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Me ce = S ois 2 = a NI NVINOSHLINS S3/Y¥VYdIT LIBRARIES SMITHSONIAN INSTITUTION NOILANLILSNI_NVINOSI Baye z a er ee z ns o : = wo = he o — o oS 3 : : — GY ° eZ 2 \ | > T= > e” fh ti. P > S i) = Bs] = Ci Da 0 = ee] S'S = oD — we Ae le - SENN o Zz m Z o Z A a — w —_ _— ES SMITHSONIAN INSTITUTION NOILOLILSNI NVINOSHLINS S3I¥VYgIT_LIBRARIES SMITHS( ” zy a7) = 77) : ala ” = < = < = a = = z a) z a ty, = = ‘ o D ra AD a YJ D . os P= fe g = SAP Fh OE 2 ce ee aes 3 = Si ee INI_NVINOSHLINS | Sa byYvdagit LIBRARI ES SMITHSONIAN INSTITUTION pNOILALILSNI_ NVINGS! ms 2 > > =a Tr) [ee uJ , Lu w : . = “ ee = = NN 5} Berd oa x tL" fii aA < ra, te ‘ier ox c “Ge h S oc Sy a = m0 = Pa ae ih ial. Pa cay ae S = 3 = S = gui a) ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IYVYEIT LIBRARIES SMITHS ro) = it res = _— ° = s) wo 2 \ 5 2 5 2 5 > 5 Ap, | E = = > 2 > Pf y & i = 2 ra a K Gy, m = m eu m 2 Uf = =, Ww 5 cam — SNI NVINOSHLINS S3SIYVYEIT LIBRARIES SMITHSONIAN NOILNLILSNI NVINOS z 7) a as ” z 7) z fz = 5 = < = =< A> = z = z = = re) = ; 2 zB B z 2 | a 2 E z iE z = = ie = > = > = > , = ers |e 2 hi ia CE eae DE emrene of A A a A eID ganas ae iat ee mmani aoe { oF Ai Mi) Nf as Me va i oe a may AY a ep ss ae i, ie a i my, di. z i ta Pah ae net ie . ee 4 ray ae / : ' eG yy ey, AV tb) Tso i 4 s oe i Hiy j i QL 46\ VOL. 97 JANUARY 1995 NO. | EGIK (ISSN 0013-8797) ~-: PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED QUARTERLY CONTENTS DIETRICH, CHRISTOPHER H. and STUART H. MCKAMEY—Two new neotropical treehop- per genera and investigation of the phylogeny of the subfamily Membracinae (Homoptera: Memb ra cidaes) ori wii atncc a near hattitiea lu os eT SACI iW sty Ue ROR Vee a A Rue Mit ae l EVANS, GREGORY A., MICHAEL E. SCHAUFF, MOH LENG KOK-YOKOMI, and RAY- MOND K. YOKOMI—A new species of Aphelinus (Hymenoptera: Aphelinidae) that para- sitizes the spirea aphid, Aphis spiraecola Patch (Homoptera: Aphididae)................ 17 FITZGERALD, SCOTT J. and BORIS C. KONDRATIEFF—A review of the mydid genus Pseudonomoneura Bequaert (Diptera: Mydidae), with the description of two new species... 22 FLOWERS, R. WILLS—Hermesia Lefévre, a resurrected genus of Neotropical Eumolpinae GeMlempterda-(CHTVSOMEMGAG) cc iirc. n tics Asche lye shh ici alah ahah s layattanel aoe ek val ae (ound RAHA Cay we eed Bp) FLOYD, MICHAEL A.—The larva and pupa of the caddisfly species, Helicopsyche paralimnella Hamilton. (icichoptera: Helicopsychidae):: ii 6c: .5 5) bese As atate age elas seat ata fe als acres Pot dle ae 46 GOULET, HENRI and DAVID R. SMITH—Four new sawflies from eastern North America, three species of Tenthredo and one of Dolerus (Hymenoptera: Tenthredinidae)........... 50 HARRISON, T. and S. PASSOA—Mirabilis-feeding Heliodines (Lepidoptera: Heliodinidae) in Cera illinois; WithGescriptlOi GF ANEW SPECIES (0! M03) Aleve baie Me elgiavact 4 eh ofa el lara 63 HEISS, ERNST—Atactocoris perneri, n. sp., a new apterous Carventinae from Jamaica (Het- CRODUC A AROULOAG) 5 calc tsincs diel lemata Stree ns nseyy $eapatis MOREE SAM Aten ta ROL REE ulema cha ob Alatic 0, Uaioe 71 KROMBEIN, KARL V.—Systematic notes on some Sri Lankan Scoliidae (Hymenoptera: PRCATLE ALA) Fea cee dd eee ee SRA oa Nieman ON Gln. coemetate ha cokoa tal slit oi AMIN Honea awa gaits Head a tadehay aie wel, Hae AMle meal nc as 77 KROMBEIN, KARL V. and BETH B. NORDEN—Notes on the behavior and taxonomy of Megachile (Xeromegachile) brimleyi Mitchell and its probable cleptoparasite, Coelioxys (Xerocoelioxys) galactiae Mitchell (Hymenoptera: Megachilidae) ..................... 86 LATTIN, JOHN D., ANNE CHRISTIE, and MICHAEL D. SCHWARTZ—Native black grass bugs (/rbisia-Labops) on introduced wheatgrasses: Commentary and annotated bibliogra- Pay CAemupleran creme pleka: eMC AE) i, Mar cis Nites Rene el Seno Nhe tert saUat chs Mca gets le uu matieG aided 90 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1995 JOHN W. NEAL, JR., President NORMAN E. WOODLEY, Treasurer RALPH P. ECKERLIN, President-Elect NATHAN SCHIFF, Program Chairman DARLENE D. JuDD, Recording Secretary M. ALMA SOLIS, Membership Chairman HOL Lis B. WILLIAMS, Corresponding Secretary PAUL J. SPANGLER, Past President JAMES PAKALUK, Custodian THOMAS J. HENRY, Editor DAVID R. SMITH, Associate Editor GARY L. MILLER, Book Review Editor Publications Committee DONALD R. DAVIS TERRY L. ERWIN F. CHRISTIAN THOMPSON A. G. WHEELER, JR., Special Publications Editor Honorary President CurTIS W. SABROSKY Honorary Members Louise M. RUSSELL ALAN STONE KARL V. KROMBEIN All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, NHB 168, Smithsonian Institu- tion, Washington, D.C. 20560. 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 8 P.M. Minutes of meetings are pub- lished regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. An- nual dues for members are $25.00 (U.S. currency). PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub- lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send ad- dress changes to the Entomological Society of Washington, % Department of Entomology, NHB-168, Smith- sonian Institution, Washington, D.C. 20560. Members in good standing receive the Proceedings of the Entomological Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscrip- tions are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remit- tances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE P. 218 OF THE JANUARY, 1991 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. Editor: Thomas J. Henry, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- West, Beltsville, MD 20705. Special Publications: A. G. Wheeler, Jr., Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, PA 17110-9408 Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 28 February 1995 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48- 1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 1-16 TWO NEW NEOTROPICAL TREEHOPPER GENERA AND INVESTIGATION OF THE PHYLOGENY OF THE SUBFAMILY MEMBRACINAE (HOMOPTERA: MEMBRACIDAE) CHRISTOPHER H. DIETRICH AND STUART H. MCKAMEY (CHD) Systematic Entomology Laboratory, USDA-ARS, % National Museum of Nat- ural History, Washington, D.C. 20560*; (SHM) Department of Ecology & Evolutionary Biology, U-43, University of Connecticut, Storrs, Connecticut 06269. Abstract.—Two new genera are described in the membracid tribe Membracini: Havi- landia, new genus, including H. pruinosa (Haviland), new combination, and H. hypso- proroides, new species; and Lewdeitzia, new genus, including L. lunata, new species. Havilandia is morphologically and behaviorally similar to Talipedini and some Leioscyta Fowler (Membracini), but may be the sister-group of Hypsoprorini. Lewdeitzia is appar- ently dimorphic for a feature of the forewing venation used to distinguish Erechtia Walker from Leioscyta. Preliminary cladistic analyses of 27 taxa in the subfamily Membracinae place the new genera among others in the tribe Membracini, but suggest that this tribe is paraphyletic, having given rise to other tribes in the subfamily. The analyses further suggest that the tribe Talipedini is paraphyletic and the genera Leioscyta and Membracis Fabricius are para- or polyphyletic. Reclassification of the Membracinae is needed, but should await elucidation of the limits of some genera of Membracini by more extensive cladistic anal- yses. A key to the tribes of Membracinae and the genera of Membracini is presented. Nomenclatural changes include restoration of the original spelling of Paragara Goding, 1926, and of the combination