emeljanov 1999b: 608, preoccupied by Hemiphanes Foerster 1868: 172 (Hymenoptera; Ichneumoni- dae). Type species: Euscelis heptneri Zachvatkin. Euscelis (Mehiphanes) heptneri (Zachvat- kin), new subgeneric placement Euscelis heptneri Zachvatkin. Euscelis (Hemiphanes) heptneri: Emeljanov 1999b: 608 [subgeneric placement]. The origin of the name Euscelis heptneri is uncertain as Emeljanov (1999b, c) did not cite its original description and attempts at finding it in other literature were unsuc- cessful. Zachvatkin described a heptneri in another athysanine taxon [Hardya (Mimo- hardya)|, but it is highly unlikely that Emeljanov was referring to that species be- cause in the same publication in which he described Hemiphanes, he elevated Mimo- hardya to full generic status, and there are no other species in that genus. Mocolinna McKamey, new name Mocolinna, nomen novum for Mocoa Lin- navuori & DeLong 1978d: 236, preoc- cupied by Mocoa Gray 1845 (Reptilia). Type species: Mocoa elegans Linnavuori & DeLong. Mocolinna elegans (Linnavuori & De- Long), new combination PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mocoa elegans Linnavuori & DeLong 1978d: 236 [n. sp.]. Saadevra McKamey, new name Saadevra, nomen novum for Saavedra Lin- navuort & DeLong 1978b: 122, preoc- cupied by Saavedra Girault 1933 (Hy- menoptera). Type species: Mesamia fas- ciata Osborn. Saadevra fasciata (Osborn), new combi- nation Mesamia fasciata Osborn 1923c: 47 [n. sp.]. Menosoma fasciata; Linnavuori 1959b: 185 [implicit n. comb. ]. Saavedra fasciata; Linnavuori & DeLong 1978b: 122 [n. comb.]. Salsocolila McKamey, new name Salsocolila, nomen novum tor Salsolicola Theron 1979a: 80, preoccupied by Sal- solicola Kuznetzov 1960 (Lepidoptera). Type species: Salsolicola plana Theron. Salsocolila calvinia (Theron), new combi- nation Salsolicola calvinia Theron 1979a: 82 [n. sp.]. Salsocolila cristata (Theron), new combi- nation Salsolicola cristata Theron 1979a: 82 [n. sp.]. Salsocolila plana (Theron), new combina- tion Salsolicola plana Theron 1979a: 80 [n. sp.]. Suarezilinna McKamey, new name Suarezilinna, nomen novum for Suarezia Linnavuort & DeLong 1978b: 121, pre- occupied by Suarezia Budde-Lund 1904 (Crustacea). Type species: Eutettix reflex- us Osborn. Suarezilinna reflexa (Osborn), new com- bination Eutettix reflexus Osborn 1923c sp.]. Menosoma reflexa; Linnavuori 1959b: 186 [n. comb.]. > 55) [int VOLUME 105, NUMBER 2 Suarezia reflexa; Linnavuori & De- Long 1978b: 121 [n. comb.]. Tribe Macrostelini Cicadulina (Idyinna) McKamey, new subgeneric name Idyinna, nomen novum for Cicadulina (dyia) Linnavuort 1960b: 58, preoccu- pied by /dyia Rafinesque 1814 (Crusta- cea). Type species: Cicadulina (Idyia) fi- jiensis Linnavuori. Cicadulina Udyinna) fyjiensis (Linnavuori), new subgeneric placement Cicadulina Udyia) fijiensis Linnayuori 1960b: 59 [n. sp.]. Tribe Paralimnini Phlebiastes (Garusa) McKamey, new subgeneric name Garusa, nomen novum for Phlebiastes (Ar- gusa) Emeljanov 1999b: 608, preoccu- pied by Argusa Kelham 1881: 530, no- men novum for Argus Temmink 1907 (Aves). Type species: Arocephalus elymi Emeljanov 1962a: 183. Phlebiastes (Garusa) elymi (Emeljanov), new subgeneric placement Arocephalus elymi Emeljanoy 1962a: 183 [n. sp.]. Phlebiastes (Argusa) elymi; Emeljanov 1999b: 608 [n. comb.]. Subfamily Selenocephalinae Dwightlini McKamey, new tribe The tribe Dwightiini was proposed by Linnavuori & Al-Ne’amy (1983a). Names of family group taxa based on junior hom- onyms are to be replaced (ICZN Art. 39), in this case by a name based on the valid name of the former type genus Dwightia. The tribe contains only Dwightla, the new replacement name for Dwightia. Dwightla McKamey, new name Dwightla, nomen novum for Dwightia Lin- navuorl & Al-Ne’amy 1983a: 24, preoc- cupied by Dwightia Wilson 1924 (Crus- 449 tacea). Type species: Dwightia acutipen- nis Linnavuori & Al-Ne’amy. Dwightla acutipennis (Linnavuori & A\l- Ne’amy), new combination Dwightia acutipennis Linnavuori & Al-Ne’amy 1983a: 26 [n. sp.]. Dwightla alecto (Linnavuori & AI- Ne’amy), new combination Dwightia alecto Linnavuori & A\l- Ne’amy 1983a: 26 [n. sp.]. Dwightla angolana Linnavuori & AI- Ne’amy), new combination Dwightia angolana (Linnavuori & Al- Ne’amy 1983a: 25 [n. sp.]. Dwightla hercules (Linnavuori & AI- Ne’amy), new combination Dwightia hercules Linnavuori & Al- Ne’amy 1983a: 26 [n. sp.]. Tribe Selenocephalini Bumka McKamey, new name Bumka, nomen novum tor Kumba Linna- vuori & Al-Ne’amy 1983a: 85, preoccu- pied by Kumba Marshall 1973 (Pisces). Type species: Selenocephalus armatissi- mus Linnavuori. Bumka_ agesilaus (Linnavuori Ne’amy), new combination Kumba_ agesilaus Linnavuori & AIl- Ne’amy 1983a: 90 [n. sp.]. & Al- Bumka_ albunea (Linnavuori & AI- Ne’amy), new combination Kumba_ albunea Linnavuori & AI- Ne’amy 1983a: 89 [n. sp.]. Bumka amphitryon (Linnavuori & A\l- Ne’amy), new combination Kumba amphitryon Linnavuori & Al- Ne’amy 1983a: 87 [n. sp.]. Bumka (Linnavuori), combination Selenocephalus armatissimus Linna- vuori 1969a: 1165 [n. sp.]. Kumba armatissima; Linnayuori & Al- Ne’amy 1983a: 86 [n. comb. ]. Bumka asera (Linnavuort & Al-Ne’amy), new combination Kumba_ asera Linnavuori Ne’amy 1983a: 90 [n. sp.]. armatissima new & Al- 450 Bumka_ bifalcata (Linnavuori & AI- Ne’amy), new combination Kumba_ bifalcata Linnavuori & AI- Ne’amy 1983a: 89 [n. sp.]. Bumka_ graphica’ (Linnavuori Ne’amy), new combination Kumba_ graphica Linnavuori & AI- Ne’amy 1983a: 89 [n. sp.]. Bumka leimonias (Linnavuori), new com- bination Selenocephalus leimonias Linnavuori 1969a: 1166 [n. sp.]. Kumba leimonias; Linnayuori & A\l- Ne’amy 1983a: 89 [n. comb.]. Bumka pylaon (Linnavuori), new combi- nation Selenocephalus pylaon Linnavuori 1969a: 1167 [n. sp.]. Kumba pylaon; Linnavuori & A\I- Ne’amy 1983a: 85 [n. comb.]. Bumka thyestes (Linnayuori), new combi- nation Selenocephalus thyestes Linnavuori 1969a: 1168 [n. sp.]. Kumba_thyestes; Linnayuori & AI- Ne’amy 1983a: 86 [n. comb.]. Bumka varia (Signoret), new combination Selenocephalus varius Signoret 1858 & Al- {in Fairmaire & Signoret 1858a: 343] [n. sp.]. Kumba varia; Linnavuori & AIl- Ne’amy 1983a: 86 [n. comb.]. Moluccazhangia McKamey, new name Moluccazhangia, nomen novum for Mol- uccasia Zhang & Webb 1996a: 8, pre- occupied by Moluccasia Rehn 1948 (Or- thoptera). Type species: Selenocephalus marmoreus Walker. Moluccazhangia marmorea (Walker), new combination Selenocephalus marmoreus Walker 1870b: 325 [n. sp.]. Moluccasia marmorea; Zhang & Webb 1996a: 8 [n. comb.]. Moluccazhangia notula (Walker), new combination PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Selenocephalus notulus Walker 1870b: 326 [n. sp.]. Moluccasia notula; Zhang & Webb 1996a: 8 [n. comb. ]. ACKNOWLEDGMENTS J. Dorsey assisted in the search for ge- neric homonyms. R. Linnavuori encouraged the replacement of the junior homonyms. M. D. Webb (The Natural History Museum, London) and FE C. Thompson (USDA/ARS Systematic Entomology Laboratory) pro- vided helpful comments on an earlier draft of the manuscript. LITERATURE CITED Asche, M. and M. D. Webb. 1994. Review of the southern Palaearctic and Palaeotropical leafhopper genus Hengchunia Vilbaste (Homoptera, Cicadel- lidae). Tijdschrift Voor Entomologie 137: 143— 154. Budde-Lund, G. 1904. A Revision of “‘Crustacea Iso- poda Terrestria” with Additions and Illustrations. 1. Eubelum. 2. Spherillionae. 3. Armadillo. H. Hagerup, Copenhagen. 144 pp. Dlabola, J. 1960b. Einige neue Zikaden aus Dagestan und Zentralasien (Homoptera). Stuttgarter Beitra- ge zur Naturkunde aus dem staatliche Museum fiir Naturkunde in Stuttgart 40: 1—5. . 1961b. Die Zikaden von Zentralasien, Dages- tan und Transkaukasien (Homopt. Auchenorrhyn- cha). Acta Entomologica Musei Nationalis Pragae 34: 241-358. Emeljanov, A. EF 1972d. New Palaearctic leafhoppers of the subfamily Deltocephalinae (Homoptera, Ci- cadellidae). Entomologicheskoe Obozrenie 51: 102—111. [In Russian.] . 1999b. [A key to the genera of the subfamily Deltocephalinae s.1. (Homoptera, Cicadellidae) from Kazakstan, Middle Asia and Mongolia with description of new genera and subgenera.] Ento- mologicheskoye Obozreniye 78(3): 589-609. [In Russian. ] . 1999c, A key to the genera of the subfamily Deltocephalinae s.l. (Homoptera, Cicadellidae) from Kazakstan, Middle Asia and Mongolia with description of new genera and subgenera. Ento- mological Review 79(5): 547-562. Fairmaire, L. and V. Signoret. 1858a. Hémipteres, Homopteres, Latreille. Deuxiéme partie, pp. 330— 343. In Thomson, J., ed. Voyage au Gabon. His- toire naturelle des insectes et des arachnides re- cueillis pendant un voyage fait au Gabon en 1856 et en 1857 par M. Henry C. Deyrolle, sous les auspices de MM. Le Comte de Mniszech et James VOLUME 105, NUMBER 2 Thomson précédée de |’histoire du voyage par M. James Thomson. Archives Entomologiques ou re- cueil contenant des illustrations d’insectes nou- veaux ou rares par M. James Thomson 2. 469 pp. Foerster, J. R. 1868. Synopsis der Familien und Gat- tungen de Ichneumonen. Verhandlungen des Na- turhistorischen Vereines der Preussischen Rhein- lande, Westfalens und des Reg. Bez. Osnabruck. Bonn. 25: 135-221. Girault, A. A. 1933. Some Beauties Inhabitant not of Commercial Boudoirs but of Nature’s Bosom, No- tably New Insects. Privately printed. 5 pp. Gray, J. E. 1845. Catalogue of the Specimens of Liz- ards in the Collection of the British Museum [Nat- ural History]. British Museum (Natural History), London. xxvili + 289 pp. Hamilton, K. G. A. 1975b. Review of the tribal clas- sification of the leafhopper subfamily Aphrodinae (Deltocephalinae of authors) of the Holarctic re- gion (Rhynchota: Homoptera: Cicadellidae). Ca- nadian Entomologist 107: 477—498. International Commission on Zoological Nomencla- ture. 1999. International Code of Zoological No- menclature. 4th ed. International Trust for Zoolog- ical Nomenclature, London. xxix + 306 pp. Kelham, H. R. 1881. Ornithological notes made in the Straits Settlements and in the Western States of the Malay Peninsula. The Ibis (4th Series) 5: 501— 532: Knight, W. J. 1975a. Deltocephalinae of New Zealand (Homoptera: Cicadellidae). New Zealand Journal of Zoology 2: 169-208. Kuznetzovy, V. I. 1960. New species of the genera Sal- solicola Kuznetz., g. nov., Pammene Hb. and Las- peyresia Hb. (Lepidoptera, Tortricidae) of the fau- na of the USSR. Entomologicheskoe Obozrenie 39: 189-199. [In Russian.] Linnavuori, R. 1959b. Revision of the Neotropical Deltocephalinae and some related subfamilies (Homoptera). Suomalaisen Elain-ja Kasvitieteeli- sen Seuran Vanamon Elaintieteellisia Julkaisuja 20: 1-370. . 1960b. Cicadellidae (Homoptera, Auchenor- rhyncha) of Fiji. Acta Entomologica Fennica 15: 1-71. 1969a. Contribution a la faune du Congo (Brazzaville). Mission A. Villiers et A. Descar- pentries XCHI. Hemipteres Hylicidae et Cicadel- lidae. Bulletin de I’ Institut Fondamental d° Afrique Noire (Series A) 31: 1129-1185. Linnavuori, R. and K.T. Al-Ne’amy. 1983a. Revision of the African Cicadellidae (subfamily Selenoce- phalinae) (Homoptera, Auchenorrhyncha). Acta Zoologica Fennica 168 105 pp. Linnavuori, R. and D.M. DeLong. 1978b. Neotropical leafhoppers of the Bahita group (Homoptera: Ci- cadellidae: Deltocephalinae). A contribution to the taxonomy. Brenesia 14—15: 109-169. 451 . 1978d. Some new or little known Neotropical Deltocephalinae (Homoptera: Cicadellidae). Bre- nesia 14&15: 227-247. Marshall, N. B. 1973. Fishes of the western north At- lantic. Genus Cetonurus Giinther 1887. Memoir, Sears Foundation for Marine Research |: 613— 623. Metcalf, Z. P. 1964a. General Catalogue of the Ho- moptera. Fascicle VI. Cicadelloidea. Bibliography of the Cicadelloidea (Homoptera: Auchenorrhyn- cha). United States Department of Agriculture, Agriculture Research Service. 349 pp. Orvig, T. 1960. New finds of acanthodians, crossop- terygians, ganoids and dipnoans in the Upper Middle Devonian calcareous flags (Oberer Plat- tenkalk) of the Begische Gladbach-Paffrath Trough. Palaontologische Zeitschrift 34: 295-335. Oman, P. W., W. J. Knight, and M. W. Nielson. 1990. Leathoppers (Cicadellidae): a Bibliography, Ge- neric Check-list and Index to the World Literature 1956-1985. CAB International Institute of Ento- mology, Wallingford, UK. [iv] + 368 pp. Osborn, H. 1923c. Neotropical Homoptera of the Car- negie Museum. Part 2. Records and descriptions of five new genera and sixty-five new species of the subfamily Jassinae. Annals of the Carnegie Museum 15: 27-79. Rafinesque, C. S. 1814. Précis des découvertes et tra- vaux somiologiques de Mr. C. S. Rafinesque- Schmaltz entre 1800 et 1814 ou Choix raisonné de ses principales découvertes en zoologie et en botanique, pour servir d’introduction a ses ouv- rages futurs. Royale typographie militaire, aux de- pens de l’auteur. Palerme. 55 pp. Rehn, J. A. G. 1948. Replacement of two preoccupied names of Tetrigidae (Orthoptera: Acridoidea). En- tomological News 59: 154-155. Theron, J. G. 1979a. Cicadellidae (Hemiptera) asso- ciated with the ganna bush, Salsola esterhuyseniae Botsch. Journal of the Entomological Society of Southern Africa 42: 77-88. Vilbaste, J. 1968a. Systematic Treatise of Cicadas Found on the Edge of the Coastal Regions. Uber die Zikadenfauna des Primorje Gebietes. Izda- tel’stvo “*Valgus,” Tallin. 195 pp. [In Russian.] Walker, EF 1870b. Catalogue of the Homopterous in- sects collected in the Indian Archipelago by Mr. A. R. Wallace, with descriptions of new species. Linnean Society of London Journal Zoology 10: 276-330. Wilson, C. B. 1924. New North American Parasite Co- pepods, New Hosts and Notes on Copepod Lit- erature. Proceedings of the United States National Museum 64 art. 17. 22 pp. Zhang Y. and M. D. Webb. 1996. A Revised Classi- fication of the Asian and Pacific Selenocephaline Leafhoppers (Homoptera: Cicadellidae). Bulletin of the Natural History Museum, Entomology Se- ries 65(1) 103 pp. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 452-459 BIOLOGY AND IMMATURE STAGES OF THE CRANE FLY PTILOGYNA (PLUSIOMYIA) HERRONI (ALEXANDER) (DIPTERA: TIPULIDAE) FROM NEW CALEDONIA, WITH DISCUSSION OF ITS PHYLOGENETIC PLACEMENT CHEN W. YOUNG AND C. DENNIS HYNES (CWY) Section of Invertebrate Zoology, Carnegie Museum of Natural History, 4400 Forbes Ave., Pittsburgh, Pennsylvania 15213, U.S.A. (e-mail: younge @carnegiemuseums. org); (CDH) 4934 Peyton Street N. Keizer, OR 97303, U.S.A. Abstract.