P. tholoidea (from Paragargara) and two new combinations, Paragara nigra (Funkhouser) (from Paragargara) and Enchenopa beebei (Haviland) (from Leioscyta). Two species most recently treated as Membracis are here considered Mem- bracini incertae sedis. Key Words: Cladistics, evolution, morphology, parsimony, paraphyly some tropical genera (Loye 1992, Wood 1993a) and may be explained partly by the The subfamily Membracinae constitutes a major component of treehopper diversity in the New World and is especially diverse in the tropics. The North American En- chenopa binotata species complex, whose females insert eggs into host plant tissue, has attracted attention as an example of host- mediated sympatric speciation (Wood 1993b). Polyphagy has been reported within * Correspondence: Christopher H. Dietrich, De- partment of Microbiology, Colorado State University, Ft. Collins, CO 80523. habit in many species, but in Membracinae only, of depositing egg masses on the host plant surface, thereby obviating many of the plant’s first-line structural and chemical de- fenses against herbivore colonization. Cladistic analyses of the family-group taxa of Membracidae (Dietrich and Deitz 1993; Dietrich, Deitz, and McKamey, unpubl.) support the monophyly of Membracinae, but, until now, there has been no explicit cladistic analysis of relationships among its 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON five tribes (Deitz and Dietrich 1993): Acon- ophorini, Hoplophorionini, Hypsoprorini, Talipedini, and Membracini. Nonetheless, three are supported by apparent synapo- morphies. The Aconophorini and Hoplo- phorionini are well-supported monophylet- ic’ groups (Dietrich and Deitz 1991; McKamey and Deitz, unpubl.). The species of Hypsoprorini are also united by apparent synapomorphies that are unique among the Membracinae: pronotum with lateral mar- gins of posterior process overlapping fore- wing at rest, and clavus of forewing acute and extensively associated with apical lim- bus. No synapomorphies have been found to support the monophyly of Talipedini or Membracini. The tribe Talipedini was erected (Deitz 1975) to draw attention to the position of Trinarea (as Talipes) appendiculata (Fon- seca) as morphologically intermediate be- tween the tribes Membracini and Hoplo- phorionini. Talipedini and Hoplophorionini both have clavate hind tibiae and small hind tarsi, a unique synapomorphy, but features used by Deitz (1975) to distinguish Tali- pedini from Hoplophorionini are present in Membracini, suggesting that Talipedini may be paraphyletic. McKamey and Deitz (1991) transferred two more species into Talipe- dini, but did not list additional diagnostic features for the tribe. The most diverse tribe in the subfamily is Membracini, which, now with 17 genera and nearly 200 described species, is twice as rich in genera and species as the next largest tribe, Hoplophorionini. Among the diagnostic features listed by Deitz (1975) for Membracini, none are unique to that tribe. Thus, Membracini (sensu Deitz 1975, ex- panded by McKamey 1992 and Sakakibara 1992), although defined by a unique com- bination of features, also may be paraphy- letic. Two new taxa were discovered that keyed to Membracini (Deitz 1975), but were not assignable to any known genus and have combinations of features traversing current concepts of tribes and genera. Species of Havilandia, new genus, share morphologi- cal features with species of Hypsoprorini, Hoplophorionini, Talipedini, and Acono- phorini. Lewdeitzia lunata, new genus, new species, may be sexually dimorphic for the forewing venation feature distinguishing Leioscyta Fowler from Erechtia Walker (tribe Membracini) and the female has a broad, obtusely rounded, marginally com- pressed, anterior pronotal horn similar to that of Aconophorini and some Hoplopho- rionini. The features of these new taxa ac- centuate problems with the generic and trib- al classification of the Membracinae. To help place the new taxa, we analyzed the rela- tionships among the genera of Membracini and representatives of other tribes of Mem- bracinae. Given the need of further taxonomic work revealed by the cladistic analysis, the nu- merous modifications to Membracini since the last key (Funkhouser 1951), and the dis- covery of some previously unknown or ne- glected intrageneric variation, we present a key to tribes of Membracinae and genera of Membracini to clarify current taxonomic concepts in the subfamily. KEY TO THE MEMBRACINAE 1. Forewing vein R initially divided into R, ,,,; and R,,, or venation reticulate, or with both CONGUIONS Se ee ee eae 2 1’. Forewing vein R initially divided into R, and Rs (=R,,3,4,;) (Figs. 1, 2), venation not re- ticulates.;.4 ee Oe ere 4 2. Metathoracic legs with tibiae clavate and with tarsi distinctly shorter than anterior tarsi .. RSs ne ED rns Minna cate Hoplophorionini! 2’. Metathoracic legs with tibiae not clavate and with tarsi as long as anterior tarsi (Fig. 2c) .. 3 3. Pronotum with lateral margins of posterior process overlapping portions of apical limbus and veins of forewing in repose; forewing cla- VuSracuiminate sn eee Hypsoprorini? ' Key to genera in prep. by McKamey and Deitz. 2 See Funkhouser’s (1951) key to genera of “‘Noto- cerini,” plus Jibarita Ramos and Hypsoprorachis Fon- seca and Diringshofen. VOLUME 97, NUMBER 1 SNe a 6’. 9¢ Pronotum evenly tapered posterolaterally, at most concealing part of apical limbus of fore- wing in repose; forewing clavus oblique api- cally Aconophorini? Metathoracic legs with tibiae clavate and with tarsi distinctly shorter than anterior tarsi; ab- domen with middorsal tuberosities Talipedini (monotypic: Trinarea Goding) Metathoracic legs with tibiae not clavate (Fig. 2c) or, if clavate, then abdomen without mid- dorsal tuberosities; metathoracic tarsi at least as long as anterior tarsi (Membracini) Metathoracic tibiae clavate, with dorsal edges flattened, and with cucullate setae numerous and small in row I, small or absent in row II (Fig. 1j) Havilandia, n. gen. Metathoracic tibiae with dorsal edges not flattened, with large cucullate setae in rows I and II (Fig. 2c) Pronotal metopidium with | to several oblique carinae on each side and with pair of dorsolateral carinae or dorsum strongly bi- sinuate posteriorly, or with all three condi- tions Pronotal metopidium without oblique cari- nae, with or without dorsolateral carinae, dorsum sublinear or once-sinuate posteriorly Re Re Pisin corer ae se ey cesses 13 Pronotal lateral carinae extending to postero- lateral margins or nearly so; pronotal integ- ument coarsely punctate (pits distinct) and thick Pronotal lateral carinae terminating before or above humeral angles; pronotal integument sometimes smooth (pits indistinct), mem- brane-like Pronotum bisinuate, declining stepwise from above humeral angle to posterior apex .... Tylopelta Fowler Pronotum declining evenly from above hu- meral angle to posterior apex ... Campylenchia Stal, Enchenopa Amy. & Serv. Pronotum and its horn depressed, pronotal integument thick and coarsely punctate (pits large, deep, and distinct above humeri) Kronides Kirkaldy Pronotum and its horn, if present, foliaceous, thin, and smooth (pits small, shallow, and indistinct above humeri at least) Pronotal horn present and distinct pe Ta ee eee Enchophyllum Amy. & Serv. Pronotal horn absent or indistinct Pronotal metopidium with a pair of dorso- 3 See Dietrich and Deitz (1991) for keys to genera and species. ie 12 13: 13%. 