—The last instar larva and pupa of the crane fly Prilogyna (Plusiomyia) her- roni (Alexander) (Diptera: Tipulidae: Tipulinae) are described and illustrated from spec- imens collected in New Caledonia. The microhabitat of the last larval instar and pupa is discussed. Comparisons are made with larvae and pupae of other Tipulinae. This repre- sents the first detailed description with illustrations for the larva and pupa of Ptilogyna Westwood. Pupal characters indicate a basal phylogenetic placement for this genus within the Tipulinae. Key Words: The crane fly genus Prilogyna Westwood (Tipulidae: Tipulinae) comprises 30 species and subspecies placed in three subgenera. Both Ctenogyna Macquart and Prilogyna Westwood are subgenera endemic to the Australasian and Oceanian regions (Ooster- broek and Jonas 1986). The third subgenus, Plusiomyia Skuse, apparently has a trans- Antarctic or trans-Pacific distribution with 18 species in Australia, 4 in New Caledonia (Oosterbroek 1989), and 1 in Brazil (AI- exander and Alexander 1970). Of the four Ptilogyna species recorded from New Ca- ledonia, the adult of P. herroni (Alexander) has the least wing coloration, and the male has the longest antennal flagellar branches (Alexander 1948: Fig. 7). Features of the immature stages are im- portant for interpreting phylogenetic rela- tionships within Tipuloidea (Oosterbroek and Theowald 1991). Knowledge of the im- mature stages of tipulid genera is very in- complete and this is especially true for gen- Diptera, Tipulidae, crane fly, Ptilogyna, larva, pupa, New Caledonia era in the Southern Hemisphere. Lack of information on immatures for genera in the Southern Hemisphere has hindered their placement in phylogenetic classifications. Of 30 genera and subgenera of Tipulinae recorded from the Australasian and Ocean- ian Regions (Oosterbroek 1989), we know larvae for only ten (33%), and even those primarily are known from species occurring in the Nearctic Region (Alexander 1920, Rogers 1949, Byers 1961, Gelhaus 1986). The greatest number of genera and subgen- era with unknown larval and pupal stages occur in the Australasian and Neotropical Regions. Recent detailed descriptions of immature stages of Brachyprema Osten Sacken (Gelhaus and Young 1991), Lepto- tarsus Guérin-Méneville (subgenera Limon- iodes Alexander, Pehlkea Enderlein, and Longurio Loew) (Gelhaus and Young 1995), and Tipula Linnaeus (subgenus 71- pulodina Enderlein) (Young 1999) has con- tributed information of phylogenetic signif- VOLUME 105, NUMBER 2 icance. The successful rearing of several last instar larvae to adults by C. Dennis Hynes while conducting research on the crane fly fauna of New Caledonia (Hynes 1993) allows the first complete, illustrated description of both last instar larva and pupa for the genus Prilogyna. Morphologi- cal characters of the pupa observed suggest a possible basal phylogenetic position for the genus within the Tipulinae. Terminology of larval and pupal charac- ters follows that of Byers (1961) and Gel- haus (1986). The term Tipulidae used here follows Alexander and Byers (1981) with three subfamilies (Tipulinae, Cylindrotom- inae, and Limoniinae). Catalogue of Pa- laearctic Diptera recognizes three families: Tipulidae (Oosterbroek and Theowald 1992), Cylindrotomidae (Sods and Ooster- broek 1992), and Limoniidae (Savchenko, Oosterbroek and Stary 1992) that corre- spond to the subfamilies just mentioned (Byers 1992). Stary (1992) proposed the fourth family Pediciidae, formerly a sub- family within the Limontidae. Accordingly, the term Tipulinae used here refers to all Tipulidae sensu stricto. Ptilogyna (Plusiomyia) herroni (Alexander) (Figs. 1-13) Larval description.—Mature larva: Body 35—38 mm long and 5—6 mm wide, tapering gradually toward both ends. Body sordid yellow, darker laterally. Head: Broad, massive, well sclerotized, oval-shaped and slightly depressed, typical tipuline head capsule (Byers 1961, Young 1981, Gelhaus and Young 1991); antenna with cylindrical basal segment and apical sensory peg, about three times as long as its basal width, slightly tapering apically; mandible bifurcate with distinct, large ven- tral tooth and smaller dorsal tooth, two smaller teeth subapically along inner mar- gin of ventral tooth, an additional tooth and hair tuft near base of mandible (Fig. 1); maxilla with hairy galea and lacinia; hy- popharynx with five teeth, median tooth 453 lowest and broad (Fig. 2); hypostomal bridge with one large, acute central tooth and four smaller ones and one corner tooth on each side (Fig. 3). Thorax: All segments covered with short microscopic hairs; prothoracic segment brown; meso- and metathoracic segments with well-defined patches of short, brown with silvery iridescence, microscopic hairs (Fig. 4); macrosetae short. Abdomen: All segments covered with short, brownish, microscopic hairs; macro- setae short to no visible hair on socket; se- tae D1 and D5 long, D6 slightly shorter and finer, D2—D4 represented by bare sockets (Fig. 5); setae V2 and V5 long, V1 and V4 slightly shorter, V3 represented by bare socket (Fig. 6); seta L3 long, L1 pale and shorter, L2 represented by bare socket, L4 absent (Fig. 7); abdominal cuticular out- growths absent. Spiracular disc (Fig. 8) sur- rounded by six fleshy lobes, each about as long as its basal width; lateral pair slightly longer than others; all lobes not fringed with long hair or setae; dorsal and lateral lobes unsclerotized; ventral lobes with an- terior surface sclerotized and each with a long apical seta; spiracles large, circular, dark brown with blackish centers; conspic- uous black spot below each spiracle, sur- rounded by short microscopic hairs; micro- scopic hairs also forming other dark patches and streaks at bases of all lobes; anal seg- ment large, anal opening transverse; four anal papillae, lateral pair conical, ventral pair broadly rounded (Fig. 9). Remarks.—The larva of P. herroni re- sembles superficially species in the most terrestrial subgenera of genus Tipula, spe- cifically the Nearctic subgenera Lunatipula Edwards, Pterelachisus Rondani, Trichoti- pula Alexander, and Triplicitipula Alexan- der, and the Neotropical subgenus Eumicro- tipula Alexander. They all share the follow- ing morphological characters: body heavily covered with short microscopic hairs; spi- racular lobes fleshy or partially sclerotized without border of setae; four or fewer dis- tinct papillae (Gelhaus 1986). Notable dif- 454 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON | er IL3 v5, V3 7 t ; ({ Wa * ee 6 Figs. 1-7. Ptilogyna herroni larva. 1, Mandible, dorsal. 2, Hypopharynx, dorsal. 3, Hypostomal bridge, ventral. 4, Thorax, lateral. 5-7, Abdominal macrosetal arrangements. 5, Dorsal setae. 6, Ventral setae. 7, Lateral setae. VOLUME 105, NUMBER 2 ferences of P. herroni from the above sub- genera are the brown with silvery irides- cence band around the anterior portion of the mesothoracic segment, the brown with silvery iridescence patches on the latero- ventral part of the meso- and metathoracic segments, the unsclerotized dorsal and lat- eral spiracular lobes, and abdominal macro- setae D2, D3, D4, V3, and L2 represented only by bare sockets. Pupal description.—Male: Length 26 mm, width 4.5 mm. Female: Length 25 mm, width 4.5 mm. Body brown, slightly darker on head and thorax. Head: Antenna greatly expanded at base and antennal branches readily visible in male (Figs. 1O—11); apex of antenna reach- ing about two-thirds length of mesothoracic tibia in male, reaching slightly beyond pro- thoracic tibia in female; paired short, wrin- kled ridges between bases of antennae; maxillary palpus short, slightly curved at base, then curved upward toward apex; apex of maxillary palpus not recurved, reaching prothoracic tibia (Fig. 10). Thorax: Length of respiratory horn 2 mm, minute annulations along its entire length; apex of wing nearly reaching end of abdominal segment II; apices of fore, mid- dle, and hind tarsi in oblique alignment, with fore tarsus shortest and adjacent to each other at ventral midline; in female apex of fore tarsus reaching anterior edge of segment IV, apices of middle and hind tarsi reaching middle of segment IV; in male, apex of fore tarsus reaching middle of segment IV, apices of middle and hind tarsi reaching posterior end of segment IV; middle tarsus slightly longer than hind tar- sus in both sexes. Abdomen: Segments II-VI with well- defined anterior and posterior annuli; small hooked spines present laterally and along posterior margins of all segments; three spines laterally on segment III—VII; 10—12 dorsal spines along posterior margin of seg- ments II-VII (Fig. 11); two ventral spines on segments I-IV and six ventral spines V—VII; terminal on segments segments 455 (VU and IX in male, VIII-X in female) with five pairs of elongate spines (Figs 12— 13); dorsal two pairs slightly blunt or acute- ly tipped, each with several smaller spines along their length; male genital sheaths (Fig. 12) curved dorsad with a lateral spine on each near bend beyond base; female with distinct cercal and hypovalvular sheaths, a single spine laterally near one- third length from each cercal apex (Fig. 13). Remarks.—The pupa of P. herroni is recognizable as Tipulinae by the wrinkles on the thoracic cuticle, the spines along posterior margins of the abdominal seg- ments, and a small spine near base of each male genital sheath (Gelhaus and Young 1995). The external structures of both male and female genital sheaths with lateral spine on the sheaths resemble those of Leptotar- sus (Limoniodes) sulphurellus (Alexander). The distinct branches of the antennal sheath found in Prilogyna are also observed in pu- pae of several subgenera of Ctenophora Meigen. This is possibly a convergent con- dition. The pupa of P. herroni can be dis- tinguished from other tipulines by the com- bined characters of the ridges between the bases of the antennae, the nearly straight (not recurved) maxillary palpal sheaths, and the distinct branches of male antennae. HABITAT AND BIOLOGY All specimens used in this study were collected in the Riviere Bleue Preserve dur- ing the months from September to Decem- ber of 1988. The preserve is primary low- land rain forest located about 20-30 km north of Ouénarou. The larvae were col- lected in three different localities assigned as Stations No. 5, No. 6, and No. 7 within the preserve (Hynes 1993). Station No. 5 (150 m) is located in the flood plain near the banks of the Riviere Bleue. This area is flooded many times during the rainy season, and water remains standing for long periods of time after such flooding. Station No. 6 (160 m), located near the edges of the river, is essentially the same vegetational type as Station No. 5, but the river does not peri- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORIN RATNER uss QUOTA Perm eaeene et fe 13 10 8-9, Ptilogyna herroni larva. 8, Spiracular area. 9, Anal papillae, ventral posterior view. 10-13, P. herroni pupa. 10, Habitus, male ventral aspect. 11, Habitus, male anterodorsal aspect. 12, Male terminal abdominal segments, lateral. 13, Female terminal abdominal segments, lateral. Figs. 8-13. VOLUME 105, NUMBER 2 odically flood it during the rainy season. Station No. 7 (170 m) is on a steep hillside and differs greatly in vegetation and aridity from Stations No. 5 and No. 6. The soil in the general collecting sites is typical of southern New Caledonia, the product of degradation of ultramafic rock. The soil consists of relatively small, nearly spherical particles, reddish in color and un- able to hold enough organic material or moisture for the development of the im- mature stages of most Tipulidae. Fallen trees in various stages of decay, along with numerous small branches entrapping leaves from various trees, form a mat of decaying organic material accumulated on the forest floor. These accumulations attain lengths of 2—3 m and widths of 1-2 m. Many remain wet or damp for long periods of time. Lar- vae of Prilogyna were collected mainly from fallen tree trunks, especially the inte- rior sections with large amounts of dark brown, decomposed material. This material remains damp throughout extended dry pe- riods. Occasionally the larvae were collect- ed within other decaying organic material, especially between layers of leaves on the forest floor. Larvae of Ptilogyna are considered as terrestrial species based on microhabitats in rotting logs away from aquatic situations, or in leaf debris on the forest floor. Larvae for rearing were kept within organic debris from rotten logs. Larvae pupated soon after capture, and adults started emerging in about eight days. Larvae, intact pupae of each sex, and exuviae of larvae and pupae were preserved in ethanol. DISCUSSION Hypotheses of phylogenetic relationships of genera within the Tipulidae have tradi- tionally relied exclusively on morphologi- cal characters of adults. Oosterbroek and Theowald (1991) proposed a phylogenetic scheme for the Tipuloidea (families Limon- iidae, Cylindrotomidae, and Tipulidae) based on larval and pupal characters. They provided informative characters in support 457 of phylogenetic relationships within the then proposed subfamilies of Limoniidae (Eriopterinae, Hexatominae, Pediciinae, and Limoniinae), and also presented strong support for a sister group relationship be- tween the Cylindrotomidae and Tipulidae s.s. However, the analysis of the family Ti- pulidae s.s. was excluded from their study due to the likely paraphyly of the currently recognized subfamilies (Dolichopezinae, Ctenophorinae, and Tipulinae). Stary (1992) has since elevated Pediciinae to family Pediciidae, and replaced Eriopteri- nae with Chioneinae, and Hexatominae with Limnophilinae due to the rules con- cerning priority. Meager knowledge of the immature stages of Tipulidae has also placed limitations on the larval key of Al- exander and Byers (1981) for the Nearctic fauna. The genera in this key do not appear in a phylogenetic sequence. The most re- cent larval key for Tipulinae is that of Gel- haus (1986), who presents a detailed taxo- nomic study of the genus Tipula in North America. No existing generic key is avail- able for immature stages of Tipulinae. The larva of P. herroni cannot be con- fused with other tipuline genera based on its distinct brown with silvery iridescence band and patches on the meso- and meta- thoracic segments. However, the arrange- ment of characteristic features around the spiracular disc with 1) unsclerotized dorsal and lateral spiracular lobes, 2) all lobes not fringed with long hairs, and 3) four reduced anal papillae, does bear a superficial resem- blance to similar structures in several spe- cies of the most terrestrial subgenera of 7i- pula; thus the larva of P. herroni cannot be easily distinguished from larvae of those subgenera, as indicated above in remarks following the larval description. Oosterbroek (personal communication) indicated that 11 teeth (nine teeth and two corners) on the hypostomal bridge of P. herroni is interesting because it is the high- est number (5—11) found in Tipulidae s.s. In Cylindrotomidae, the supposed sister group of the Tipulidae s.s., there are about 458 17 teeth. Many teeth may be considered plesiomorphic for the Tipulidae s.s. How- ever, larvae of Ctenophora and Dendroti- pula apparently have retained this plesiom- orphic number though they are not consid- ered primitive, but the larvae live in dead or decaying wood, as do those of P. her- ron. Alexander (1927) proposed some pre- sumed primitive generic lineages, including Ptilogyna, within Tipulinae based on the ra- dial field of the wing. Other workers have attempted to place various lineages within Tipulinae based on pupal characters such as the shape of the maxillary palpal sheath, the length of the distal section of the antennal sheath, and the position of the antennal sheath. A straight or curved maxillary pal- pal sheath, short distal section of antennal sheath, and apices of antennal and palpal sheaths closely approximated (for example, Megistocera Wiedemann, Brachypremna Osten Sacken, and Leptotarsus Guérin- Méneville) are considered plesiomorphic within the Tipulinae (Oosterbroek and Theowald 1991, Gelhaus and Young 1995). A strongly recurved apex of the maxillary palpal sheath, an extended distal section of the antennal sheath, and apices of antennal and palpal sheaths widely separated (for ex- ample, Dolichopeza Curtis, Nobilotipula Alexander, Platytipula Matsumura, Tipulo- dina Enderlein, and most other Tipulinae) are considered apomorphic (Gelhaus and Young 1995). In Prilogyna, the shape of the maxillary palpal sheath is slightly curved without a strongly recurved apex (plesiomorphic), the distal section