14. Wee ie}. . Forewing with only | r-m crossvein lateral carinae and a ventral lobe produced in front of the vertex of the head Peer GP EY LF be TY Se Se Phyllotropis Stal Pronotal metopidium without dorsolateral carinae or without ventral lobe, or with nei- ther feature Pronotal dorsum strongly foliaceous, sides extensively compressed into a single plate- like median carina Folicarina Sakakibara* Pronotal dorsum not so strongly compressed, sides not extensively fused .. Membracis (in part; e.g. M. carinulata Richter) Pronotum foliaceous and smooth (pits small, shallow, and indistinct above humeri at least); forewing surface coriaceous throughout (ar- cuate chaetoids conspicuous) Sa Ne Membracis (in part; e.g. M. foliata [L.]) Pronotum depressed, at least posterolateral- ly, and more coarsely punctate (pits larger, deeper, and distinct above humeri); distal forewing surface more glossy (arcuate chae- toids absent or inconspicuous) ........... 14 Anteromedial pronotal horn present (except in some ¢ Tritropidia, which are indistin- guishable‘from ieioscyia)) oe 2.2 15 . Anteromedial pronotal horn absent ....... 16 Pronotal horn narrow in lateral aspect; clyp- eus distally narrowed (as in Fig. 1h) Pee ate Ae ean ella neha nee n a Tritropidia Stal . Pronotal horn broad in lateral aspect; clypeus distally truncate (Fig. 2g) ... Lewdeitzia, n. gen. Forewing with 2 or more r-m crossveins .. 17 rer 18 Pronotum rugosely carinate, steeply decli- vous posteriorly, humeri without transverse Caninae:3s0 Fae ee Bolbonotodes Fowler Pronotum evenly carinate, gradually decli- vous posteriorly, humeri often with trans- VEISE CANINA wenn tee ere ter te Erechtia Pronotal dorsum with irregular gibbosities throughout, transversely carinate or gibbous subapically, and without long straight lateral carinae; mesothoracic tibiae foliaceous .... Bolbonota Amy. & Serv. . Pronotal dorsum without irregular gibbosi- ties, usually with | pair sublinear lateral ca- rinae extending posteriorly over dorsum; me- sothoracic tibiae cylindrical to foliaceous .. 19 Pronotal contour in lateral aspect strongly SINUAte we caee. Pte Stee ae eee Paragara? 4 New distribution record for Folicarina nr. bicolor: Trinidad, Arima Valley, St. Andrew’s Trace, 16-24 June 1981, leg. S.H. McKamey. ’ Paragara Goding, 1926a; original spelling here re- stored from Paragargara. Type species: Paragara tho- 4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 19’. Pronotal contour in lateral aspect linear or nearly so 20. Hind wing without r-m and m-cu crossveins, withavem)Mofreene => aneerieere Eunusa Fonseca 20'. Hind wing with | r-m and | m-cu crossvein Lee Ae Ee AE a a ee 72 Leioscyta DESCRIPTIONS OF NEw TAXA Morphological terms, techniques, and the convention for quoting labels follow Deitz (1975), Dietrich (1989), and Dietrich and Deitz (1991), except forewing crossvein s (that connects veins R,,; and R,,,;) equals “r” of Deitz (1975). Specimens for this study were provided by The Natural History Mu- seum, London (BMNH); North Carolina State University, Raleigh (NCSU); the per- sonal collection of S. H. McKamey (SHMC); and the United States National Museum of Natural History, Washington (USNM). Character states for taxa other than those described below are based on material iden- tified and labeled in the NCSU and USNM collections. Tribe Membracini Rafinesque Havilandia, NEw GENUS (Figs. la—k) Type species: Haviland. Diagnosis.—Metathoracic tibia clavate and compressed along dorsal edge, with se- tal row I bearing numerous small cucullate setae and row II with few or none; forewing with cell membranes clothed by erect mac- rotrichia, without crossvein s. Description.— Head: Vertex (Fig. lh) impressed mesad of ocelli, ventrolateral margins weakly produced; ocelli approxi- mately 2x as far from each other as from mesal margins of eyes and approximately 1 ocellar diameter from dorsal margin of ver- Tropidoscyta pruinosa loidea Goding, 1926a, by original designation. Re- stored combination: Paragara tholoidea Goding. New combination: Paragara nigra (Funkhouser 1940). Par- agargara (Goding 1926b) was an incorrect subsequent spelling and has no availability. Paragargara Goding, 1929, was an unjustified emendation and is an objec- tive junior synonym of Paragara. tex; frontoclypeus flat, diamond shaped, margins weakly produced, cibarial muscle scars parallel to ventrolateral margins; fron- toclypeus, clypellus, and rostrum, in lateral view (Figs. la, i) forming continuous arc, rostrum extended to base of abdomen; in anterior view with distance between lateral margins of eyes slightly less than distance between pronotal humeri. Thorax: Pro- notum (Figs. la, h-1) elongate and with mid- line strongly carinate and with 1 dorsolat- eral pair of strong carinae diverging from apex, extending to posterolateral margin of pronotum. Forewing (Figs. la, 1) punctate in basal half between veins C and M and in anal area, vein R initially divided into R, and Rs, crossveins s and r-m, absent, 2 m-cu crossveins present, apical limbus relatively narrow, contiguous with clavus for short distance only. Hind wing with crossveins r-m and m-cu present. Legs: Pro- and me- sothoracic tibiae (Fig. 11) compressed, translucent, margins of prothoracic tibia not expanded, mesothoracic tibia with poste- rior margin slightly expanded; mesothorac- ic coxa without acute process; metathoracic femur with pair of dorsoapical cucullate se- tae; metathoracic tibia (Figs. 1b, j) clavate, posterior margin compressed, row I with 15 or more small cucullate setae, row II with 5 or fewer, row III absent; all tarsi subequal in length; metathoracic tarsus relatively slender, tarsomere I with small apical cu- cullate seta. Abdomen: Sternum III without median tubercle, transverse carina indis- tinct or absent; terga without tuberosities or fenestrae. Male: Pygofer (Fig. 1d) with weak vertical lateral carina; lateral plate free, un- armed; aedeagus (Figs. 1 f—-g) with shaft slen- der, tapering apically, anterior face of apex evenly denticulate; gonopore membrane, in posterior view, occupying apical half of shaft; shank of style (Fig. le) slightly expanded preapically, shank sparsely setose, apex re- curved, acute, in posterior view oriented dorsolaterally; subgenital plate with preap- ical dorsal lobe. Female: Posterior margin of sternum VII with arcuate emargination; VOLUME 97, NUMBER 1 5 Fig. 1. Havilandia, new genus. a—g, H. pruinosa (Haviland). a—c, holotype ?: a, habitus, lateral view; b, left metathoracic tibia and tarsus, lateral view; c, second valvulae, lateral aspect; d—g, #7-90-337g 6; d, terminalia, lateral view (genitalia removed); e, shank of left style, ventrolateral view; f-g, aedeagus, posterior and lateral views, respectively. h-k, H. hypsoproroides, new species, holotype 2: h, head and pronotum, anterior view; i, habitus, lateral view; j, left metathoracic tibia and tarsus, lateral view; k, second valvulae, lateral view. avec, anteroventral carina; dlc, dorsolateral carina; dsc, dorsal submedial carina; p, pygofer; mc, median carina; Ip, lateral plate; sp, subgenital plate; tc, transverse sternal carina. 