of the antennal sheath is elon- gated (apomorphic), and the apices of an- tennal and palpal sheaths are widely sepa- rated (apomorphic). Based on this limited information we postulate that the genus Ptilogyna is a member closes to the basal assemblage (Brachypremna, Leptotarsus, Megistocera, Ozodicera Macquart, Clyto- cosmus Skuse) of the subfamily Tipulinae and suggest a possible phylogenetic place- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ment of it between the primitive and the advanced genera within Tipulinae. The discovery of the immature stages of P. herroni provides much-needed informa- tion in order to narrow the gap in our knowledge of the immature stages of Southern Hemisphere crane fly taxa. Our findings concur with several previous stud- ies by various authors that many of the lar- val characters (including spiracular disc, spiracular lobes, anal papillae, hypostomal teeth), and the size and form of the respi- ratory horns of the pupa provide taxonom- ically important information for identifica- tion and probably have a close correlation of structure to habitats. These larval char- acters probably have evolved in their spe- cific habitat to such a degree that they show a greater divergence than do some of the more conservative pupal characters such as maxillary palpal sheath, antennal sheath, and spines on genitalial sheaths. It is likely that the similarities of larval characters have evolved convergently because of hab- itat requirements, while the similarities of pupal characters have evolved by reason of common ancestry. This must be considered when using these characters in the place- ment of the genus within the subfamily Ti- pulinae. Chaetotaxy and additional pupal characters as mentioned in Gelhaus and Young (1995) may have applicability for establishing relationships but are not yet evaluated phylogenetically against a broad range of tipuline taxa. Further work is need- ed on the morphology of the immature forms of Tipulinae before such information can contribute to the understanding of the postulated relationships among these line- ages. ACKNOWLEDGMENTS This paper is based on specimens col- lected by Dennis Hynes while conducting research in New Caledonia. We thank Jean Chazeau (French Institute of Scientific Re- search and Development through Coopera- tion, Nouméa, New Caledonia) for his sup- port during Hynes’ studies in New Cale- VOLUME 105, NUMBER 2 donia. Thanks are due to Pyotr Oosterbroek, John E. Rawlins, and Robert Davidson for suggestions on an earlier draft of the man- uscript; also to three anonymous reviewers for their comments. LITERATURE CITED Alexander, C. P. 1920. The crane-flies of New York. Part II. Biology and Phylogeny. Cornell Univer- sity Agricultural Experiment Station, Memoir 38: 691-1133. . 1927. The interpretation of the radial field of the wing in the nematocerous Diptera, with spe- cial reference to the Tipulidae. Proceedings of the Linnean Society of New South Wales 52: 41-72. . 1948. The crane-flies of New Caledonia (Dip- tera, Tipulidae). Transactions of the royal Ento- mological Society of London 99: 361-393. Alexander, C. P. and M. M. Alexander. 1970. Family Tipulidae. Fascicle 4: 1-259. Jn Papavero, N., ed. Catalogue of the Diptera of the Americas South of the United States, Museu de Zoologia, Univ- ersidade Sao Paulo. Alexander, C. P. and G. W. Byers. 1981. Tipulidae, pp. 153-190. In McAlpine, J. F et al., eds. Manual of Nearctic Diptera, Vol. 1. Agriculture Canada Monograph No. 27. Byers, G. W. 1961. The crane fly genus Dolichopeza in North America. University of Kansas Science Bulletin 42: 665—924. 1992. Crane flies—Three families or one? Acta Zoologica Cracoviensia 35(1): 37—41. Gelhaus, J. K. 1986. Larvae of the crane fly genus Tipula in North America (Diptera: Tipulidae). University Kansas Science Bulletin 53: 121-182. Gelhaus, J. K. and C. W. Young. 1991. The immature instars and biology of the crane fly genus Brachy- premna Osten Sacken (Diptera: Tipulidae). Pro- ceedings of the Entomological Society of Wash- ington 93: 613-621. . 1995. Pupae of the crane fly genus Leprotar- sus (Diptera: Tipulidae) in the New World, with discussion of the monophyly of the genus. Annals of Carnegie Museum 64: 135-145. Hynes, C. D. 1993. The crane-flies of New Caledonia (Diptera Tanyderidae, Tipulidae), pp. 73-121. In 459 Matile L., J. Najt, and S. Tillier, eds. Zoologia Neocaledonica, Vol. 3. Mémoires du Muséum Na- tional d’Histoire Naturelle 157: 73-121. Paris, France. Oosterbroek, P. 1989. Tipulidae, pp. 53-116. /n Ey- enhuis, N. L., ed. Catalog of the Diptera of the Australasian and Oceanian Regions. Bishop Mu- seum Press, Honolulu, and E. J. Brill, Leiden. Oosterbroek, P. and T. Jonas. 1986. Catalogue of the Australian-Oceanian Tipulidae (Insecta, Diptera), Including a Geographical Index and References to the Species. Privately published, Amsterdam. 242 PP- Oosterbroek, P. and Br. Theowald. 1991. Phylogeny of the Tipuloidea based on characters of larvae and pupae (Diptera, Nematocera), with an index to the literature except Tipulidae. Tijdschrift voor Ento- mologie 134: 211-267. . 1992. Family Tipulidae, pp. 56-178. Jn Soos A., L. Papp, and P. Oosterbroek, eds. Catalogue of Palaearctic Diptera, Vol. |. Hungarian Natural History Museum, Budapest, Hungary. Rogers, J. S. 1949. The life history of Megistocera longipennis (Macquart) (Tipulidae, Diptera), a member of the neuston fauna. Occasional Papers of the Museum of Zoology, University of Michi- gan, no. 521, 14 pp. Savchenko, E. N., P. Oosterbroek, and J. Stary. 1992. Family Limoniidae, pp. 183-369. Jn Sos A., L. Papp, and P. Oosterbroek, eds. Catalogue of Pa- laearctic Diptera, Vol. 1. Hungarian Natural His- tory Museum, Budapest, Hungary. Stary, J. 1992. Phylogeny and classification of Tipu- lomorpha, with special emphasis on the family Li- moniidae. Acta Zoologica Cracoviensia 35(1): 11-36. Sods, A. and P. Oosterbroek. 1992. Family Cylindro- tomidae, pp. 179-182. Jn Sods A., L. Papp, and P. Oosterbroek, eds. Catalogue of Palaearctic Dip- tera, Vol. 1. Hungarian Natural History Museum, Budapest, Hungary. Young, C. W. 1981. The immature instars of Tipula (Platytipula) ultima Alexander (Tipulidae, Dip- tera). Journal of the Kansas Entomological Soci- ety 54: 409-415. . 1999. New species and immature instars of crane flies of subgenus Tipulodina Enderlein from Sulawesi (Insecta: Diptera: Tipulidae: Tipula). Annals of Carnegie Museum 68: 81—90. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 460—466 ERICAPHIS LOUISAE, N. SP. (HEMIPTERA: APHIDIDAE: APHIDINAE: MACROSIPHINI) ON LUETKEA PECTINATA (PURSH) (ROSACEAE), AND A KEY TO ERICAPHIS SPECIES KEITH S. PIKE AND PETR STARY (KSP) Entomologist, Washington State University, Irrigated Agriculture Research and Extension Center, 24106 N Bunn Rd, Prosser, WA 99350, U.S.A; (PS) Entomologist, Institute of Entomology, Academy of Sciences of the Czech Republic, Ceské Budéjovice, Czech Republic Abstract.—Ericaphis louisae Pike, n. sp. (Hemiptera: Aphididae: Aphidinae: Macro- siphini) is described and illustrated. The aphid is holocyclic monoecious, found on Luetkea pectinata (Pursh) (Rosaceae) in subalpine settings in the Cascade and Olympic Mountains of Washington, USA, and in the Coastal Mountains of northwestern British Columbia, Canada. A key to the species of Ericaphis is also provided. Key Words: Research on aphid parasitoids in north- western North America (Pike et al. 2000) led to the discovery of a new species of aphid on Luetkea pectinata (Pursh) (Rosa- ceae) in subalpine settings near timberline in the Cascade and Olympic Mountains of Washington and Coastal Mountains of northwestern British Columbia. Diagnosis, descriptions, and illustrations are provided, together with a key to Ericaphis species. MATERIAL AND METHODS The aphid was collected through Berlese extractions of Luetkea pectinata, and by hand shaking plants over a white surface. The aphid was found at five sites in the Cascades Mountains of Washington, from Mt. Baker to Mt. Rainier, at elevations ranging from about 1,600 to 1,800 m; at one site in the Olympic Mountains at 1,750 m; and at one site in northwestern British Columbia, Tormented Valley near Fraser at 940 m. Material was cleared and mounted on slides in Canada balsam; part was processed aphid, Ericaphis, Luetkea, new species using techniques published by Hille Ris Lambers (1950) with modifications by D. Voegtlin of the Illinois Natural History Sur- vey (see Pike et al. 1991), and part was cleared and mounted following techniques by Foottit and Maw (http://www.zoology. ubc.ca/~mawe/bcaphid). The aphid de- scription is attributed to the first author. Line illustrations were hand drawn from images taken with a Nikon Coolpix 990 digital camera. Scanning electron micros- copy (SEM) was used to illustrate antennal rhinaria of the male. SEM procedures were as follows: aphids were placed in Kahle’s solution overnight, then progressively de- hydrated through an ethanol series (70, 80, 90, 95, 100%), 30 minutes per dilution. Specimens were transferred to 12 ml screw- cap vials containing 3 ml undiluted HMDS (hexamethyldisilazane) and left to soak for 1 hour. HMDS was decanted and 3 ml of fresh, undiluted HMDS was added. Speci- mens were removed after | hour and al- lowed to air-dry overnight. Specimens were mounted on aluminum SEM_ stubs and VOLUME 105, NUMBER 2 461 Figs. 1-8. frontal tubercles of head. 3, Antenna. 4, Ultimate rostral segment. 5, Siphunculus. 6, Cauda. 7, Ovipara, hind tibia and tarsus. 8, Apterous male, antenna. Bar scale: Fig. | = 0.4 mm; Figs. 2-8 = 0.05 mm. gold-coated to a thickness of 15—20 nm us- ing a Technics Hummer V_ sputtercoater. These were scanned at an accelerating volt- age of ISKV on a Hitachi S-570 scanning electron microscope. Photographs were tak- en using Polaroid™ Type 55 film. The SEM work was performed at the Electron Mi- croscopy Center, Washington State Univer- sity (WSU), Pullman, Various features of Ericaphis louisae. 1-6, Apterous vivipara. 1, Whole mount. 2, Median and Ericaphis louisae Pike, new species (Figs. 1-9) Etymology.—The species was named af- ter the type locality, Louise Lake, Mount Rainier National Park, Washington, USA. Description.—Apterous vivipara (Figs. 1—6): Coloration in life, green. Adult: body length 1.22 to 1.81 mm. Body pale, ap- pendages pale or sometimes slightly darker 462 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON than body, especially tarsi and distal por- tions of tibiae. Dorsal integument faintly to moderately wrinkled. Dorsal and ventral ce- phalic sutures sometimes vaguely indicated. Setae on head short, pointed or blunt, usu- ally less than half the basal diameter of an- tennal segment III. Frontal tubercles (= an- tennal tubercles) moderately developed, di- verging, rugose, particularly inner margins: median tubercle moderately developed, with similar height and surface sculpturing as frontal tubercles. Antenna short, 0.40— 0.67 body length; 6-segmented without secondary rhinaria; primary rhinaria with conspicuous ciliate margins. Antennal seg- ment II 0.13—0.22 mm, TV 0.05—0.15 mm, V 0.07-0.15 mm, base of VI 0.08-—0.11 mm, processus terminalis 0.09—0.19 mm. Ultimate rostral segment (0.06—0.08 mm) tapering, rounded distad of preapical pri- mary setae, with 2 accessory setae. Hind tibia 0.46—0.65 mm; setae pointed, length of longest about % width of segment at wid- est point. Tarsal I chaetotaxy 3,3,3. Claws simple; empodial setae acuminate, not quite reaching to tip of claws. Abdominal seg- ments I-V apparently fused; spiracles re- niform to subcircular without opercula, ex- cept abdominal segments VI—VII spiracles with some marginal anterior overlap. Si- phunculus 0.19—0.30 mm, imbricated throughout, subcylindrical, and distal ¥, to ¥; part narrowing slightly to pre-flange con- striction. Cauda 0.10—0.18 mm, triangular, with 4—6 setae. For additional measure- ments, see Table 1. Ovipara (Fig. 7): Adult: body length 1.41—-1.79 mm. Similar to apterous vivipara, but typically slightly larger (see Table | for complete range of measurements). Integu- ment faintly to moderately wrinkled. Lat- eral ocelli absent or sometimes with a trace indicated. Hind tibia with 21 to 64 pseu- dorhinaria. Abdominal dorsum membra- nous, pale, sometimes with very light pig- mentation on marginals, less frequently on posterior terga. Marginals sometimes with small flat subcircular tubercles, usually one per marginal when present. Apterous male (Figs. 8-9): Adult: body length 1.17 to 1.35 mm. Body somewhat elongate compared with vivipara and ovi- para. Dorsal integument faintly or moder- ately wrinkled. Abdominal dorsum some- times partially pigmented, antenna and tarsi darkened, femora and tibiae partially dark- ened, siphunculi and cauda usually slightly darker than abdomen. Small median and lateral ocelli present. Antenna with second- ary rhinaria on III (15—22), [TV (7-16), and V (5-12). Primary antennal rhinaria with distinct marginal cilia; secondary antennal rhinaria without marginal cilia (Fig. 9). For additional measurements, see Table 1. Material.—Holotype: apterous vivipa- rous 2—USA, Washington, Lewis Co., Mt. Rainier National Park, Louise Lake, 15-[X- 1994, on Luetkea pectinata (Pursh) (WSU Code 94R227—4), collectors K. S. Pike and R. Miller. Paratypes (all from Luetkea pec- tinata): 6 viviparae and 2 oviparae collected with holotype; 2 ¢ and 5 oviparae collected on 29-IX-1994 at type locality; 11 viviparae collected on 1-VII-1995, 16-VIII-1995, and 8-IX-1995 at type locality; 1 d collected on 8-LX-1995 at Chinook Pass, Yakima Co., Washington; 3 oviparae collected on 17-X- 1995 at Mt Baker near ski area, Whatcom Co., Washington; 12 viviparae, 11 oviparae, and 3 ¢ collected on 11 and 27-L[X-1996 at Chinook Pass, Dewey Lake Trail, Yakima Co., Washington; | vivipara collected 9-X- 1996, Yakima Nation, Goat Butte, Yakima Co., Washington; | vivipara collected on 6- IX-2001 at Quartz Mountain, Kittitas Co., Washington; 15 oviparae, and 15 ¢ col- lected on 25-IX-2001 at Chinook Pass, Dewey Lake Trail, Yakima Co., Washing- ton; 3 vivipara collected on 22-VII-2002, Olympic National Park, Hurricane Ridge, Clallam Co., Washington; and 2 ovipara and | 6 collected on 14-VIHI-2002, Tor- mented Valley near Fraser, British Colum- bia. Holotype deposited in National Muse- um of Natural History, Smithsonian Insti- tution, aphid collection, Beltsville, MD (USNM). Paratypes, two each of the vivi- parae, oviparae, and males, deposited in VOLUME 105, NUMBER 2 USNM,; Canadian National Collection, Ot- tawa; The Natural History Museum, Lon- don; and Muséum National d’ Histoire Na- turelle, Paris. Additional paratypes depos- ited in the Washington State University aphid collection. Biology and distribution.—Ericaphis louisae is holocyclic monoecious on Luet- kea pectinata (Pursh) (Rosaceae). Based on present collections, it is known only from subalpine areas in the Cascades and Olym- pic Mountains of Washington and at one site in the Coastal Mountains of British Co- lumbia. Its distribution is likely more ex- tensive, probably covering the full range of its host; L. pectinata is found from the Alaska/Yukon region southward in the Coastal Mountains to southern British Co- lumbia, in the Cascades to northern Cali- fornia, in the Rocky Mountains to southern British Columbia, southwestern Alberta, and the Bitterroot Mountains of western Montana and eastern Idaho (Hitchcock and Cronquist 1973), and in the Olympic Mountains of Washington. The plant is found in moist or shaded areas, usually in sandy soil, mostly where snow persists until late into the season, from subalpine to well above timberline (Hitchcock and Cronquist 1973); Two species of primary parasitic wasps (Hymenoptera, Braconidae, Aphidiinae) were found attacking Ericaphis louisae: Aphidius sp. and Lysiphlebus testaceipes (Cresson); associated secondary parasitoids (Hymenoptera, Pteromalidae) included Asa- phes californicus Girault and an undeter- mined pteromalid. Other confirmed parasit- oid associations with members of the genus Ericaphis include: Aphidius nr. Marshall, Aphidius sp., Ephedrus lacertosus hortensis (Haliday), Lysaphidus sp., Praon unicum Smith and Praon sp. on E. gentneri; Aphi- dius sp. on E. scammelli; and Aphidius po- lygonaphis (Fitch), and Aphidius sp. on E. wakibae (Pike et al. 2000). With the addition of the new species, the genus Ericaphis is comprised of about 12 species in North America, 3 of which are 463 of European origin. Members of the genus collectively are associated with plants in six families: Ericaceae, Rosaceae, Liliaceae, Empetraceae, Scrophulariaceae and Capri- foliaceae [from the latter, only the alatae of E. harmstoni are known from a single col- lection (Knowlton 1943)—possibly errant specimens]. Diagnosis—The new species shares some similarities (moderately developed, divergent, rough sculptured frontal tuber- cles; wrinkled integument; and short pro- cessus terminalis) with Ericaphis latifrons (Borner 1942) a European species (Remau- diére 1952 [Myzodium lagarriguei = latif- rons], Ossiannilsson 1954 [E. empetri = la- tifrons|, Heie 1992), but differs from the latter in the shape of the siphunculus (with FE. louisae, siphunculus distally narrows slightly; with £. Jatifrons, siphunculus is more or less cylindrical, curving outward apically), cilia in antennal secondary rhi- naria (absent in FE. /ouisae), and number of ultimate rostral segment accessory setae (2 in E. louisae vs 4—6). The new species shares the peculiar shaped siphunculus with Abstrusomyzus (Jensen and Stoetzel 1999), but differs from the latter in body integu- ment (E. louisae is wrinkled throughout, with tergum unpigmented in apterous vivi- parae, while Abstrusomyzus spp. are dis- tinctly pentagonal/hexagonal reticulate, with tergum often completely pigmented), number of ultimate rostral segment acces- sory setae (2 vs. 3-9), and tarsal I chaeto- taxy (3,3,3 vs. 3,3,2). The new species shows some resemblance to Myzodium spp. (Smith and Robinson 1975) in shape and sculpturing of the frontal tubercles, but dif- fers from the latter in the shape of the cauda (triangular vs. abruptly narrowed apically) and host plant association (Luetkea vs. moss). Other genera, 1.e. Au/acorthum, Ela- tobium, Hyalomyzus, Myzus, Ovatus, Pho- rodon, and Rhodobium, which bear some slight resemblance to E. louisae, differ in the shape of the siphunculus, usually in the lack of development of the median tubercle, 464 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table |. Morphological measurements (mm) and comparisons for adult morphs of Ericaphis louisae. Apterous Vivapara Ovipara n= 35 n = 23 Mean (Range) Mean Body (length) 1.42 (1.15-1.81) 1.61 Head Head (width) 0.37 (0.32-0.43) 0.40 Antenna (length) 0.76 (0.60—0.93) 0.90 Il 0.18 (0.13-0.22) 0.21 IV O.11 (0.05—0.15) 0.13 Vv 0.11 (0.07—0.15) 0.14 Base of antennal segment VI 0.09 (0.08—0.11) 0.10 Processus terminalis 0.15 (0.09-0.19) 0.17 Setal length (antennal segment III) 0.005 (0.004—0.007) 0.006 Ultimate rostral segment (length) 0.07 (0.06—0.08) 0.07 Ultimate rostral segment (width) 0.05 (0.04—0.06) 0.05 Antenna III secondary rhinaria 0) — 0 Antenna IV secondary rhinaria 0 — 0 Antenna V secondary rhinaria 0 — 0 Antenna socket ventral margin setae 2.1 (1-4) 233 Ultimate rostral segment assessory setae 2 — 2 Thorax Hind tibia (length) 0.55 (0.46—0.65) 0.61 Hind tarsus II (length) 0.09 (0.07—0.10) 0.09 Tibial rhinaria 0 — 36.3 Abdomen Siphunculus (length) 0.24 (0.19—0.30) 0.27 Cauda (length) 0.14 (O.10—0.18) 0.16 Cauda (width) 0.12 (O.11—0.14) 0.12 Abdominal VIII dorsal setae 2:3) (2-4) 4.0 Caudal setae 5.0 (4-6) 5.6 Comparisons Antenna/body 0.54 (0.40—0.67) 0.56 Processus terminalis/base of antenna VI 1.56 (1.00—2.11) 1.69 Ultimate rostral segment (length/basal width) 1.51 (1.17-1.94) 1.60 Ultimate rostral segment/hind tarsus II 0.81 (0.67—1.00) 0.83 Ultimate rostral segment/body 0.05 (0.04—0.06) 0.05 Siphunculus/body 0.17 (O0.13—0.20) 0.17 Siphundulus/antennal segment VI 1.01 (0.81-1.32) 1.00 Cauda/body 0.10 (0.08—0.12) 0.10 Cauda/siphunculus 0.60 (0.42-0.79) 0.58 and usually in the shape of the frontal tu- bercles. With regard to the genus Ericaphis, some species currently in the genus may belong elsewhere, and some species currently in other myzine genera may belong in Erica- phis. A comprehensive study and revision of the genus is needed. The key below is provided as a tool for identification of the apterous viviparae of the species presently included in Ericaphis by Remaudiere and Remaudiere (1997). KEY TO ERICAPHIS (APTEROUS VIVIPARAE) [Ericaphis harmstoni not in key, since apterous viviparae are unknown (Knowlton 1943)] ile Abdominal dorsum with sclerotic pigmen- (TSTOYeh & hotemauskaus Shen ota eke ond aucneioen as 2 = Abdominal dorsum membranous ....... 3 VOL UME 105, NUMBER 2 Table 1. Extended. Ovipara Apterous Male n 23 n=9 Mean (Range) (1.41-1.79) 1.18 (1.09-1.34) (0.36—0.44) 0.37 (0.34—0.42) (O.80—1.00) 1.19 (1.09—1.26) (0.19—0.25) 0.27 (0.26-0.29) (O.10—0.16) 0.20 (0.18—0.21) (O.11—0.17) 0.20 (0.17—0.21) (0.09—0.13) 0.12 (0.11—0.14) (0.14—0.20) 0.24 (0.21—0.30) (O.005—0.007 ) 0.006 (0.005—0.007) (0.06—0.08) 0.07 (O.07—0.08) (O.04—0.06) 0.05 (0.04—0.06) — 18.6 (15-22) — 10.4 (7-16) — 98 (5-12) (1-5) 253 (1-5) —_ 2? _ (O.55—0.65) 0.69 (O0.66—0.73) (O.08—0. 10) 0.09 (O.08—0.10) (21-64) 0 = (0.25—0.30) 0.21 (O.18—0.23) (0.14—0.19) 0.10 (O0.08—0.11) (0.10—0.13) 0.11 (0.09—0.12) (3-5) 3:3 (24) (3-6) ae (4-7) (0.5 1—0.64) 1.02 (0.91—1.06) (1.27-2.00) 2.01 (1.67-2.50) (1.21—2.05) 1:59 (1.21—1.80) (0.70-0.91) 0.79 (0.78—0.88) (0.04—0.05) 0.06 (O.05—0.06) (0.15—0.19) OPI, (0.15-0.21) (0.83—1.17) 0.57 (0.43—0.66) (O.06—0.11) 0.08 (0.07—0.10) (0.52—0.70) 0.48 (0.40—0.54) 2(1). 3(1). Tarsal I chaetotaxy 5,5,5: on Ericaceae (Er- ica); distribution: Europe (single record from California) ericae (Borner) Tarsal I chaetotaxy 3,3,3; on Ericaceae (Kal- mia), Rosaceae (Fragaria, Rosa, and Rubus) and Scrophulariaceae (Pedicularis); distri- bution: N. Amer., England wakibae-siphunculata group Siphunculus distally narrowing (distal 4, to ) g 6 '/; narrowed) to pre-flange constriction, ° without reticulation (Fig. 5); on Rosaceae 465 a) Fig. 9 ary rhinaria on antennal segment II (SEM photo). Bar = 0.005 mm. Ericaphis louisae, apterous male. Second- scale (Luetkea pectinata). Distribution: Washing- ton and British Columbia = Siphunculus more or less cylindrical, some- louisae Pike tumes curved outward distally, with or with- out reticulation; not on Luetkea pectinata .. 4 4(3). Processus terminalis/base of antennal seg- 4.5 Processus terminalis/base of antennal seg- ment VI > 4.5 Processus terminalis/base of antennal seg- a ment VI < 5(4) ment VI = 2.2; antenna not reaching to base of siphunculus; on Empetraceae (Empetrum) and Ericaceae (Calluna, Vaccinium). Distri- bution: Europe, Iceland dad fetta h Dyetrads abans faeces ds Syed latifrons (Borner) = Processus terminalis base of antennal seg- ment VI > yond base of siphunculus; on Ericaceae (An- Vac- Rosa). Distribution: N. Amer, Europe ........ 6 Ultimate -1.4; on Ericaceae, Arctostaphylos, distri- 2.4; antenna reaching to or be- dromeda, Arctostaphylos, Pernettya, cinium), and Rosaceae (Fragaria, 6(5). rostral segment/hind tarsus II bution: France, French Alps (Remaudiere 1971) - Ultimate leclanti (Remaudieére ) rostral segment/hind tarsus II 1.0; reported from various Ericaceae (An- Vac- Rosa). dromeda, Arc tostaphylos, Pernettya, cinium), and Rosaceae (Fragaria, 466 Distribution: N. Amer., Europe GOhe ei Sic 6 Sole Oro fimbriata-scammelli group 7(4). Ultimate rostral segment with 2 secondary or accessory setae; on Rosaceae (Amelan- chier, Crataegus, Photinia, Mespilus, Pyrus, Sorbus). Distribution: N. Amer. Pg ce Mensa Cotta renee eiroknorcte cs gentnert (Mason) - Ultimate rostral segment with 6—8 second- ary or accessory setae; on Liliaceae (Lilium, Scoliopus, Smilacina, Streptopus). Distribu- tion: western N. Amer. lilii-scoliopi group Note that although E. harmstoni was not keyed, inter-morph comparisons with E. louisae indicate clear distinctions between the species, with differences in such char- acters as siphuncular distal shape and sculp- turing, processus terminalis/base of anten- nal segment VI (4.6—4.7 in E. harmstoni vs. 1.0—2.5 in E. louisae) and overall size (2.3— 2.5 mm in E. harmstoni vs. 1.2—1.8 mm in E. louisae) (see Knowlton 1943 for original description of E. harmstoni). ACKNOWLEDGMENTS We express thanks to the D. Allison, L. Boydston, G. Graf, and D. Graf (Washing- ton State University Technical Support Staff), R. Miller (University of Guam, for- merly with Washington State University), and L. Pike for assistance in collecting, par- asitoid rearings, mounting, and/or illustra- tions; R. F Foottit and E. Maw (Agriculture and Agri-Food Canada, Ottawa), G. Re- maudiere (Muséum National d’ Histoire Na- turelle, Paris), and M. Stoetzel (Systematic Entomology Laboratory, USDA, Beltsville, MD), A. Jensen (formerly with the System- atic Entomology Laboratory, Beltsville, MD), Cho-Kai Chan (Vancouver, BC), and P. Jepson (Department of Entomology, Oregon State University, Corvallis) for loan or access to Ericaphis spp.; and R. F. Foot- tit, E. Maw, and A. Jensen for manuscript reviews. We thank G. Ahlstrand and R. Lechleitner for facilitating collections at Mt. Rainier National Park; B. Baccus and G. Hunter for facilitating collections at the Olympic National Park; and the Yakama Nation for allowing access to their lands. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON We also thank J. Rumph (Washington State University, graduate student) for scanning electron micrographs. LITERATURE CITED Borner, C. 1942. Weitere neue europaische Blattlau- sarten. Ver6ffentlichungen der Deutschen Kolon- ie—Ubersee-Museum von Bremen 3: 259-266. Heie, O. E. 1992. The Aphidoidea (Hemiptera) of Fen- noscandia and Denmark. IV. E. J. Brill/Scandina- vian Science Press Ltd., Kobenhavn. 189 pp. Hille Ris Lambers, D. 1950. On mounting aphids and other soft-skinned insects. Entomologische Beri- chten No. 298(del XII): 55-58. Hitchcock, C. L., and A. Cronquist. 1973. Flora of the Pacific Northwest. University of Washington Press, Seattle. 730 pp. Jensen, A. S., and M. B. Stoetzel. 1999. An exami- nation of the North American aphid species cur- rently placed in Ovatus van der Goot (Hemiptera: Aphididae) with the description of a new genus. Proceedings of the Entomological Society of Washington. 101: 39-56. Knowlton, G. EK 1943. Three new western aphids. En- tomological News 54: 122-124. Ossiannilsson, F 1954. Ericaphis empetri, n. sp., anew Swedish aphid (Hem. Hom.). Opuscula Entomo- logica 19: 18-20. Pike, K. S., L. Boydston, and D. Allison. 1991. Alate aphid viviparae associated with small grains in North America: a key and morphometric charac- terization. Journal of the Kansas Entomological Society 63: 559-602. Pike, K. S., P. Stary, T. Miller, G. Graf, D. Allison, L. Boydston, and R. Miller. 2000. Aphid parasitoids (Hymenoptera, Braconidae, Aphidiinae) of North- west USA. Proceedings of the Entomological So- ciety of Washington 102: 688-740. Remaudiére, G. 1952. Contribution a |’étude des Aphi- doidea de la faune frangaise, Description de qu- elques Aphididae nouveaux et addition a la liste des Myxinae et Dactynotinae. Revue de Patholo- gie Végétale et d’Entomologie Agricole de France 31(4): 232-263. . 1971. Un Ericaphis nouveau vivant sur Arc- tostaphylos dans les Alpes frangaises (Hom. Aphi- didae). Annales de la Societé Entomologique de France, N.S., 7(4): 771-778. Remaudiére, G. and M. Remaudieére. 1997. Catalogue des Aphididae du Monde, Homoptera Aphidoidea. Institut National de la Recherche Agronomique (INRA), Paris. 473 pp. Smith, C. F and A. G. Robinson. 1975. The genus Myzodium with the description of M. knowltoni, new species (Homoptera: Aphididae). Proceedings of the Entomological Society of Washington 77: 481-480. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 467-483 STEPHANIDAE (HYMENOPTERA) OF AMERICA NORTH OF MEXICO ALEXANDRE PIRES AGUIAR AND NORMAN E JOHNSON Universidade de Sao Paulo, Museu de Zoologia, Avenida Nazaré 481, Ipiranga, Sao Paulo, SP 04263-000, Brazil (e-mail: apaguiar@usp.br); (NFJ) Department of Entomol- ogy, The Ohio State University, 1315 Kinnear Road, Columbus, OH 43212-1192, U.S.A. (e-mail: johnson.2 @osu.edu) Abstract.—Eight species of the family Stephanidae are found in Canada and the con- tinental United States. Two new species are described, Megischus annectens Aguiar, n.sp., and Megischus brevicaudatus Aguiar, n.sp., from the western U.S.A. Megischus bicolor Westwood is redescribed, and regional color forms are not recognized as formal subspecies. A key to species is provided. Data on geographical distribution, flight period, and biological associations are summarized. Key Words: species, Nearctic The family Stephanidae is a basal group of apocritan Hymenoptera that are idiobiont parasitoids of the larvae of wood-boring Coleoptera (Cerambycidae, Buprestidae) and Hymenoptera (Siricidae). The family 1s fairly small, encompassing only 290 de- scribed species around the world. Our ob- jective here is to discuss the taxonomy of stephanids from the United States and Can- ada, an area last treated by Townes (1949). We include a review and update of the spe- cies covered by Townes as well as the de- scription of two new species. The first Nearctic species of stephanid to be described was Megischus bicolor West- wood (1841). Cresson (1872, 1880) later described two additional species, one of which, Stephanus cinctipes Cresson, was designated as the type species of Schletter- erius Ashmead (Ashmead 1900a). Three species described by Schletterer (1889) and Davis (1897) are synonyms of M. bicolor. Townes (1949) described two species and recorded the Cuban species Megischus brunneus Cresson for the first time in the Hymenoptera, Stephanidae, Schlettererius Ashmead, Megischus Brullé, new U.S.A. In total, Townes recognized six Ne- arctic species and divided M. bicolor into two subspecies. Carlson (1979) did not rec- ognize of the validity of these subspecies. MATERIAL AND METHODS A total of 833 specimens were studied from North American and European mu- seums. The Mexican fauna of Stephanidae is both more diverse in species, and the spe- cies are more variable in structure and col- or. Therefore we have deferred a consider- ation of the specimens found in Mexico un- til they can be considered in the context of the full diversity of that fauna. Morpholog- ical terminology and generic concepts gen- erally follow Aguiar (1998). Measurements of total length are made from the base of the anterior tubercle on the head to the apex of the abdomen. The ratio of pronotal length to width compares the maximum length to minimum width (Fig. 1). The terms semiannular and preannular refer to differentiated parts of the pronotum (Elliott 1922). The semiannular (Fig. 3, sa) is the 468 large, inflated posterior part of the prono- tum. The preannular (Fig. 3, pa) is the short transverse area between the pronotal fold and the semiannular. The scientific and common names of