2nd valvulae (Figs. Ic, k) in lateral view gradually expanded toward midlength, then tapering slightly toward apex, dorsal margin with arcuate emargination preapically, without distinct crenulae or teeth. /ntegu- mental vestiture: Vertex and pronotum evenly punctate; forewing membrane with semi-erect pale setae. Abdominal terga coarsely punctate, most pits associated with lateral seta; acanthae simple to dentate. Notes and etymology. —Short-horned specimens of this genus resemble Leioscyta Fowler, but are distinguishable by the pres- ence of a pair of anteroventral longitudinal carinae on the pronotum and the non-foli- aceous front tibiae. Long-horned specimens resemble Hypsoprora Stal (Hypsoprorini), but lack apicolateral expansions on the pos- terior pronotal process. Although Havilan- dia best fits Deitz’s (1975) concept of the tribe Membracini, certain features of the new genus resemble those of other tribes. Like species of Hoplophorionini and Talipedini, Havilandia has clavate hind tibiae with a reduced number of cuculate setae in row II. Like some species of Hypsoprorini, the new genus has a ventrolateral pair of carinae on the pronotal horn. Generic recognition is based primarily on the tibial and abdominal features, but both species also have: vertex densely clothed with pale setae; thoracic pleuron pilose, se- tae clothed with white waxlike material; pronotum produced anterodorsally, with | ventrolateral pair of strong carinae diverg- ing from the apex; and forewings densely clothed with arcuate chaetoids. This genus is named in honor of Maud D. Haviland, a pioneer in the ecology and taxonomy of Neotropical Membracidae. Havilandia pruinosa (Haviland), NEw COMBINATION (Figs. la—g) Synonymy: Tropidoscyta pruinosa Hav- iland, 1925:237. Type locality: Kartabo, Cuyuni District, Guyana [BMNH]. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Material examined.— Holotype 2 [BMNH]. Other specimens: Nestor, Trinidad; San Mar- tin and Shapajilla, Peru; and Culebra, Ama- zonas, Venezuela [all USNM]; road to Foco Mine, 33 km SE El Dorado, Bolivar, Vene- zuela [SHMC]. Dates of collection: February, April, May, August. A teneral 6 specimen from Maron, French Guiana [Figs. | d—g; Dietrich Research #7-90-337g, NCSU] is probably conspecific with the holotype of H. pruinosa but lacks a pale macula on the forewing at the apex of vein Cu that is present in the holotype (Fig. 1a). Notes.—Females of H. pruinosa guard their eggs and are not ant-attended (Havi- land 1925), characteristics shared with Ho- plophorionini and some Aconophorini and Membracini (Haviland 1925, Wood 1984, Dietrich and Deitz 1991). H. pruinosa fe- males also deposit a pale waxlike material in spirals along the twig and leaf edges in the vicinity of their egg masses (Haviland 1925; McKamey, unpubl.). Similar deposits are produced by Leioscyta spiralis Haviland (Haviland 1925, Striimpel 1986) (tribe Membracini), an additional (unidentified) Leioscyta species (McKamey, unpubl.), Ochropepla triangulum (Germar) (tribe Ho- plophorionini; Wood 1984 [as O. pallens Stal]), and Aconophora mexicana Stal (tribe Aconophorini; Wood 1984, Dietrich and Deitz 1991). Nymphs of H. pruinosa, L. spiralis, and A. mexicana are also clothed with irregular waxlike exudates that facili- tate crypsis among the spiral deposits (as in Striimpel 1986: Fig. 2). Many other species of Membracinae have nymphs with white waxlike exudates but lack waxy deposits ad- jacent to their egg masses. Such features have not been reported for species of subfamilies other than Membracinae. Havilandia hypsoproroides, NEw SPECIES (Figs. 1h—-k) Type locality: Fonteboa, Amazonas, Brazil [BMNH]. Description. — Head, pronotum, and ab- VOLUME 97, NUMBER 1 domen reddish brown, pronotal horn darker than rest of pronotum, legs yellow; head with vertex densely clothed with pale setae; tho- racic pleuron pilose, setae clothed with white waxlike material; forewing membrane uni- formly smoky hyaline. Pronotum (Fig. 11) produced into an elongate, slender antero- medial horn, apex in lateral view expanded and obliquely rounded, with | ventrolateral pair of strong carinae diverging from apex; posterior process approximately even with crossvein r-m of forewing at rest, apex slightly elevated. Forewing densely clothed with arcuate chaetoids. Metathoracic tibia (Fig. 1j) with setal row I bearing 15-18 cu- cullate setae. Dimensions (mm): Body length (head to apex of forewing at rest) 5.1, width across humeri 1.7; vertex height 0.7, width 1.0; pronotum length 5.3; head to horn apex 2.1; forewing length 4.5; prothoracic tibia length 1.0; metathoracic tibia length 1.5; ovipositor length 1.0. Material examined.— Holotype @ labeled: ‘‘Amazon./ Fonteboa; Riksmuseum/ Stock- holm; Dietrich Res./ 7-90-33a?; HOLO- TYPE/ Havilandia/ hypsoproroides/ Die- trich and McKamey” [BMNH]. Etymology.—The trivial name was se- lected to indicate the similarity of this spe- cies to members of the genus Hypsoprora Stal. Key TO SPECIES OF HAVILANDIA 1. Anterior pronotal process, in lateral view, short, broad, and tapered, apex rounded (Fig. la) ... Ste PRO RI OC OT eee pruinosa (Haviland) . Anterior pronotal process, in lateral view, elongate and slender, apex obliquely truncate (1g Ep iI) PA eS ee hypsoproroides, n. sp. _ Lewdeitzia, NEw GENUS (Figs. 2a—m) Type species: Lewdeitzia lunata, new species. Diagnosis. — Anterior region of pronotum in lateral view elevated into broad, rounded process with | carina on each side, in female further developed into large, marginally compressed horn; forewing with | or 2 r-m crossveins, with vein R initially divided into R, and Rs. Description.— Head: Vertex (Fig. 2g) not impressed mesad of ocelli, ventrolateral margins foliaceous and strongly produced; ocelli approximately 2x as far from each other as from mesal margins of eyes and approximately 2 ocellar diameters from dorsal margin of vertex; frontoclypeus flat, lateral corners rounded and apex truncate, margins foliaceous; rostrum extended to base of abdomen; frontoclypeus, in lateral view, forming shelf above clypellus; in an- terior view (Fig. 2g) with distance between lateral margins of eyes slightly less than dis- tance between humeri. Thorax: Pronotum with broad, marginally compressed anter- odorsal process, much larger in female (Fig. 2a) than in male (Fig. 2e), bearing pair of lateral carinae extending ventrolaterally from apex but not attaining posterolateral margins. Forewing (Figs. 2a, e) punctate in basal half between veins C and M and in base of anal area, vein R initially divided into R, and Rs, crossveins s and r-m, pres- ent, with crossvein r-m, present (Fig. 2a) or absent (Fig. 2e), 2 m-cu crossveins present, m-cu, perpendicular to veins M and Cu, Ist and 2nd M cells subequal in length, apical limbus relatively wide, contiguous with cla- vus for short distance only. Hind wing crossveins r-m and m-cu present. Legs: Pro- and mesothoracic tibiae (Figs. 2h, i) com- pressed and foliaceous; mesothoracic femur with anteroapical cucullate seta; metatho- racic tibia (Fig. 2c) straight, not compressed, row I with 7-8 enlarged cucullate setae, row II with 4-5 enlarged cucullate setae, row III without cucullate setae; metathoracic tarsus slightly longer than others, tarsomere I with 4 apical cucullate setae. Abdomen: Ster- num III without median tubercle, sternum IV with distinct transverse carina; terga without tuberosities or fenestrae. Male: Pygofer with vertical lateral carinae; lateral plate free, with prominent rounded vertical ridge over entire height (Fig. 2j); aedeagus 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Lae cise RAE ROTI says st AS Fig. 2. Lewdeitzia lunata, new genus, new species. a—d, holotype 2: a, head, pronotum, and forewing, lateral view (texture); b, head and pronotum, dorsal view (color); c, reversed right metathoracic tibia and tarsus, lateral view, same scale as 2h-i; d, second valvulae, lateral view; e—m, 6; e, head, pronotum, and forewing, reversed lateral view (texture); f-g, head and pronotum, dorsal (color) and