plants follow the Plants Database of the U.S. Department of Agri- culture (plants.usda.gov). The graphs of phenology display the number of specimens for each week of the year as well as the number of collecting events from which specimens are derived. A collecting event is defined as the unique combination of time, place, collector, and method of col- lection. Specimens were kindly loaned by the fol- lowing institutions and individuals alpha- betized by codens used in the text: ABSC, Archbold Biological Station Collection, Lake Placid, FL; AEIC, American Ento- mological Institute, Gainesville, FL: AMNH, American Museum of Natural His- tory, New York, NY; ANSP, Academy of Natural Sciences, Philadelphia, PA; CDAE, California State Collection of Arthropods, Sacramento, CA; CHAH, Henry A. Hes- penheide collection, Los Angeles, CA; CIDA, Alberston College of Idaho, Cald- well, ID; CMNH, Carnegie Museum of Natural History, Pittsburgh, PA; CNCI, Ca- nadian National Collection of Insects, Ot- tawa, Ontario; CSUC, Colorado State Uni- versity, Ft. Collins, CO; EDNC, North Car- olina Department of Agriculture, Raleigh, NC; EMEC, University of California, Berkeley; EMUS, Utah State University, Logan, UT; FCDA, Fresno County Depart- ment of Agriculture, Fresno, CA; FMNH, Field Museum of Natural History, Chicago, IL; FSCA, Florida State Collection of Ar- thropods, Gainesville, FL; IRCW, Univer- sity of Wisconsin, Madison, WI; ISUI, lowa State University, Ames, IA; LACM, Natu- ral History Museum, Los Angeles, CA; LSUC, Louisiana State University, Baton Rouge, LA; MCPM, Milwaukee Public Museum, Milwaukee, WI; MEMU, Missis- sippi State University, Mississippi State, MS; MTEC, Montana State University, Bozeman, MT; NCSU, North Carolina State PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON University, Raleigh, NC; NHMW, Naturhis- torisches Museum, Vienna, Austria; ODAC, Oregon Department of Agriculture, Salem, OR; OSEC, Oklahoma State Uni- versity, Stillwater, OK; OSUC, Ohio State University, Columbus, OH; PMNH, Pea- body Museum of Natural History, New Ha- ven, CT; PPCD, West Virginia Department of Agriculture, Charlestown, WV; PSUC, Pennsylvania State University, University Park, PA; ROME, Royal Ontario Museum, Toronto, Ontario; RUIC, Rutgers State Uni- versity, New Brunswick, NJ; RWSC, Rob- ert W. Surdick collection; SEMC, Univer- sity of Kansas, Lawrence, KS; SFVS, San Fernando Valley State University, Northrid- ge, CA; TAMU, Texas A&M University, College Station, TX; UADE, University of Arkansas, Fayetteville, AR; UAIC, Univer- sity of Arizona, Tucson, AZ; UCDC, Uni- versity of California, Davis; UCFC, Uni- versity of Central Florida, Orlando, FL; UCMC, University of Colorado, Boulder, CO; UCMS, University of Connecticut, Storrs, CT; UCRC, University of California, Riverside; UGCA, University of Georgia, Athens, GA; ULKY, University of Louis- ville, Louisville, KY; UMMZ, University of Michigan, Ann Arbor, MI; UMRM, Uni- versity of Missouri, Columbia, MO; UNSM, University of Nebraska State Mu- seum, Lincoln, NE; USNM, National Mu- seum of Natural History, Smithsonian In- stituuion, Washington, DC; WFBM, Univer- sity of Idaho, Moscow, ID; ZSMC, Zoolo- gische Staatssammlung, Munich, Germany. KEY TO NEARCTIC SPECIES OF STEPHANIDAE is Tergite and sternite of first metasomatic seg- ment distinctly separated, both relatively short (see Fig. | in Townes 1949); pronotum short, in lateral view with distinct 90° break in profile immediately in front of mesono- tum to form a flat, polished wall; fore wing with vein 1M arched; hind tibia white ba- Sally! sic.ssa%8 Schlettererius cinctipes (Cresson) = Tergite and sternite of petiole fused, petiole distinctly longer than wide (see Fig. 2 in Townes 1949); pronotum long, evenly sub- conical; fore wing vein 1M straight; hind VOLUME 105, NUMBER 2 Figs. 1-5. 3, Same, left side, pa = bars: 0.25 mm. Drawings by Glaucia Marconato. ZEyY tibia black or brown basally (Megischus Brulle) Petiole and pronotum polished; petiole with conspicuous transverse strigavion Petiole and pronotum microreticulate; peti- ole otherwise smooth, without transverse Sinigationl.. Favs.) Sirsa. eur eke, seeten an Hind femur matte laterally; pronotum in dorsal view with deep polished constriction, interrupting gradual convergence of sides i) 4(3). anteriorly; semiannular weakly swollen: hind tibia moderately constricted medially (BigsO) ie ace oes ae at ae Hind femur polished laterally; pronotum ap- pearing conical in dorsal view, without an- terior constriction; semiannular distinctly swollen; hind tibia with distinct nearly semi- circular constriction medially (Fig. 11) Ovipositor sheath with distinct preapical white band; gena usually smooth, some- 469 Pronotum. 1, Dorsal measurements: Pnl = length; Pnw = width. 2, Megischus annectens, dorsal. preannular, sa = semiannular. 4, M. brevicaudatus, dorsal. 5, Same, left side. Scale 6 470 5(4). 6(3). 7(2). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON times with sparse, shallow foveolae; femoral sulcus distinct on pronotum; petiole rela- tively long, length 0.62—0.77 x length of metasoma beyond petiole Ovipositor sheath uniformly dark; gena usu- ally foveolate, in small specimens foveolae may be sparse, but deep, distinct; femoral sulcus on pronotum absent or inconspicu- ous; petiole short, length 0.49—0.62 length of metasoma beyond petiole (Cali- fornia) .... Megischus annectens Aguiar, n.sp. Ovipositor longer than body, length 1.08— 1.28 * body length: hind basitarsus brown- ish-yellow; median femoral tooth inclined towards apex, triangular (Fig. 8); inter- and postfoveolar areas of mesopleuron (see Aguiar 1998) distinctly transversely strigate; length of dilated part of hind tibia 1.5 length of compressed basal portion: wings weakly infuscate; dorsal portion of meta- pleuron smooth only in narrow central area see seete Coe cut Megischus bicolor Westwood Ovipositor shorter than body, length 0.80— 0.99 body length; hind basitarsus reddish; median femoral tooth long perpendicular, round (Fig. 9); inter- and postfoveolar areas of mesopleuron usually smooth or weakly transversely strigate, only rarely distinctly strigate; length of dilated part of hind tibia 2-3 * length of compressed basal portion; wings clear; dorsal portion of metapleuron entirely smooth, polished ........... ethane 3). Megischus brevicaudatus Aguiar, n.sp. Ovipositor sheath with preapical white band; pronotum anteriorly with pair of short carinae on each side (Townes 1949: Plate 25, Fig. 6); mesopleuron (Fig. 6) and pros- ternum with scattered foveolae, generally separated by distance greater than their di- ametereecrers t - Megischus arizonicus Townes Ovipositor sheath uniformly dark; pronotum with four sharp transverse carinae anteriorly (Townes 1949: Plate 25, Fig. 7); mesopleu- ron (Fig. 7) and prosternum with dense fo- veolae, each separated by less than their di- ameter (Safe. = Megischus brunneus Cresson Posterior half of pronotum with fine trans- verse strigation; mesopleuron rugulose; me- tapleuron nearly or entirely glabrous; max- illary palp short, reaching approximately to apex of mandible; vein 2-1A darkly pig- mented, tubular or nebulous; body dark brownsins COlObr gate wars yesl aay RE On ron Megischus californicus Townes Posterior half of pronotum microreticulate, without transverse strigation; mesopleuron with large shallow foveolae; metapleuron with dense tuft of setae; length of maxillary palp subequal to height of head; vein 2-1A very weakly pigmented, nebulous; body light brownianiColor 2.2 cert) aie. = es ceees PicAcoac Shree PMeme kere Megischus texanus Townes Schlettererius cinctipes (Cresson) (igs 12517) Stephanus cinctipes Cresson 1880: XVIII (original description); Cresson 1887: 52 (keyed); Schletterer 1889: 156 (repeat of Cresson (1880), distribution, keyed); Ashmead 1900b: 597 (listed). Schlettererius cinctipes: Ashmead 1900a: 150 (generic transfer, keyed); Dalla Torre 1902: 5 (listed); Enderlein 1905: 474 (listed); Enderlein 1906: 290 (listed, keyed); Viereck 1914: 132 (type species); Elliott 1922: 714 (description, taxonomic discussion, keyed); Leonard 1928: 961 (listed); Townes 1949: 362, fig. 1 (de- scription, distribution, ecology, keyed); Kirk, 1975: 59, 60 (hosts, seasonality): Meyer et al. 1978: 326 (biology, behav- ior); Carlson 1979: 741 (listed, distribu- tion, biology, ecology); Mason 1990: 94, 95 (wing venation nomenclature, figure); Mad! 1991: 120 (redescription, figure, keyed); Smith 1997: 377, 378 (distribu- tion, rearing records, keyed). Distribution.—We have seen 227 speci- mens from southern British Columbia, Washington, Idaho, Montana, South Dako- ta, Oregon, Utah, California, and southeast- ern Arizona, Fig. 12 (AEIC, CNCI, FSCA, IRCW, AMNH, UCRC, ANSP, EMUS, ROME, WFBM, EMEC, CIDA, UCDC, LACM, MTEC, ODAC, FCDA, CDAE, USNM). A single specimen in the USNM was collected in northern Virginia by D. R. Smith (Smith 1997). This specimen was from a Malaise trap in a suburban yard and may have emerged from wood from the western U.S. or Canada. Most likely it is not yet an indication of an expansion of its range to the East. It was also introduced into Australia (Victoria, New South Wales, Tasmania). Biological associations.—Kirk (1975) reared this species from ponderosa pine (P1- VOLUME 105, NUMBER 2 471 Figs. 6-11. bicolor. 9, M. brevicaudatus. nus ponderosa P. & C. Lawson), Engel- mann spruce (Picea engelmannii Parry ex Engelm.), and white fir (Abies concolor Lindl. ex Hildebr.) infested by species of Sirex Linnaeus, Urocerus Geoffroy, and Xeris Costa (Hymenoptera: Siricidae); and it has also been reared from Douglas-fir, Pseudotsuga menziesii (Mirbel) Franco. According to Townes (1949), S. cinctipes has a range similar to that of Douglas-fir and seems to be most common in forests of this tree. Taylor (1967) established a laboratory culture of Schlettererius cinctipes on Sirex noctilio Fabricius from adults collected in California. Additional material was later imported from Arizona and New Mexico, and the parasitoid was established in Tas- mania (Taylor 1976). However, Sirex noc- tilio does not occur within the natural range of S. cinctipes (Carlson 1979). Meyer et al. (1978) suggested that Monochamus orego- Mesopleuron, left. 6, Megischus arizonicus. 7, M. brunneus. 8—9, Hind left femur, lateral. 8, M. 10-11, Hind left tibia, lateral. 10, M. bicolor. 11, M. arizonicus (detail shows posterior view). Scale bars: 0.5 mm. Drawings by Glaucia Marconato. nensis (LeConte) (Coleoptera: Cerambyci- dae) may also serve as a host. Collection dates for populations in the U.S.A. and Canada extend from June | to August 29 (Townes 1949, see Fig. 17). Label data on specimens additionally re- cord specimens collected on or near grand fir, Abies grandis (Douglas ex D. Don) Lindl; specimens emerging from Jeffrey pine, Pinus jeffreyi Grey. et Balf., and sugar pine, P. lambertiana Douglas; as well as a possibly erroneous record of a rearing from thistle, Cirsium sp. (Asteraceae). Megischus annectens Aguiar, new species (Figs. 2—3, 13, 17) Etymology.—From the Latin to link, to join; in reference to the intermediate taxo- nomic status of this species. Description of @.—Reddish black, in- cluding ovipositor sheaths. Gena ventrally 472 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2 ees oT | ea oS Pepe Ay fr ¢ G cig, Oe | 5 | mw Collecting Events o 44 2 | GT Ze | oe en Li er es 1A i AOS: 161922) 25:028 31°34" 37° 4043, 46. 4952 Week of year Megischus bicolor 40 ; 35 4 30 + 25 | 20 | 15 | 10 | (0) o_o Ee oe ee 1,4 7% 10 13716, 19) 22.°25) 28 31,34 37 40+43)-46..49" 52 O Specimens g Collecting Events Number of specimens events Week of year Fig. 17. Seasonal flight periods as indicated by total number of specimens collected and number of inde- 2 g ) I pendent collecting events. Collecting events are unique combinations of localities, dates, collectors, and methods. Only specimens or events with the date recorded as a single day are illustrated. 478 straight femoral sulcus, and the presence of fine alutaceous sculpture or lines along the apical margin of the scutellum. Megischus bicolor may be distinguished from M. an- nectens by the polished smooth or sparsely punctate gena, the alutaceous microsculp- ture between the propodeal foveolae, and the white subapical band on the ovipositor sheaths. Males are very similar, in some cases indistinguishable, from males of M. brevicaudatus. They differ from males of M. annectens by the distinctly foveolate propodeum, with the foveolae widely sep- arated from one another. Very small speci- mens have the propodeal sculpture similar to M. annectens. Biological associations.—Collected on scrub hickory, Carya floridana Sarg.; pe- can, Carya illinoinensis (Wangenh.) K. Koch; unspecified Carya Nutt. sp. (Juglan- daceae); juniper, Juniperus L. (Cupressa- ceae); pond pine, Pinus serotina Michx. (Pinaceae); unspecified pine, Pinus sp.; swamp white oak, Quercus bicolor Willd.; sand live oak, Quercus geminata Small; unspecified oak, Quercus (Fagaceae). Emerged from papaya, Carica papaya L. (Caricaceae); shagbark hickory, Carya ova- ta (P. Mill.) K. Koch; unspecified Carya sp.; beech, Fagus L. (Fagaceae); swamp white oak; bur oak, Quercus macrocarpa Michx. (Fagaceae). Recorded by Townes (1949) in association with bigtooth aspen, Populus grandidentata Michx. (Salicaceae) and American beech, Fagus grandifolia Ehrh. (Fagaceae). Comments.—Townes (1949) recognized two subspecies: M. bicolor bicolor West- wood from the southeastern U.S.A. and M. bicolor sickmanni (Schletterer) throughout the more northern portion of the distribu- tion (Fig. 15). We have found no structural characters that can be used to substantiate their formal nomenclatural differentiation. The two color forms overlap but little in their geographic distribution, but significant gaps in our knowledge remain. For exam- ple, we have not seen specimens from Ten- nessee, Kentucky, Mississippi, and most of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Illinois, Indiana, West Virginia, Virginia, South Carolina, and Alabama. We see no practical reason to continue to recognize subspecies solely on the basis of color with such relatively poor sampling of the spe- cies. Therefore, we concur with Carlson’s (1979) treatment of this as a single undif- ferentiated species. Megischus brevicaudatus Aguiar, n. sp. (Figs. 4, 55,95 113, 1/8) Etymology.—From the Latin brevis, short; cauda, tail; in reference to the rela- tively short ovipositor. Description of °.—Dark brown or black, with yellowish spot ventrally on gena under eye. Reddish on pronotal lobe, trochanters, fore- and midtibiae, compressed portion of hind tibia, hind tarsus, apex of petiole; sometimes also reddish near petiolar fora- men, base of petiole. Lighter reddish to yel- lowish on basal half of flagellum; fore- and midfemora basally; fore- and midtarsi; sometimes hind trochantellus. Wing veins dark brown, membrane clear. Gena moderately foveolate on ventral half, polished smooth dorsally. Vertex lat- erally areolate rugose, posteriorly trans- versely strigate, sculpture not or barely reaching temple. Pronotum (Figs. 4—5) dis- tinctly rugose, without median longitudinal sulcus, with deep, anterior, smoothly pol- ished constriction. Semiannular weakly dis- tinguished from preannular; femoral sulcus distinct, narrow, polished smooth, sinuous or with at least posterior apex curved (Fig. 5). Apex of scutelium normally polished smooth. Mesopleuron distinctly foveolate, sometimes partially rugose foveolate. Me- tapleuron distinctly areolate rugose central- ly, densely pilose posteriorly, polished smooth dorsally from posterior margin to pleuropropodeal fovea. Interfoveolar area polished smooth to moderately transversely strigate; postfoveolar area polished smooth or with very weak strigation; very rarely both areas distinctly transversely strigate. Propodeum deeply foveolate, area between foveolae distinctly, finely alutaceous; VOLUME 105, NUMBER 2 479 Megischus brevicaudatus 18 10 5 o = 8 3 | mi os 6 0 Specimens a) Pal @ Collecting Events 2 | E 24 h = | 2 (| a io oon ono _ 1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 Week of year Megischus californicus 15 Specimens m Collecting Events Number of specimens events i oo (om 7: 1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 Week of year Megischus texanus Specimens w Collecting Events Number of specimens events (0) fo a |? en fo 1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 Week of year Fig. 18. Seasonal flight periods as indicated by total number of specimens collected and number of inde- pendent collecting events. Collecting events are unique combinations of localities, dates, collectors, and methods Only specimens or events with the date recorded as a single day are illustrated. 480 coarsely transversely rugose near petiolar foramen. Posterior spiracular plate densely foveolate, foveolae contiguous or fused to- gether. Hind femur weakly microreticulate, nearly smooth basally, progressively more distinct toward apex, sculpture much less distinct than that on dilated part of tibia. Basal tooth nearly perpendicular to long axis of femur, elongate in lateral view, rare- ly triangular, rounded in cross-section, apex blunt; apical tooth long, acute; margin of femur concave beyond apical tooth (Fig. 9). Metasomatic T2—TS8 very finely microre- ticulate, sculpture progressively more dis- tinctly developed apically. Pygidial sulcus distinctly V-shaped, without basal exten- sion; T8 sometimes depressed at base of “Vv formed by sulcus. Total length 17.6—22.8 mm; ovipositor length/total length O0.80—0.99; ovipositor length/length of metasoma beyond petiole 2.09—2.66; ovipositor length/petiole length 3.26—3.96 ; ovipositor length/length meso- soma 2.89—3.68; length of petiole/length of metasoma beyond petiole 0.62—0.73; length/minimum width of pronotum 2.17— 3.15 (N = 11 for all measurements). Description of d.—Generally similar to female, but diagnostic characters less con- spicuous because of significantly smaller body size. Diagnosis.—Megischus brevicaudatus may be separated from the closely related M. bicolor by the distinctly shorter ovipos- itor, reddish hind tarsi, V-shaped pygidial sulcus, perpendicular basal tooth, and pres- ence of a distinct concavity on the femur beyond its apical tooth in the former spe- cies. The reddish color, especially on the hind tarsi, is the only stable diagnostic fea- ture found for male M. brevicaudatus. This new species may be distinguished from M. annectens by the lack of a median longi- tudinal sulcus dorsally on the pronotum, the V-shaped pygidial sulcus, and the presence of a white preapical band on the ovipositor sheaths. Some species in Central and South America also have the deep constriction in the pronotum. Megischus brevicaudatus PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON may be distinguished from these by the dis- tinctly sculptured semiannular and hind coxa, the presence of punctures or foveolae on the ventral half of the gena, and the densely foveolate sculpture of the meso- pleuron. Biological associations.—Collected on ash, Fraxinus L. (Oleaceae); sweetpotato cactus, Cereus P.Mill. (Cactaceae); mes- quite, Prosopis juliflora (Sw.) DC. (Faba- ceae); and blue paloverde, Parkinsonia flor- ida (Benth. ex Gray) S. Wats. (Fabaceae). Material examined.—Holotype 2°: ARI- ZONA: Chiricahua Mts., 2-VI-1935, J. N. Knull; OSUC 0020309 (OSUC). Paratypes. ARIZONA: Pima Co., Tucson; 10-X-1959; F Werner; Megischus bicolor sickmanni &, det. E Werner, 1960; OSUC 0020156; 2 (UAIC). Tucson, vic. Ina/Oracle; 7-X-1985; on Cereus bloom; 2200 h; L. Nutting; OSUC 0020154; ¢ (UAIC). IBP Research Area, Silver Bell, 10 mi W Marana; 1-IX- 1972: D. S. Chandler: Megischus bicolor sickmanni det. D. S. Chandler; OSUC 0020123; 2 (UCDC). Maricopa Co., Rain- bow Valley Rd., 2—5 mi S Gila R.; 3-V- 1990; ex flowers of Cercidium floridum Benth.; Megischus arizonicus Townes, det. J. R. Wiley, 1995; OSUC 0020211; 9% (FSCA). TEXAS: Bexar Co., Fort Sam Houston; 15-IX-1953; B. J. Adelson; OSUC 0020157; 2 (UCMS). San Antonio; 6-X-1968; W. H. Tyson; OSUC 0020119; 2 (LACM). Dickens Co., White River Re- serve, Fert. Bait, 7-11-VII-1988, R. Morris; OSUC 0020363; 2 (UGCA). Hidalgo Co., Bentsen Rio Grande Valley S. P. near Mis- sion; l1O-VII-1981; €. Porter; OSUC 0020180; 2 (FSCA). Karnes Co.; 12-ILI- 1934; C. E. Heard: 5443; Megischus bicol- or sickmani det. H. M. Greenbaum 1974; OSUC 0020153; 6 (TAMU). Real Co., Rio Frio, Hwy. 90; 14-VIII-1959; OSUC 0020314; 2 (CNCI). Starr Co., Salineno, along Rio Grande; spring 1992; reared from dead ash limbs; E. G. Riley; OSUC 0020202; ¢ (TAMU). Uvalde Co., 10 mi N of Uvalde; 19-VI-1968; G. H. Nelson; on dead limbs; on Prosopis juliflora (Sw.)DC; VOLUME 105, NUMBER 2 OSUC 0020209; 5 (FSCA). Val Verde Co., Devil’s River, Dolan Falls area, elev. 360 m; 18-V-1993; Gelhaus No. 589, Nelson, Koenig; OSUC 0020311; 2 (ANSP). One additional non-type female without locality labels from UAIC. Megischus brunneus Cresson (Figs. 7, 14) Megischus brunneus Cresson 1865: 84 (original description); Townes 1949: 364, 369 (generic transfer, redescription, dis- tribution, figure, keyed): Carlson 1979: 141 (listed, generic transfer, distribution); Gauld, 1995: 184 (listed, distribution south of U.S.A.). Stephanus bruneus: Dalla Torre 1902: 6 (listed, generic transfer, distribution, un- justified emendation). Stephanus brunneus: Enderlein 1905: 475 (listed, distribution); Kieffer 1908: 4 (list- ed, distribution); Elliott 1922: 716, 718, 725 (description, keyed); Orfila 1951: 273 (generic transfer, keyed). Distribution.—Seven specimens from southernmost Florida (Monroe, Dade coun- ties: AEIC, FSCA, USNM); extralimital material from Cuba, Fig. 14. Biological associations.—Collected on dead white mangrove, Laguncularia race- mosa (Linnaeus) Gaertn. f. (Combretaceae); “at” American sycamore, Platanus occi- dentalis L. (Platanaceae); resting on dead fig, Ficus L. (Moraceae). Recorded in lit- erature in association with buttonwood (Townes 1949), possibly Conocarpus erec- tus L. (Combretaceae); and “‘diseased Jal- ia’? (Cresson 1865). Megischus californicus Townes (Figs. 14, 18) Megischus californicus Townes 1949: 364, 1 2. (original description, proton- um figured, keyed); Carlson 1979: 741 (listed, distribution); Halstead 1986: 101 (distribution, seasonality in California, cited as californica). 481 Distribution. —Oregon, California, Ne- vada, Utah, Colorado (56 specimens: AEIC, CDAE, CNCI, EMEC, EMUS, LACM, ODAC, OSUC, UCDC, UCRC, USNM). Biological associations.—Emerged from buckbrush, Ceanothus cuneatus (Hook.) Nutt. (Rhamnaceae); interior live oak, Quercus wislizeni A. DC.; and an unspeci- fied oak, Quercus L. (Fagaceae). Collected on mule’s fat, Baccharis salicifolia (Ruiz & Pavon) Pers. (cited as B. glutinosa) (Aster- Ceanothus and Ceano- aceae); cuneatus, thus L. Megischus texanus Cresson (Figs. 16, 18) Megischus texanus Cresson 1872: 190, @ (original description); Elliott 1922: 761 (possible placement in Hemistephanus Enderlein); Townes 1949: 364 (figure, description, distribution); Carlson 1979: 741 (listed, distribution); Halstead 1986: 103 (distribution, seasonality in Califor- nia; biology); Gauld 1995: 184 (distri- bution south of U.S.A.). Stephanus texanus: Dalla Torre 1902: 9 (ge- neric transfer, distribution); Elliott 1922: 7G; 730 distribution, keyed). Stephanus (Hemistephanus) texanus: Ro- man 1917: 14 (keyed, subgeneric assign- ment). Hemistephanus texanus: Elliott 1922: 755 (generic transfer, similar to Parastepha- nellus pictipes Roman). (description, Distribution.—California, Nevada, Ari- zona, New Mexico, Colorado, Oklahoma, Texas, Louisiana, Fig. 16 (124 specimens: AEIC, AMNH, ANSP, CHAH, CNCI, CSUG, EMEC, EMUsS, FCDA, LACM, LSUC, MCPM, ODAC, OSUC, SEMC, SEV S, TAMU,,.UAIC, -UCDE, UCRE; UGCA, USNM, WFBM). Biological associations.—Reared as a lar- val parasitoid of Acmaeoderopsis junki (Thery) (Coleoptera: Buprestidae). Emerged from paloverde species, Parkinsonia Linnae- us, (Fabaceae; recorded on label as Cerci- 482 dium Tulasne); blue paloverde, P. florida (Benth. ex Gray); desert ironwood, Olneya tesota Gray (Fabaceae); mesquite, Prosopis Linnaeus (Fabaceae), interior live oak, Q. wislizeni. Collected on paloverde; pine, Pi- nus Linnaeus (Pinaceae), mesquite; oak; and sumac, Rhus Linnaeus (Anacardiaceae). Col- lected on site dominated by pine. Comments.—Small specimens of M. tex- anus do not have the sculpture and color patterns typical of the species. They tend to be darker and have coarser pronotal sculp- ture, and they may easily be mistaken for M. californicus. The relative length of the maxillary palp and the pilosity on the me- tapleuron are stable, size-independent char- acters not mentioned in Townes (1949). Megischus texanus may be distinguished from M. californicus by having the maxil- lary palp as long as the height of the head and the presence of a dense tuft of hairs centrally on the metapleuron. ACKNOWLEDGMENTS Thanks to the many curators for the loan of specimens for study; Luciana Musetti provided important help with the databasing of specimens and useful suggestions to the manuscript, and Glaucia Marconato skill- fully prepared all drawings. This material is based upon work supported by the National Science Foundation under Grant No. DEB- 9521648 and by a scholarship from CAPES (“‘Fundagao Coordenagao de Aperfeigoa- mento de Pessoal de Nivel Superior’), Bra- zil. LITERATURE CITED Aguiar, A. P. 1998. Revisao do genero Hemistephanus Enderlein, 1906 (Hymenoptera, Stephanidae), com consideragdes metodologicas. Revista Brasi- leira de Entomologia 41: 343-429. Aguiar, A. P. and A. Sharkov. 1997. Blue pan traps as a potential method for collecting Stephanidae (Hymenoptera). Journal of Hymenoptera Research 6: 422-423. Ashmead, W. H. 1900a. Classification of the ichneu- mon flies, or the superfamily Ichneumonoidea. Proceedings of the United States National Muse- um 23: 1-220. . 1900b. Hymenoptera. Stephanidae, p. 597. In PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Smith, J. B., ed. Insects of New Jersey. A List of Species Occurring in New Jersey, with Notes on those of Economic Importance. Supplement to the twenty-seventh annual report of the State Board of Agriculture 1899. Trenton, New Jersey. 755 pp. Carlson, R. W. 1979. Stephanidae, pp. 740-741. In Krombein, K. V., P. D. Hurd Jr., D. R. Smith, and B. D. Burks, eds. Catalog of Hymenoptera in America North of Mexico, Vol. Institution Press, Washington, DC. xvi + 1198 pp. Cresson, E. T. 1865. Catalogue of Hymenoptera in the collection of the Entomological Society of Phila- delphia from Colorado Territory. Proceedings of the Entomological Society of Philadelphia 4: 242— 313. . 1872. Hymenoptera Texana. Transactions of the American Entomological Society 4: 153-292. . 1880. [New species of Hymenoptera-No orig- inal title]. Transactions of the American Entomo- logical Society Proceedings 8: XVU-—X VIII. . 1887. Synopsis of the Families and Genera of the Hymenoptera of America, North of Mexico, Together with a Catalogue of the Described Spe- cies, and a Bibliography. Transactions of the 1. Smithsonian American Entomological Society Supplement. 351 pp. Dalla Torre, C. G. de. 1902. Catalogus hymenoptero- rum hucusque descriptorum systematicus et syn- onymicus. Volumen III: Trigonalidae, Megalyri- dae, Stephanidae, Ichneumonidae, Agriotypidae, Evaniidae, Pelecinidae. Sumptibus Guilelmi En- gelmann, Lipsiae. 1141 pp. Davis, G. C. 1897. Descriptions of new species of Tri- Stephanidae and Transactions of the American Entomological So- ciety 24: 349-372. Elliott, E. A. 1922. Monograph of the hymenopterous family Stephanidae. Proceedings of the Zoological Society of London 92: 705-831. Enderlein, G. 1905. Uber die Klassifikation der Ste- phaniden. Zoologische Anzeiger 28(13): 473-477. . 1906. Neue Beitriige zur Kenntnis und Klas- sifikation der Stephaniden. Stettiner Entomologis- che Zeitung 47: 289-306. Gauld, I. D. 1995. Stephanidae, pp. 181—184. Jn Han- son, P. and I. D. Gauld, eds. Hymenoptera of Cos- ta Rica. Oxford University Press, Oxford. 893 pp. Halstead, J. A. 1986. Distribution and seasonality of Megischus spp. (Hymenoptera: Stephanidae) in California. Entomological News 97: 101-103. Kieffer, J. J. 1908. Insecta. Hymenoptera (Fam. Ste- phanidae). Genera Insectorum 77: 1—10. Kirk, A. A. 1975. Siricid woodwasps and their asso- ciated parasitoids in the southwestern United States. The Pan-Pacific Entomologist 51: 57-61. Leonard, M. D. 1928. A list of the insects of New York, with a list of the spiders and certain other gonalidae, Ichneumonidae. VOLUME 105, NUMBER 2 allied groups. Cornell University Agricultural Ex- periment Station Memoir 101: 1—1121. Madl, M. 1991. Zur Kenntnis der palaéarktischen Ste- phanidae (Hymenoptera, Stephanoidea). Entomo- fauna, Zeitschrift ftir Entomologie 12(9): 117— 128. Mason, W. R. M. 1990. Cubitus posterior in Hyme- noptera. Proceedings of the Entomological Soci- ety of Washington 92: 93-97. Meyer, R. P, T. L. McKenzie, and K. Davis. 1978. Observations on a population of Schlettererius cinctipes (Cresson) (Hymenoptera: Stephanidae) in a selective cut of white fir (Abies concolor) in the Sierra Nevada of California. Natural control of Monochamus oregonensis, wood boring in- sects. The Pan-Pacific Entomologist 54: 326. Orfila, R. N. 1951. Sinopsis de los * Stephanus” neo- tropicos, con descripcion de una nueva especie de Argentina. Revista Brasileira de Biologia 11: 271— 274. Roman, A. 1917. Schlupfwespen aus Amazonien. Ar- kiv for Zoologi 11(4): 1-15. Schletterer, A. 1889. Monographie der Hymenopteren- Gattung Stephanus Jur. Berliner Entomologische Zeitschrift 33: 71-160. 483 Smith, D. R. 1997. Collections of Stephanidae (Hy- menoptera) in the mid-Atlantic states including an eastern record for Schlettererius cinctipes (Cres- son). Proceedings of the Entomological Society of Washington 99: 377-378. Taylor, K. L. 1967. The introduction, culture, liberation and recovery of parasites of Sirex noctilio in Tas- mania, 1962-67. CSIRO Technical Paper (Ento- mology) 8: 1-19. . 1976. The introduction and establishment of insect parasitoids to control Sirex noctilio in Aus- tralia. Entomophaga 21: 429-440. Townes, H. K. 1949. The Nearctic species of the fam- ily Stephanidae (Hymenoptera). Proceedings of the United States National Museum 99: 361-370. Viereck, H. L. 1914. Type species of the genera of ichneumon flies. Bulletin of the United States Na- tional Museum. 83: 1-186. Westwood, J. O. 1841. On the Evaniidae and some allied genera of hymenopterous insects. Annals and Magazine of Natural History (1)7: 535-538. . 1843. On Evania and some allied genera of hymenopterous insects. Transactions of the Royal Entomological Society of London 3(4): 237-278. PROC. ENTOMOL. SOC. WASH. 105(2), 2003, pp. 484-498 SEVEN NEW SPECIES OF POLYCENTROPODIDAE (TRICHOPTERA) FROM NICARAGUA AND COSTA RICA MARIA LOURDES CHAMORRO-LACAYO Department of Entomology, University of Minnesota, Saint Paul, MN 55108, U.S.A (e-mail: cham0138 @umn.edu) Abstract.