anterior views, respectively; h-i, left pro- and mesothoracic tibiae and tarsi, lateral views; j, left lateral plate, dorsal view (ant, anterior); k, left style, ventrolateral view; I-m, aedeagus, anterior and lateral views, respectively. (Figs. 21, m) with shaft slender, in anterior view subparallel, in lateral view tapering apically, anterior face of apex evenly den- ticulate laterally; gonopore membrane, in posterior view, occupying nearly entire width of shaft; shank of style (Fig. 2k) with numerous long setae, apex recurved and acute; subgenital plate evenly tapering api- cally, without lobes. Female: Posterior margin of sternum VII with arcuate emar- gination; 2nd valvulae, in lateral view (Fig. 2d), subparallel, apical 4 irregularly cren- ulate. Integumental vestiture: Vertex and pronotum coarsely punctate, sparsely clothed with pale recumbent setae; thoracic pleuron sparsely setose, setae not clothed with waxlike material; forewing membrane glabrous. Abdominal terga coarsely punc- tate, pits without lateral setae; acanthae dentate. Notes and etymology. — The female of this genus resembles Aconophora Fairmaire (Aconophorini), while the male resembles Leioscyta. The genus is named to honor VOLUME 97, NUMBER 1 Lewis L. Deitz, whose higher classification and bibliographies have greatly facilitated research on the Membracoidea. Lewdeitzia lunata, NEw SPECIES (Figs. 2a—m) Type locality: Sao Paulo, Sao Paulo State, Brazil [NCSU]. Description.—Ground color brown; pronotal carinae, crescent-shaped band around posterior base of horn (female; Fig. 2b) or mottling between carinae and humeri (male; Fig. 2f), 2 transverse bands farther posterad on dorsum, transverse band on metopidium between humeri, and legs paler orange brown; forewing smoky hyaline with 2 pale transverse bands (1 near base, 1 aligned with end of clavus). Female pron- otal horn with apically bifid lateral longi- tudinal gibbosity, apex obtusely rounded, lateral carinae weak, terminating slightly posterad of humeri; male horn short and rounded, carinae distinct, marginal com- pressed area narrower. Other morphology as described for genus. Dimensions (mm).— Body length 2 6.3, 6 5.3; maximum width 2 2.8, ¢ 2.4; pronotal length 2 7.0, 6 4.5; head to horn apex @ 3.3; forewing length 2 5.5, ¢ 4.8; prothoracic tib- ia length 2 1.5, 6 1.2; metathoracic tibia length @ 2.2, 6 1.9; ovipositor length 1.9. Material examined.— Holotype 2 labeled: “Sao Paulo/ S. Paulo Brazil/ Nov. 1930; A.Maller,Coll./ FrankJohnson/ Donor; 1531 2/ S. Paulo/ S. Paulo/ 10.30; HOLOTYPE/ Lewdeitzia/ lunata/ Dietrich and Mc- Kamey” [NCSU]. Other material: ¢ from Corupa, Santa Catarina, Brazil, XI-1929, A. Maller, Coll. (Dietrich Research 15-91-40d 6) [NCSU]. Notes and etymology.—The holotype lacks its left metathoracic leg. The male, which lacks its left hind tarsus and right hind tarsomeres II and III, was excluded from type status because it may not be con- specific, given the considerable differences compared to the female. Nonetheless, be- cause extreme sexual dimorphism is com- mon in other species of Membracinae (e.g. see Sakakibara 1976 and Striimpel and Striimpel 1978), we consider the nearly identical color pattern and similar size of both Lewdeitzia specimens sufficient to re- gard them as a single species. The species name is derived from Latin for the pale cres- cent-shaped band around the base of the female’s horn. CLADISTICS Morphological characters Head 1. Frontoclypeus: 0, apex in lateral view not forming a shelf over clypellus, margins not expanded laterally (Fig. 1h); 1, apex in lateral view not forming a shelf over cly- pellus, margins expanded laterally; 2, apex in lateral view forming a shelf over clypellus (Sakakibara 1992: Figs. 1, 2, 4). Pronotum 2. Shape: 0, depressed or rounded (Figs. la, i); 1, compressed (Deitz 1975: Fig. 15S); 2, foliaceous (op. cit.: Fig. 15U). 3. Anteromedial horn: 0, absent or weak (Fig. la); 1, well developed (Fig. 11). Al- though entirely enclosed by the foliaceous pronotal margins, the horns of Phyllotropis and Folicarina are nonetheless well indi- cated (Sakakibara 1992: Figs. 3, 4). 4. Transverse metopidial carinae: 0, ab- sent; 1, restricted to humeri; 2, complete, two sides meeting medially. 5. Dorsolateral carinae: 0, absent; 1, one submedial pair (Figs. la, 1); 2, two or more pairs (Deitz 1975: Fig. 15R). The pronotal carinae of Hoplophorionini converge at the base of the metopidium rather than at its apex, as in other Membracinae, and there- fore may not be homologous. 6. Posterior extension of dorsolateral ca- rinae: 0, absent; 1, terminated above hu- meri; 2, extended beyond humeri. 7. Dorsal submedial carinae: 0, absent; 1, meeting medial carina anteriorly (Fig. 1a); 2, not meeting medial carina anteriorly. 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 8. Anteroventral pair of carinae, which are contiguous with dorsolateral carinae: 0, absent; 1, present (Figs. la, h, 1). 9. Irregular, oblique metopidial carinae: 0, absent; 1, present (Sakakibara 1992: Fig. 4). 10. Posterior transverse carina: 0, ab- sent; 1, present (Haviland 1925: Pl. II, Fig. 6). 11. Posterior sublateral longitudinal ca- rinae: 0, absent; 1, present (Fonseca and Di- ringshofen 1969: Fig. 3). Our treatment of this feature as a separate character from character 6 is supported by the presence of both dorsolateral and sublateral carinae in some taxa (e.g. Erechtia). 12. Punctation: 0, coarse—distance be- tween pits less than pit diameter, pits dis- tinct above humeri; 1, fine—distance be- tween pits greater than pit diameter, pits indistinct above humeri. Considerable vari- ation exists among taxa coded as state 0, but state | (e.g. Membracis) is distinctive (compare Figs. 7 and 10 of Wood and Mor- ris 1974). Pleuron and legs 13. Pleural wax: 0, absent; 1, present. State 1 is considered to represent a homo- log, although Havilandia has the pleuron densely pilose with waxy setae while Hyp- soprora has the pleuron sparsely setose with waxlike deposits on the surface of the scler- ite. 14. Pro- and mesothoracic tibiae: 0, nar- row, semicylindrical; 1, subfoliaceous (Figs. ZhSi): 15. Mesothoracic acute coxal process: 0, absent; 1, present. State 1 is a synapomor- phy of the tribe Hoplophorionin1. 16. Metathoracic tibia: 0, not clavate, row II with numerous cucullate setae (Fig. 2a); 1, distinctly clavate, row II with few or no cucullate setae (Figs. 1b, j). Some Erechtia and Leioscyta species have the metathoracic tibia weakly clavate, but with numerous cu- cullate setae in row II, perhaps representing an intermediate state of this character. 17. Length of metathoracic tarsi relative to pro- and mesothoracic tarsi: 0, longer; 1, subequal; 2, distinctly shorter (Deitz 1975: Fig. 14A). Forewing 18. Vein R: 0, initially divided into R, and Rs (Fig. la); 1, initially divided into Roso.5 and. Ray. (Deitz. 197 5:.Figl rAy: 19. Crossvein s: 0, absent; 1, present (Figs.2a;e). 20. Crossvein(s) r-m: 0, one (Figs. 1a, i); 1, two or more (Fig. 2a). 21. Membrane, erect macrotrichia: 0, absent; 1, present. 22. Membrane, arcuate chaetoids: 0, ab- sent, surface glabrous; 1, present, surface shagreen (Dietrich and Deitz 1993: Fig. 27). Abdomen 23. Sternum III: 0, unarmed; 1, with me- dial tubercle; 2, with transverse carina (Fig. la). 24. Sternum IV: 0, unarmed; 1, with transverse carina (Deitz 1975: Fig. 3A). 