—Seven new species of Polycentropodidae (Trichoptera) are described and illustrated from collections made in Nicaragua and Costa Rica: Cernotina riosanjuanen- sis, Cyrnellus zapateriensis, Polycentropus garfio, Polycentropus hamiltoni, Polycentro- pus phraterus, Polyplectropus maesi, and Polyplectropus nicaraguensis. Polycentropus holzenthali Bueno-Soria and Hamilton also is illustrated and redescribed. Key Words: tral America The family Polycentropodidae is com- prised of 372 species in 26 genera world- wide (Morse 2001). Six genera occur in the Neotropics: Antillopsyche Banks 1941, with 4 extant and | extinct species from the Greater Antilles; Cernotina Ross 1938 with 50 species and Cyrnellus Banks 1913 with 9 species are distributed only in the New World: and Nyctiophylax Brauer 1965, Po- lycentropus Curtis 1835, and Polyplectro- pus Ulmer 1905 with 4, 64, and 42 species, respectively, in the Neotropics (Flint et al. 1999). Costa Rica and Nicaragua have rep- resentatives of all these genera, except An- tillopsyche and Nyctiophylax. Thirty-five species of Polycentropodidae have been re- corded for Costa Rica and 19 for Nicara- gua: Cernotina, 2 species in Costa Rica, 4 in Nicaragua; Cyrnellus fraternus, (Banks) found in both countries; Polycentropus, 12 species in Costa Rica, 6 in Nicaragua; and Polyplectropus, 8 species in Costa Rica, 7 in Nicaragua (Flint et al. 1999). In this pa- per, I describe two new species of Polycen- tropus from Costa Rica and five new spe- cies in the genera Cernotina, Cyrnellus, Po- lycentropus, and Polyplectropus trom Nic- Trichoptera, Polycentropodidae, Nicaragua, Costa Rica, new species, Cen- aragua. In addition, Polycentropus holzenthali Bueno-Soria and Hamilton 1986 is illustrated and redescribed, and a new distribution record is provided. Dr. Ralph W. Holzenthal, University of Minnesota, and colleagues collected the new species from Costa Rica during an in- ventory of the caddisflies of that country conducted from 1986 through 1995. Dr. Jean-Michel Maes, Museo Entomologico, Leon, Nicaragua, provided the Nicaraguan caddisflies, which he collected during an ongoing inventory of Coleoptera and noc- turnal Lepidoptera of Nicaragua. I collected the remaining Nicaraguan caddisflies dur- ing two trips from June through August 2000 and from July through August 2001. Types are deposited in the University of Minnesota Insect Collection, Saint Paul, Minnesota (UMSP), the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (NMNH), and the Insti- tuto Nacional de Biodiversidad, Heredia, Costa Rica (INBIO). The terminology used in this paper follows that of Hamilton (1986) for Polycentropus and Polyplectro- pus, and Flint (1971) for Cernotina and Cyrnellus, with minor modifications. VOLUME 105, NUMBER 2 Cernotina riosanjuanensis Chamorro-Lacayo, new species (Fig. 1) This new species appears to be closest to Cernotina zanclana Ross 1951, but dif- fers in the more slender shape of the dor- sal branches of the intermediate append- ages, in the evenly rounded apex of the inferior appendages, and in the medial or- igin of the basodorsal lobes of the inferior appendages. Male.—Length of forewing 4.5 mm. Col- or in alcohol, yellowish brown. Genitalia as in Fig. 1. Sternum IX short, produced an- terolaterally; anterior margin, in ventral view, deeply excavated. Tergum X semi- membranous, long, narrow, setose; in lat- eral view, curved posterad; in dorsal view, curved posteromesally. Intermediate ap- pendages bipartite; dorsal branches with ba- ses broad, curved mesally and tapering into pigmented acute apices; ventral branches setose, in lateral view, digitate with broad bases and rounded thumblike apices, in dor- sal view, square, mesal margins deeply con- vex, posterior margins truncate, excavated, expanded posterolaterally. Preanal append- ages apparently absent. Inferior appendages elongate, oval, in lateral view, posterolat- eral margins rounded, apicoventral and ap- icomesal lobes concealed; basodorsal lobes clublike each with long, thick apical setae and a small posteriorly directed submedial process; apicoventral lobes pigmented, in dorsal and ventral views triangular; apico- mesal lobes, in dorsal view, square, trans- parent with posterior margins lightly pig- mented. Phallus long, slender, tubular, membranous internally. Female.—Unknown. Type material.—Holotype, ¢d. NICA- RAGUA: Rio San Juan, Refugio Bartola, small creek, 300 m NW of station, 10°58’N, 84°21'W, el. 35 m, 7.viii.2000, Chamorro, Dobbins (UMSP) (UMSP000066732). Etymology.—This species is named for the magnificent national treasure that is the 485 San Juan River (Rio San Juan), Nicaragua, by which this species was collected. Cyrnellus zapateriensis Chamorro-Lacayo, new species (Fig. 2) This new species is closely related to Cyrnellus fraternus (Banks 1905). Cyrnel- lus zapateriensis, can be distinguished from C. fraternus by the smaller size and truncate apices of the inferior appendages, and by the membranous ventrolateral margins of the intermediate appendages. Male.—Length of forewing 4.5 mm. Col- or in alcohol, body and legs yellowish brown; wings brown. Male genitalia as in Fig. 2. Sternum IX, in lateral view, reversed L-shaped, apparently fused with interme- diate appendages dorsally. Tergum IX and X membranous, fused with setose interme- diate appendages; fused structure, in lateral view, nearly square with ventrolateral mar- gins membranous and expanded posterad (in some specimens not as expanded, but reduced to a narrow apex), in dorsal view, trapezoidal. Body of bipartite preanal ap- pendages digitate, setose, not exceeding in- termediate appendages, produced mesally into digitate, posteroventrally curved me- sobasal processes, with pigmented apex, bordering phallus. Inferior appendages, in lateral view, broad basally, curved poster- odorsally and narrowing to rounded apices; in ventral view, posterior margins truncate, apicomesal lobes a narrow, acute mesally directed spine with pigmented apex; mesal margin concave basad of apicomesal lobes, expanding into medial setose circular lobes. Phallus short; phallic sclerite in lateral view somewhat indistinct, basally broad, slightly curved; in dorsal view distinct, Y-shaped. Subgenital plate digitate, fused basally, uni- form width throughout, apex lightly pig- mented, slightly curved ventrally, almost same size as mesobasal processes of preanal appendages. Female.—Unknown. Type material.—Holotype: 6. NICA- RAGUA: Granada: Isla Zapatera, El 486 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cernotina riosanjuanensis Fig. 1. Cernotina riosanjuanensis, male genitalia. A, Lateral view. B, Ventral view. C, Dorsal view. D, Inferior appendage, caudal view. E, Inferior appendage, dorsal view. VOLUME 105, NUMBER 2 487 2E Cyrnellus zapateriensis Fig. 2. Cyrnellus zapateriensis, male genitalia. A, Lateral view. B, Ventral view. C, Dorsal view. D, Caudal view. E, Phallus. EK Phallic sclerite, dorsal view. Bambu, Frente a Lago de Nicaragua, Same as holotype, 1 ¢ (NMNH); Granada: 11°45.829'N, 85°51.991’W, el. 42 m, Isla Zapatera, 20.v.1988, S. Valle, 1 6 19.vii.2001, Chamorro, Martinez, Ruiz (UMSP). (UMSP) (UMSPO000066725). Paratypes: Etymology.—This species is named for 488 the type locality, Isla Zapatera, located in Lake Nicaragua, Granada, Nicaragua. Polycentropus garfio Chamorro-Lacayo, new species (Fig. 3) This new species is a member of the gertschi group, as defined by Hamilton (1986). Polycentropus garfio, is similar to Polycentropus spicatus Yamamoto 1967 and Polycentropus zurqui Holzenthal and Hamilton 1988 (both unplaced species within the gertschi group) in that the in- termediate appendages have laterally di- rected apices and bear a large basodorsal tooth, and in the overall shape of the pre- anal appendages. Polycentropus garfio is distinguishable from these species by the tight curvature of the intermediate append- ages, the overall shape of the inferior ap- pendages, the presence of paired plates within the phallus, and by the shape of the mesoventral process of the preanal append- ages. Male.—Length of forewing 7.5—8.0 mm. Color of body brown, legs with darker setae on femora and tarsi; dorsum of head brown with long, erect, dark brown setae; fore- wings covered by brown setae and scattered patches of golden setae. Male genitalia as in Fig. 3. Sternum IX, in lateral view, with anterior margin nearly straight, broad ven- trally, tapered posterodorsally and posterior margin moderately sinuate; in ventral view, rectangular. Tergum X membranous, basal margin broad in dorsal view, trapezoidal. Intermediate appendages thick basally, with basodorsal tooth, main body tightly curved anterolaterally tapering into acute apex. Preanal appendages dorsally positioned and apically truncate, tapering into narrow me- soventral processes; processes curved pos- teroventrally, apices thin and pointed. In- ferior appendages, in lateral view, triangu- lar, bearing dorsally directed mesoventral tooth, mesal ridge with numerous thick spine-like setae, in ventral view, oval shaped, expanded laterally, mesoventral tooth directed medially. Phallobase long; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON apicoventral process broad basally, sharply narrowed apically with pair of small subap- icolateral teeth; four to ten phallic spines embedded in endothecal membrane; phallic sclerite, in dorsal view, rectangular, broad anteriorly, anterior margin concave; endoth- ecal membrane with pair of large dorsal in- ternal triangular plates meeting dorsome- sally. Subphallic sclerite broad. Female.—Unknown. Type material—Holotype: ¢. COSTA RICA: Alajuela: Rio Agrio, ca. 3.5 km NE Bajos del Toro, 10.243°N, 84.279°W, el. 1,290 m, 20.vii.1990, Holzenthal, et al. (UMSP) (UMSP000049376). Paratypes: Same data as holotype, 57 6 (UMSP); COSTA RICA: Alajuela: Rio Toro, 3.0 km (road) SW Bajos del Toro, 10.204°N, 84.316°W, el. 1,530 m, 3—4.ix.1990, Hol- zenthal, Blahnik, Huisman, 2 6 (INBIO); Heredia: Parque Nacional Braulio Carrillo, Estacioén Magsasay, Rio Peje, 10.402°N, 84.050°W, el. 130 m, 25—26.vili.1990, Hol- zenthal, Blahnik, Huisman, 8 d (UMSP), 3 do (NMNH); San José: Parque Nacional Braulio Carrillo, Estacion Carrillo, Quebra- da Sanguijuela, 10.160°N, 83.963°W, el. 800 m 27.11.1987, Holzenthal, Hamilton, Heyn, 1 6 (UMSP). Etymology.—This species is named for the diagnostic, sharply curved body and acute apex of the intermediate appendage, which resembles a garfio, Spanish for hook. Used as a noun in apposition. Polycentropus hamiltoni Chamorro-Lacayo, new species (Fig. 4) Polycentropus n.sp. 10: Hamilton 1986: 110; Holzenthal and Hamilton 1988: 335. This new species is a member of the bo- nus complex of the gertschi group and is closely related to Polycentropus fortispinus Holzenthal and Hamilton 1988. Polycentro- pus hamiltoni differs from P. fortispinus in that the bodies of the preanal appendages are approximately equal in size to the in- ferior appendages. It also differs in the 489 VOLUME 105, NUMBER 2 Rt Z Ss ae S [ing v < 3D re) 2) Fig. 3. Polycentropus garfio, male genitalia. A, Lateral view. B, Dorsal view. C, Ventral view. D, Inferior appendage, caudal view. E, Phallus. F Phallic sclerite, dorsal view. G, Apex of apicoventral process of phallus, dorsal view. 490 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Polycentropus hamiltoni Fig. 4. Polycentropus hamiltoni, male genitalia. A, Lateral view, everted tergum X. B, Dorsal view everted tergum X. C, Ventral view. D, Inferior appendage, caudal view. E, Phallus. smaller and more medially positioned me- nally, the numerous phallic spines of P. soventral processes of the preanal append- hamiltoni are longer and thinner than in P. ages, the slightly smaller intermediate ap- fortispinus. pendages, and the strongly ovoid and less Male.—Length of forewing 5.0—6.6 mm. angularly shaped inferior appendages. Fi- Body generally pale brown to yellow be- VOLUME 105, NUMBER 2 low, dorsum of head and thorax dark brown with long, dark setae and some paler setae between antennal bases; forewing bases with long erect setae, with general vestiture of fine, dark brown setae, and numerous patches of fine golden setae scattered over wing. Male genitalia as in Fig. 4. Sternum IX in lateral view nearly rounded; in ventral view, rectangular, anterior and posterior margin broadly and shallowly concave. Ter- gum X membranous; short, in dorsal view rectangular. Intermediate appendages broad, fused basally to membrane below tergum X, tapering posteromesally to very thin api- ces, each with single apical seta; length not exceeding preanal appendages. Body of preanal appendages, in lateral view, nearly square, mesoventral process rod-like, orig- inating laterally, then directed anteriorly and then curving posteromesally. Inferior appendages short, in lateral view, nearly square appearing subquadrate; in ventral view appearing nearly elliptical, expanded posterolaterally; surfaces irregular with broad mesoventral tooth. Phallobase short with projecting ventral apex; apico- ventral process tapering to rounded ventral- ly directed apex; group of phallic spines embedded in endothecal membrane; phallic sclerite, in lateral view, broad anteriorly, curved posteroventrad to narrow apex, in ventral view narrow posteriorly and broad anteriorly. Subphallic sclerite extending lat- erally to contact preanal appendages baso- ventrally. Female.—Unknown. Type material.—Holotype: d. NICA- RAGUA: Jinotega: Cerro Mazu, 14°33'N, 85°07'W, el. 220 m, 7—10.1x.1997, J. M. Maes and B. Hernandez (UMSP) (UMSP000066734). Paratypes: MEXICO: Chiapas: Rro Contento, 7 km N. of Ocos- ingo, 20.v.1981, C. M. and O. S. Flint, Jr., 2 6 (NMNH). Etymology.—Named in honor of Dr. Steven W. Hamilton for his contribution to the study of Neotropical caddisflies, espe- cially in the family Polycentropodidae. mesal 491 Polycentropus phraterus Chamorro-Lacayo, new species (Fig. 5) This new species is a member of the bar- tolus complex of the gertschi species group and is closely related to Polycentropus ne- bulosus Holzenthal and Hamilton 1988. Po- lycentropus phraterus, ditfers from P. ne- bulosus in the triangular shape and_ the acute posteromesal point of the inferior ap- pendages and the gradual curvature of the intermediate appendages. After close ex- amination of the 5 paratypes of P. nebulo- sus in UMSP, it became apparent that 3 of these paratypes differ from the holotype and are actually P. phraterus. Consequent- ly, these 3 paratypes are here assigned as the paratypes series for P. phraterus. Male.—Length of forewings 6.0—6.5 mm. Body sclerites and setae generally pale brown to yellow; legs, especially femora and tarsi, with fine, dark brown setae; dor- sum of head and thorax dark brown with long, erect, golden-brown setae; forewings covered with fine, brown setae and with scattered patches and mottled areas of gold- en and brown setae; forewing bases with long erect, golden-brown setae. Genitalia as in Fig. 5. Sternum IX, in lateral view, with anterior margin nearly straight, ventral mar- gin broad, tapering posterodorsally, poste- rior margin moderately sinuate, posterov- entral corner rounded, in ventral view, rect- angular. Tergum X membranous, trapezoi- dal. Intermediate appendages broad basally, tapering into acute apices, gently curved mesoventrally. Preanal appendages in lat- eral view, nearly semicircular, mesoventral processes absent. Inferior appendages, in lateral view, triangular, ventral nearly straight, posteroventral corners an- gulate and bearing acute posteromesal pig- mented point; in ventral view, nearly tri- angular, posteromesal margins irregular, posterolateral margins straight, mesal mar- gins nearly straight, expanded laterally, with mesobasal spines. Phallobase short, apicoventral process broad basally, nar- margins 492 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Polycentropus phraterus Fig. 5: appendage, caudal view. E, Phallus. rowed apically; four phallic spines embed- ded in the endothecal membrane. Subphal- lic sclerite weakly developed. Female.