25. Dorsal tuberosities or fenestrae, terga V-VIII: 0, absent; 1, paired (op. cit.: Fig. 3A); 2, unpaired medial (Dietrich 1989: Fig. 18). The presence, in 7rinarea and some Erechtia, of paired tuberosities on the an- terior terga and unpaired medial tuberosi- ties on the posterior terga suggests that one feature was derived from fusion or separa- tion of the other. Second valvulae 26. Length: 0, elongate; 1, short and broad (Deitz 1975: Figs. 17H—-J). 27. Dorsal teeth (op. cit.: Figs. 17C, M): 0, absent; 1, one; 2, two. The serrations found in many membracid groups (op. cit.: Fig. 8) were not considered homologous. 28. Dorsoapical emargination: 0, absent; 1, present (Figs. Ic, k). Analysis Our data matrix (Table 1) included at least one representative of each genus of Mem- VOLUME 97, NUMBER 1 Table 1. represented by ?’s. 11 Character state matrix for numerical cladistic analysis of the Membracinae. Missing values are Character OTU 1-5 6-10 11-15 16-20 21-28 Heteronotinae (outgroup) Dysyncritus intectus Fowler 20000 00000 00000 00010 00000000 Membracinae Aconophorini (Guayaquila Goding) 00100 00000 00010 01110 01011000 Hypsoprorini (Hypsoprora Stal) 20101 11100 00110 00110 01010020 Hoplophorionini (Potnia Stal) 00102 22000 10001 12110 00012000 Membracina Bolbonota sp. 10002 22001 00010 01010 01010100 Enchenopa binotata (Say) 21101 21010 00010 00010 01011020 Enchophyllum sp. A 21101 11010 01010 00010 01011020 Enchophyllum sp. B 21101 11000 01010 00010 01011020 Erechtia bicolor Walker 20022 21000 10010 01011 01112000 Erechtia sp. 20022 21000 10010 01010 01112770 Eunusa concolor Fonseca 00000 00000 00000 00010 00110000 Havilandia pruinosa (Haviland) 00001 21100 00110 11000 11100001 H. hypsoproroides gen. & sp. n. 00101 21100 00110 11000 11000001 Kronides sp. 00101 11010 00010 00010 01010020 Enchenopa beebei (Haviland) 21001 21010 00010 00010 01011020 Leioscyta sp. A 10001 21000 00010 00010 00010010 Leioscyta sp. B 20002 21000 10010 00010 01112000 Lewdeitzia lunata 4 gen. & sp n. 20101 21000 00010 00010 01010000 Lewdeitzia lunata 2 20101 21000 00010 00011 01010000 ‘“*Membracis” ferruginea (Funkh.) 21000 00000 00010 00010 11110000 Membracis foliata (L.) 22000 00000 01010 00010 01011020 Membracis carinulata Richter 21001 11010 01010 00010 01011000 Phyllotropis cingulata (Germar) 22101 11000 01010 00010 01011020 Folicarina bicolor Sakakibara 22101 11010 01010 00010 01010000 Paragara nigra (Funkhouser) 00001 22000 00010 00010 00110100 Tritropidia sp. ALOT 21001 00010 00010 00110100 Tylopelta gibbera (Stal) 20002 21011 00010 00010 01010000 Talipedini Trinarea appendiculata (Fonseca) 20112 21000 10010 12011 01112010 Trinarea sp. A 20022 21000 10010 12010 01112??? bracini except Bolbonotodes, which is known only from the 2 holotype, and Campylen- chia, which is identical to Enchenopa bin- otata for the included characters. Because Lewdeitzia is dimorphic for character 18, we included the male and female in the data as separate OTUs. The tribe Talipedini was represented by two species. The other mem- bracine tribes, which are invariant for most of the characters, were each represented by one OTU. A species belonging to the sister group of Membracinae (Dietrich and Deitz 1993), Dysyncritus intectus Fowler (type species of the genus; subfamily Heteronotinae) was chosen as the outgroup. Although clearly a member of Heteronotinae by virtue of the single r-m crossvein in its forewing, D. in- tectus shares certain features with Membra- cinae that are absent in other Heteronotin- ae. The metathoracic tibiae have cucullate setae enlarged in rows I and II but reduced 12 Table 2. List of apomorphies for the cladogram (Fig. 3) based on ACCTRAN character state optimi- zation (Swofford 1990); other equally parsimonious optimizations are possible. Terminal taxa without apo- morphies in the data are not listed. Nonhomoplasious changes are indicated by *. Node Apomorphies 52 14 (1), 22 (1) 51 5)(2) 3612); 7 @) 50 LRG) 25K(O— 2) 49 4 (2), 17 (1) 47 16 (1), 17 (2) 46 3 (1), 4 (0) 45 23 (0) 44 10 (1) 43 > ()) 42 1 (1), 22 (0) 4] 23 (1), 26 (1) 40 3 (1) 39 6 (1), 27 (2) 38 9 (1) 37 Di) 2510) 36 N72 GO} 35 27 (0) 34 9 (0) 33 2 (2) 32 6 (2) 31 1 (0), 17 (1) 30 8 (1), 13 (1), 16 (1), 19 (0)*, 21 (1), 24 (0), 28 (1)* Heteronotinae 23 (0), 24 (0) Aconophorini 5 (0), 6 (0), 7 (0), 18 (1), 25 (1) Havilandia pruinosa 3 (0), 23 (1) Hypsoprorini 8 (1), 13 (1), 18 (1) Enchenopa beebei 3 (0) Membracis foliata 3 (0), 5 (0), 6 (0), 7 (0) Membracis carinulata 3 (0) Folicarina 2 (2), 25 (0) Kronides 1 (0) Lewdeitzia ° 20 (1) Paragara 1 (0), 7 (2) Tritropidia PAOD SOW S HOO) Leioscyta sp. A 27 (1) Bolbonota WG) (2) a iF (226 (1) Tylopelta 9 (1) Hoplophorionini 1°(0); 7 (2), 14°), 15 (1)*} 185(1); 22°), 23 (0) 4 (1), 20 (1), 27 (1) 20 (1) PY A PALEY) 1 (QO) Trinarea appendiculata Erechtia bicolor Membracis foliata Eunusa 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in row III, a condition heretofore unknown outside of Membracinae, and forewing vein R is initially divided into R, and Rs. We analyzed the data (Table 1) using Hennig86 version 1.5 (Farris 1988) and PAUP version 3.0s (Swofford 1990). Exact algorithms (guaranteed of finding minimal length trees) proved too time-consuming, so we used the command sequence “mh; bb*”’ of Hennig86 and the heuristic search with Tree-Bisection-Reconnection branch swap- ping (TBR) and MULPARS of PAUP. Among the 10 multistate characters, only characters 2 and 25 were arranged a priori in transformation series; thus only these 2 characters were treated as ordered (addi- tive). Initially, we assigned all characters weight = 1. To find trees supported by the most consistent characters, we used the suc- cessive weighting facility of Hennig86 (command sequence “‘xs w; mh; bb*;’’). We compared the lengths of trees resulting from weighted and unweighted analyses by re- setting the weight of each character to 1 (command sequence “‘ccode /1.; xsteps 1;”’). RESULTS AND DISCUSSION Hennig86 and PAUP yielded identical re- sults for the unweighted analyses. Both found 4 equally most parsimonious trees of length = 88, consistency index (excluding autapomorphies) = 0.402, and retention in- dex = 0.651. One of four most parsimoni- ous trees is presented in Fig. 3. The others differ in whether the Erechtia species form a clade together or with 7rinarea sp. A, and whether node 49 is supported. None of these 4 trees was preferred by successive weight- ing, which stabilized after 2 iterations at 71 trees of lengths 90-92, based on all character weights = |. Component 47 (Fig. 3) confirms Deitz’s (1975) hypothesis that the closest relative of Hoplophorionini is Talipedini, which is a para- or polyphyletic group. Both tribes are apparently derived from Membracini, which is also apparently ancestral to Hyp- soprorini and Aconophorini. VOLUME 97, NUMBER 1 39 37 40 36 38 43 53 45 51 49 52 50 31 42 47 30 32 35 41 46 Heteronotinae (Dysyncritus) Aconophorini Havilandia pruinosa Havilandia hypsoproroides Hypsopronini Enchenopa binotata Enchenopa beebei Enchophyllum sp. A Enchophyllum sp. B Membracis foliata Phyllotropis Membracis carinulata Folicarina Kronides Lewdeitzia & Lewadeitzia 9 Leioscyta sp. A Paragara Tritropidia Bolbonota Tylopelta Hoplophorionini Trinarea appendiculata } Talipedini Trinarea sp. A Erechtia bicolor Erechtia sp. A Leioscyta sp. B Membracis ferruginea Eunusa Fig. 3. Hypothesized cladistic relationships among Membracinae. Apomorphies are listed in Table 2. 14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Aconophorini, Hypsoprorini, and Ho- plophorionini are diverse monophyletic groups with distinct morphologies and be- haviors, so recognition of additional tribes for genera currently placed in Membracini seems preferable to treating all the tribes of Membracinae as synonyms. However, be- cause some genera of Membracini (Mem- bracis and Leioscyta) are polyphyletic and others are defined by individual features of the pronotum or forewing (Funkhouser 1951), the creation of new tribes should await species-level analyses that clarify the limits of genera within the two complexes. One problematic complex of genera in- cludes Folicarina, Phyllotropis, Membracis, and Enchophyllum, which, apart from the plesiomorphic horn in the latter genus, are distinguished from each other only by com- binations of four pronotal features: (1) me- dian carina extensively compressed into a plate-like carina or not; (2) metopidium with ventral lobe or not; (3) dorsolateral carinae present or not; and (4) oblique metopidial carinae present or not. Membracis exhibits the greatest variation, including among its species both states of all four characters (as examples of [1] and [2], M. foliata vs. M. mexicana Stal; of [3], M. flava Richter vs. M. mexicana; of [4], M. carinulata vs. M. humilis Fowler). The evolutionary polari- ties of these features need to be determined to establish better generic limits in this com- plex. The stability of the current genera is especially precarious because none is de- fined by a single feature—only by combi- nations. The other problematic complex includes Leioscyta and its relatives (key couplets 14— 20, and Havilandia). Leioscyta includes species with and without dorsolateral cari- nae and foliaceous mesothoracic tibia, and differs from the other genera only by the absence of their diagnostic (and seemingly apomorphic) traits, rather than by any syn- apomorphies of its own. Even the outgroup, Dysyncritus intectus, would fit the present concept of Leioscyta were it not for the fea- tures distinguishing their respective sub- families. Thus, Leioscyta probably com- prises a diverse and polyphyletic set of species retaining different ancestral features, and is in great need of further work. Our preliminary estimate provides a framework for such studies. Although some problems remain to be solved, other nomenclatural changes are al- ready due. The Leioscyta-complex differs from the Membracis-complex in having coarser pronotal punctation. This feature was disregarded by Richter (1947) when he described the new species Membracis mi- caniaae and by Fonseca and Diringshofen (1969) when they referred L. ferruginea Funkhouser to Membracis. Both species should be considered Membracini incertae sedis until generic limits in the Leioscyta- complex are better resolved. The Leioscyta- complex differs from Enchenopa in lacking oblique metopidial carinae, but not in lack- ing an anterior horn (e.g. males of E. per- mutata Van Duzee are hornless). We there- fore refer L. beebei, whose holotype (BMNH) has oblique metopidial carinae but lacks a horn, to Enchenopa, creating the new com- bination EF. beebei (Haviland). Our analyses support recognition of the new genera Havilandia and Lewdeitzia. The most parsimonious trees indicate that Hav- ilandia is the sister group of Aconophorini, but the genus shares at least one derived feature with some Hypsoprorini (character 8: pronotal horn with a pair of anteroventral carinae) and trees placing Havilandia as the sister group of Hypsoprorini required only one additional step. Although we included no characters in the analysis to unite the male and female of Lewdeitzia, the genus clearly represents a lineage distinct from other Membracini and therefore merits for- mal recognition. Our provisional placement of Havilandia and Lewdeitzia in Membra- cini reflects Deitz’s (1975) concept of the tribe, which, for nomenclatural stability, is retained at present. VOLUME 97, NUMBER 1 ACKNOWLEDGMENTS We are indebted to R. L. Blinn (NCSU), Mick Webb, and Peter Broomfield (BMNH) for lending specimens. L. L. Deitz, R. W. Hodges, P. M. Marsh, H. H. Neunzig, T. K. Wood, and two anonymous reviewers crit- ically reviewed earlier versions of the manu- script. This research was supported, in part, by the North Carolina Agricultural Re- search Service, North Carolina State Uni- versity, Raleigh, the University of Con- necticut Department of Ecology and Evolutionary Biology, and a National Sci- ence Foundation Graduate Fellowship (to SHM). LITERATURE CITED Deitz, L. L. 1975. Classification of the higher cate- gories of the New World treehoppers (Homoptera: Membracidae). North Carolina Agricultural Ex- periment Station Technical Bulletin 225: 1-177. Deitz, L. L. and C. H. Dietrich. 1993. Superfamily Membracoidea (Homoptera: Auchenorrhyncha): I. Introduction and revised classification with new family-group taxa. Systematic Entomology 18: 287-296. Dietrich, C. H. 1989. Surface sculpturing of the ab- dominal integument of Membracidae and other Auchenorrhyncha (Homoptera). Proceedings of the Entomological Society of Washington 91: 143-152. Dietrich, C. H. and L. L. Deitz. 1991. Revision of the Neotropical treehopper tribe Aconophorini (Homoptera: Membracidae). North Carolina Ag- ricultural Research Service Technical Bulletin 293: 1-134. 1993. Superfamily Membracoidea (Homop- tera: Auchenorrhycha). IT. Cladistic analysis and conclusions. Systematic Entomology 18: 297-311. Farris, J.S. 1988. HENNIG86 reference, version 1.5. Computer program and documentation, pub- lished privately, not paginated. Fonseca, J. P. da, and R. von Diringshofen. 1969. Contribuicao ao conhecimento dos membracideos neotropicos (Homoptera: Membracidae, VI). Ar- quivos do Instituto Biologico, Sao Paulo 36: 143- 161. Funkhouser, W. D. 1940. New Peruvian Membra- cidae (Homoptera). Journal of the New York En- tomological Society 48: 275-292. 1951. Homoptera fam. Membracidae. Gen- era Insectorum 208: 1-383. Goding, F. W. 1926a. New Membracidae, I. Journal 15 of the New York Entomological Society 34: 243- 246. . 1926b. Classification of the Membracidae of America. Journal of the New York Entomological Society 34: 295-317. 1929. The Membracidae of South America and the Antilles. 1V. Subfamilies Hoplophorion- inae, Darninae, Smiliinae, Tragopinae (Homop- tera). Transactions of the Entomological Society of America 55: 197-330. Haviland, M. D. 1925. The Membracidae of Kar- tabo, Bartica District, British Guiana, with de- scriptions of new species and bionomical notes. Zoologica [New York] 6: 229-290. Loye, J. E. 1992. Ecological diversity and host plant relationships of treehoppers in a lowland tropical rainforest (Homoptera: Membracidae and Nicom- iidae), pp. 280-289. Jn Quintero, D. and A. Aiello, eds., Insects of Panama and Mesoamerica: Se- lected Studies. Oxford University Press, Oxford. McKamey, S. H. 1992. Reappraisal of the Neotrop- ical treehopper genus Eunusa Fonseca (Homop- tera: Membracidae), with ecological notes. Annals of the Entomological Society of America 85: 253- D511. McKamey, S. H. and L. L. Deitz. 1991. Nomencla- tural changes in the treehopper tribes Hoplopho- rionini, Smiliini, and Talipedini (Homoptera: Membracidae). Proceedings of the Entomological Society of Washington 93: 193-196. Richter, L. 1947. Membracidae Colombianae. Re- vision de las especies Colombianas del género Membracis. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 7: 382- 403. Sakakibara, A.M. 1976. Sphongophorus gracilis Sak- akibara, 1971 —notas complentares. (Homoptera- Membracidae). Atas da Sociedade de Biologia do Rio de Janeiro 18: 1-2. 