—Unknown. Type material.—Holotype: ¢. COSTA RICA: Alajuela: Reserva Forestal San Ra- Polycentropus phraterus, male genitalia. A, Lz iteral view. B, Dorsal view. C, Ventral view. D, Inferior mon, Rio San Lorencito and _ tributaries, 10.216°N, 84.606°W, el 980 m, 6— 10.11.1991, Holzenthal, Munoz, Huisman (UMSP) (UMSP000049471). Paratypes: Guanacaste: Parque Nacional Guanacaste, Rio San Josecito, Estaci6n Mengo, VOLUME 105, NUMBER 2 10.922°N, 85.470°W, 28-29.vii.1987, el. 960 m, Holzenthal, Morse, Clausen, 1 d (NMNH), 1 6 (UMSP), 1 56 (INBIO). Etymology.—This species is named phraterus from the Greek phratero, broth- ers, in honor of Dr. Ralph W. Holzenthal and Dr. Steven W. Hamilton for their con- tributions to the study of Neotropical cad- disflies. Polyplectropus maesi Chamorro-Lacayo, new species (Fig. 6) This new species is a member of the t/i- lus group of Yamamoto (1967), which in- cludes Polyplectropus deltoides (Yamamoto 1967), P. carolae Bueno-Soria 1990, P. denticulus Bueno-Soria 1990, and P. thilus (Denning 1962), as defined by Bueno-Soria (1990). Polyplectropus maesi differs from the other members of this group in the fol- lowing ways: the ventrolateral processes of the preanal appendages are expanded pos- terad, each bearing two ventrally directed short subapical lobes: in the posterior mar- gins of the dorsolateral lobes of the inferior appendages are bilobed; and the inferior ap- pendages have broadly triangular spine bearing ventromesal lobes. Male.—Length of forewing 5.0 mm. Col- or in alcohol, yellowish brown. Genitalia as in Fig. 6. Sternum IX, in lateral view, short, deltoid, posterior margin sinuate, anterior margin produced ventrally; in ventral view, rectangular, anterior margin concave. Ter- gum X membranous, broad and elongate. Intermediate appendages digitate, setose, not exceeding inferior appendages. Preanal appendages tripartite; dorsolateral processes heavily sclerotized, long, directed antero- mesally, from base recurved posterolater- ally, then mesoventrally, finally postero- mesally tapering into acute point; mesola- teral processes rectangular, broad, setose, lightly sclerotized, produced into ventrolat- eral processes meeting mesally to form nar- row bridge below phallus; ventrolateral pro- cesses directed posteriorly, subapically each bearing two fused ventrally-directed highly 493 sclerotized lobes. Inferior appendages di- vided into two lobes; dorsolateral lobes, in lateral view, club-shaped with posterior margins bilobed, setose; ventromesal lobes short, in lateral view, triangular with round- ed ventral margins, setose, in ventral view, broadly triangular tapering mesally; mesal margins scalloped bearing robust sclero- tized spines, seven visible, in caudal view. Phallobase short, apically membranous; dorsal phallic sclerite cylindrical, narrowing apically; endophallus cylindrical, ventrally located, embedded in endothecal mem- brane. Female.—Unknown. Type material.—Holotype, ¢d. NICA- RAGUA: Zelaya: Rio Las Latas, 14°04'N, 88°33'W, el. 220 m,; 2.vi.1998, J: M. Maes and B. (UMSP) (UMSP0000667 24). Etymology.—It gives me great pleasure to name this species in honor of Dr. Jean- Hernandez Michel Maes, Belgian entomologist, in rec- ognition of the many years he has dedicated to the study of Nicaraguan insects, for his unrelenting efforts, and for his wholeheart- ed support, collaboration, and friendship. Polyplectropus nicaraguensis Chamorro-Lacayo, new species (Fig. 7) This new species is a member of the charlesi group, as defined by Bueno-Soria (1990) and is closely related to Polyplec- tropus mignonae Bueno-Soria 1990. Poly- plectropus nicaraguensis resembles P. mig- nonae in the overall shape of the inferior appendages, in the reduction of the dorso- lateral processes of the preanal appendages, and in the shape of the intermediate ap- pendages and phallus. Polyplectropus_ ni- caraguensis can be distinguished from P. mignonae by the shape and position of the ventrolateral processes of the preanal ap- pendages and by the lack of spines on the posterior margins of the ventromesal lobes of the inferior appendages. Male.—Length of forewing 4.5—5.0 mm. Color in alcohol yellowish brown. Genitalia 494 Fig. 6. as in Fig. 7. Sternum IX, in lateral view, short, deltoid, posterior margin sinuate, an- terior margin nearly straight. Tergum X membranous, short, in dorsal view, rectan- gular, in lateral view deltoid. Intermediate appendages digitate, setose, directed dor- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Polyplectropus maesi, male genitalia. A, Lateral view. B, Ventral view. C, Dorsal view. D, Caudal view. E, Phallus. solaterally. Preanal appendages bipartite and migrated dorsally; mesolateral process- es short, setose, oval, broadly tapering pos- terodorsally, produced into ventromesal processes which meet mesally below phal- lus, forming narrow bridge; ventromesal VOLUME 105, NUMBER 2 495 Polyplectropus nicaraguensis Fig. 7. Polyplectropus nicaraguensis, male genitalia. A, Lateral view. B, Ventral view. C, Dorsal view. D, Caudal view. E, Phallus. processes round, setose, pigmented. Inferior appendages divided into two lobes; dorso- lateral lobes, in lateral view, long, slender with median hump, posterior margin trun- cate, bearing ventrolateral spine; ventro- mesal lobes short, in lateral view, broad ba- sally tapering to truncate apex, in ventral view, nearly rectangular, basally broad, di- 496 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Polycentropus holzenthali Fig. 8. Polycentropus holzenthali, Bueno and Hamilton 1986, male genitalia. A, Lateral view. B, Dorsal view. C, Ventral view. D, Phallus. E, Inferior appendage, caudal view. rected laterally. Phallus narrow; phallic sclerite cylindrical; phallus surrounded by membrane, situated well dorsally in genital capsule. Female.—Unknown. Type material.—Holotype, ¢. NICA- RAGUA: Zelaya: Rio Las Latas, el. 220 m, 14°04’N, 88°33’W, 2.vi.1998, J. M. Maes and B. Hernandez (UMSP) (UMSP000066727). Paratype: Same data as holotype, | d (NMNH). Etymology.—This species is named for VOLUME 105, NUMBER 2 the country where the types where collect- ed. Polycentropus holzenthali Bueno-Soria and Hamilton 1986 (Fig. 8) Polycentropus holzenthali Bueno-Soria and Hamilton 1986: 300. [Type locality: MEXICO: Chiapas: Tributario del Rio Teapa situado en la carretera 195 a 3 km al N de Ixhuatan; NMNH, See recent Proceedings for style of figure legends. Literature Cited.—Give only papers re- ferred to in the text; list alphabetically by author. Spell out references (journal names etc.) completely, including conjunctions and prepositions—no abbreviations. Do not underline; journal names and book titles are set in Roman type. See recent issues for styles for journals, books, book chapters, etc. When referring to references in text, use the following; Henry (1990), (Henry 1990), (Henry 1987, 1990), (Smith 1990, Henry 1992), (Smith 1990; Henry 1990, 1991), (Miller 1990a, 1990b) where two articles are published by the same author in the same year, or Henry et al. 1990 where more VOLUME 105, NUMBER 2 than two authors are involved (do not ital- icize *“‘et al.”*). Citations *‘in press’’ should be cited as “‘in press,’’ not with year of expected publication. Names of persons providing unpublished information should include initials and be referenced in the text as: W. Mathis (per- sonal communication) [information ob- tained orally] or W. Mathis (in litt.) [infor- mation obtained in a writing]. Names of organisms.—The first men- tion of a plant or animal should include the full scientific name including the authority. Use only common names approved in Com- mon Names of Insects and Related Organ- isms published by the Entomological Soci- ety of America. Spell out the entire scien- tific name the first time it is mentioned, thereafter abbreviate generic names; how- ever, do not abbreviate a genus name at the beginning of a paragraph or sentence, or if two or more genera being discussed have the same first letter. Within sentences, al- ways use the genus name or initial preced- ing a species name. Taxonomic papers.—Taxonomic papers must conform to requirements of the Inter- national Code of Zoological Nomenclature. In addition, type specimens must be desig- nated for new species described, type de- positories must be clearly indicated, and new taxa must be clearly differentiated from existing taxa by means of keys or dif- ferential diagnoses. In short, these papers must conform to good taxonomic practices. Book Reviews.—Send to Book Review Editor (see inside front cover) and see re- cent Proceedings for format. Charges.—Authors will be assessed a charge to cover editing and publication ex- penses. The current charge to members is $40.00 per printed page. These charges are in addition to those for reprints and author’s correction in proof. The charge for pages in excess of 15 printed pages, for non-mem- bers, and papers for immediate publication is $60.00 per printed page. Member authors who are retired or not affiliated with an in- stitution may request to have charges waived and should do so when the manu- script is submitted. A complete or partial waiver may delay publication. Charges for author errors and/or changes in proof, for reprints if ordered, for immediate publica- tion, and for non-members cannot be waived. Acceptance of papers is based only on scientific merit without regard to the au- thor’s financial support. ENTOMOLOGICAL SOCIETY OF WASHINGTON http://entomology.si.edu/ES W/ES W Menus. lasso Infe~ >ation and Officers Bylaws History First 100 Years Past Presidents History of the ESW Seal Information for Authors Available Publications Meetings Members Database Membership Application Subscription PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael E. Schan fis Sippy 99 Oe sce Sees ee eS ee ee ee ee A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric Grisselitand Michaelibsschautty 87) pp! 1997, = =e ee eee Revision of the Oriental Species of Aphthona Chevrolat (Coleoptera: Chrysomelidae), by Alexander S. KKonstantinoy andi Steven W: Linpatelter, 349!pp. 200226 en ee ee eee Revision of the Genus Anoplophora (Coleoptera: Cerambycidae), by Steven W. Lingafelter and E. Richard (Hoebeke:.236;pp: 2002.2 Ses = he. ee ee ee 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 __. 6 The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi Makahasr 230 pp O69 ce se se RO Ss 5 ee ee ee a Se ee No. 8. The North American Predaceous Midges of the Genus * »omyia Meigen (Diptera: Cerato- pogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 ; FU ete sy ANG ee Ee eee ee No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: ( alcidoidae), by Michael E. Schauff. GTMPP OSA: Tee See eee TR ae re fot os eee eo ke en i No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 PPP L995 | 2 8 ie a a oe ee No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent AUC RW el Grogan shirulO Sypps USO Sissies ee oe ee ae ee ee ee No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 __ No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited by.B..BsNordenjandiAss: Menke: 2116 ppylooon 2s a ee ee eee 25.00 No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and Willtaniicy Grocanh Ie297 ps WOOT, Sie ee es ee ae he ne eee No. 19. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by Thomas]. Henryl49ipp: 1997) S'S i ee No. 20. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- afelterdUUSipp wl 8 ye see ee ee Ree ce oe ee hee fe ee ey ee eee No. 21. New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. Lexb rte VII GIO) GoTo pil Oe testa eae eee, ee cee ee a ee ene ee ear a Se ee eee No. 22. Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: inichogtammatidae)s by Joho DePinto: 28i7epp sl GOO eee ee ee eee No. 23. Revision of the Net-Winged Midges of the Genus Blepharicera Macquart (Diptera: Blepha- riceridae) of Eastern North America, by Gregory W. Courtney. 99 pp. 2000 ____.- No. 24 Holcocerini of Costa Rica (Lepidoptera: Gelechioidea: Coleophoridae: Blastobasinae), by David Wdarnskiv P47 pp, 2002) cee Wie te ee Se ae ee ee RS ee a $10.00 15.00 40.00 30.00 $15.00 15.00 15.00 15.00 12.00 5.00 25.00 25.00 25.00 25.00 18.00 12.00 12.00 28.00 14.00 18.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume to non-members and $25.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra on al] orders. Dealers are allowed a discount of 10 percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Custodian, Entomological Society of Washington, “ Department of Entomology, MRC 168, Smithsonian Institution, Washington, D.C. 20560-0168. CONTENTS (Continued from front cover) McKAMEY, STUART H.—Review of the New World leafhopper genus Excultanus Oman (Hemiptera: Cicadellidae: Deltocephalinae) McKAMEY, STUART H.—Some new generic names in the Cicadellidae (Hemiptera: Deltocephalinae, Selenocephalinae) PIKE, KEITH S. and PETR STARY—Ericaphis louisae, n. sp. (Hemiptera: Aphididae: Aphidinae: Macrosiphini) on Luetkea pectinata (Pursh) (Rosaceae), and a key to Ericaphis species .... PIKE, KEITH S., P. STARY, and G. GRAF—Description of a new species of Braconidae, Monoctonus allisoni (Hymenoptera: Braconidae: Aphidiinae), and discussion of the aphid- parasitoid guild for Nasonovia spp. (Hemiptera: Aphididae) in northwestern United States .. POGUE, MICHAEL G. and J. BOLLING SULLIVAN—Re-evaluation of the Elaphria fes- tivoides (Guenée) species complex (Lepidoptera: Noctuidae) QI, BAOYING, CARL W. SCHAEFER, NONNAIZAB BAI, and ZHEMIN ZHENG—Miridae (Heteroptera) recorded from China since the 1995 World Catalog by R. T. Schuh RABAGLIA, ROBERT J.—Annotated list of the bark and ambrosia beetles (Coleoptera: Scolytidae) of Maryland, with new distributional records RABAGLIA, ROBERT J. and MICHAEL A. VALENTI—Annotated list of the bark and ambrosia beetles (Coleoptera: Scolytidae) of Delaware, with new distributional records .... ROBINSON, HAROLD—Five new species of Xanthina Aldrich (Diptera: Dolichopodidae) from Mexico and Central America ; SHAFFER, JAY C.—New Peoriini (Lepidoptera: Pyralidae: Phycitinae) from Brazil WHEELER, A. G., JR.—Plagiognathus reinhardi Johnston (Hemiptera: Miridae): Distribution, habits, and seasonality of a hawthorn (Crataegus) specialist YOUNG, CHEN W. and C. DENNIS HYNES—Biology and immature stages of the crane fly Ptilogyna (Plusiomyia) herroni (Alexander) (Diptera: Tipulidae) from New Caledonia, with discussion of its phylogenetic placement NOTES POINAR, GEORGE, JR.—Cecidomyiidae (Diptera) as insect parasitoids ROBBINS, RICHARD G., SHARON L. DEEM, and JAMES L. OCCI—First report of Amblyomma humerale Koch (Acari: [xodida: Ixodidae) from Bolivia, with a synopsis of col- lections of this tick from the South American yellow-footed tortoise, Geochelone denticulata (L.) (Reptilia: Testudines: Testudinidae) BOOK REVIEW NICKLE, DAVID A.—Katvdids & Bush-Crickets. Reproductive Behavior and Evolution of the Tettigoniidae, by Darryl T. Gwynne OBITUARY NORRBOM, ALLEN N. and F. CHRISTIAN THOMPSON— Richard Herbert Foote (1918-2002) REPORTS OF OFFICERS 452 499 i a aye erie rath IAN INSTITUTION LIBRARIES INEM 3 9088 01073 7948