1992. Sobre alguns Membracini (Homop- tera, Membracidae): Notas tax6nomicas e descri- ces de género e espécies novos. Revista brasileira Entomologica 36: 93-100. Strimpel, H. 1986. Bemerkungen zur protektiven Mimese von Larven der neotropischen Membra- cide Leioscyta spiralis Haviland, 1925 (Homop- tera, Membracidae). Entomologische Mitteilun- gen aus dem Zoologischen Museum Hamburg 8: 281-284. Strimpel, H. and R. Striimpel. 1978. Die Membra- ciden-Fauna Kolumbiens 4. Die Gattung Tvritro- pidia Stal, 1869. Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 6: 133-149. Swofford, D. L. 1990. PAUP, Phylogenetic Analysis Using Parsimony, Version 3.0. Computer pro- gram distributed by the Illinois Natural History Survey, Champaign, Illinois. 16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Wood, T. K. 1984. Life history patterns of tropical membracids (Homoptera: Membracidae). Socio- biology 8: 299-344. 1993a. Diversity in the New World Mem- bracidae. Annual Review of Entomology 38: 409- 435. _ 1993b. Speciation of the Enchenopa binotata complex (Insecta: Homoptera: Membracidae), pp. 299-317. In Lees, D. R. and D. Edwards, eds., Evolutionary Patterns and Processes. Academic Press, London. Wood, T. K. and G. K. Morris. 1974. Studies on the function of the membracid pronotum (Homop- tera). I. Occurrence and distribution of articulated hairs. Canadian Entomologist 106: 143-158. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 17-21 A NEW SPECIES OF APHELINUS (HYMENOPTERA: APHELINIDAE) THAT PARASITIZES THE SPIREA APHID, APHIS SPIRAECOLA PATCH (HOMOPTERA: APHIDIDAE) GreGorY A. EvANs, MICHAEL E. SCHAUFF, MOH LENG KOK-YOKOMI, AND RAYMOND K. YOKOMI (GAE) University of Florida, Entomology and Nematology Department, Gainesville, Florida 32611; (MES) Systematic Entomology Laboratory, USDA-ARS, PSI, % USS. National Museum of Natural History, NHB 168, Washington, D.C. 20560; (MLK-Y) University of Florida, IFAS, Central Florida Research and Education Center, Leesburg, Florida 34748; (RK Y) USDA, ARS, Horticultural Research Laboratory, 2120 Camden Road, Orlando, Florida 32803. Abstract. — A new species of an aphid parasitoid, Aphelinus spiraecolae Evans and Schauff (Hymenoptera: Aphelinidae) is described and figured. This species is parasitic on the spirea aphid, Aphis spiraecola Patch, and is being investigated for possible use in a bio- logical control program against the spirea aphid and other citrus aphids including the brown citrus aphid, 7oxoptera citricida (Kirkaldy). The new species is very similar to Aphelinus gossypii and characters to differentiate it from related species are given. Key Words: During 1992, collections of aphid para- sitoids from citrus were made in four prov- inces of the People’s Republic of China namely, Sichuan, Hunan, Fujian, and Guangdong. Some of the emerging parasit- oids collected frem the Guangzhou area (Guangdong Province) were brought to Florida for study as potential biological con- trol agents of citrus aphids. This research was undertaken because the brown citrus aphid, Toxoptera citricida (Kirkaldy), the most efficient vector of citrus tristeza virus has spread throughout the Ca- ribbean Basin except the Bahamas (Yokomi et al. 1994) and is likely to be introduced into the continental U.S. in the foreseeable future. This aphid generally attacks only cit- rus and citrus relatives and is thought to be native to Asia where citrus originated. Be- cause the aphid has potential to cause major damage to citrus, research is under way to Parasitoid, biological control, citrus, citrus tristeza virus find control measures that would suppress the aphid population and mitigate the dam- age. The species described in this paper was originally believed to be Aphelinus gossypii Timberlake, a common, widespread para- site of the melon or cotton aphid, Aphis gossypii Glover, and related aphids. How- ever, investigation of the biology of the spe- cies from China revealed differences be- tween it and that of A. gossypii including its ability to attack the spirea aphid, Aphis spi- raecola Patch (Yokomiet al. 1993). Further examination revealed characters that were different from those of A. gossypii and we are, therefore, proposing a new species name for this taxon. The importance of this new species is due to its potential as a biological control agent of the spirea aphid which has few effective parasitoids in the United States (Cole 1925, 18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-7. Aphelinus spiraecolae, female except Figs. 6, 7: 1, fore wing, disk setae excluded and stigmal vein magnified. 2, thorax. 3, antenna. 4, gaster (venter) and hind leg. 5, mandible. 6, antenna. 7, aedeagus. VOLUME 97, NUMBER 1 Miller 1929, Tang et al. 1994). The spirea aphid is abundant on citrus worldwide and has a wide host range. It transmits citrus tristeza virus (Yokomi and Garnsey 1987), as well as some potyviruses (Adlerz 1987). Aphelinus spiraecolae Evans and Schauff, New SpPEcIES Figs. 1-7 Diagnosis. — Head and thorax dark brown; legs yellow except dark brown coxae and hind tibiae; gaster dark brown with yellow- ish base; Fl and F2 broader than long, F3 quadrate; costal cell of fore wing with two lines of ventral setae, area proximal to linea calva bordered by one complete row of 13- 16 setae and one incomplete row of 2-5 setae. Female: Length, 0.95-1.2 mm (Holo- type, 0.98 mm). Color: Head and thorax dark brown, legs yellow except dark brown coxae and hind tibiae; gaster dark brown with base (terga I) yellowish; endophragma fuscous; third valvulae pale with dark brown lateral margins; antennae light brown; wings hyaline. Structure: Head dorsum about as broad as thorax; mandibles bidentate (Fig. 5) with an internal tooth, a blunt middle tooth and a truncation; lateral ocellus sep- arated from eye margin by one ocellus di- ameter; occiput reticulate. Antenna (Fig. 3) with short radicle; scape about 4 to 5.5 times as long as broad; pedicel (ventral length) less than 2 times its width; Fl annuliform, with ventral margin longer than dorsal margin; F2 broader than long, as long as F1; F3 quadrate, about 2 times longer than F2 with one linear sensillum; club 2.4 times longer than broad, about 3 times longer than F3 with 6-7 linear sensillae and 7-8 papillae. Thorax (Fig. 2) with fine reticulations; mid- lobe of mesoscutum with two pairs of pri- mary setae and approximately 40 setae, pri- mary setae MC (posterior central) short, as long as ML (anterior lateral) setae, and ax- illar setae (AS), reaching base of SC1 setae; each side lobe with 2-3 setae; scutellum with 19 2 pairs of black setae, placoid sensillae small and widely separated, with pale, diamond- shape markings around each; endophragma short, about 0.7 times as long as the length of the thorax; tibial spur of middle leg slight- ly shorter than corresponding basitarsus; hind tibia with 7 conspicuous conical setae at apex, tibial spur of leg III about one-half as long as corresponding basitarsus. Fore wing (Fig. 1) broad, more than 2 times as long as wide (0.8:0.36 mm); costal cell 1.2 times longer than marginal vein and with 20-25 ventral setae in 2 rows and 11-13 dorsal setae; submarginal vein with 5-6 se- tae; marginal vein with 9 setae along the margin, all about 1.5 times longer than width of marginal vein; basal cell bare; stigmal vein short with stigma rounded; area prox- imal to linea calva bordered by one com- plete row of 1 1-15 setae and one incomplete row of 1-5 setae; marginal fringe very short, less than 0.05 times as wide as fore wing disc; ciliation dense after the linea calva. Gaster (Fig. 4) longer than thorax (0.56:0.35 mm); ovipositor (0.35 mm) inserted at mid- dle of gaster, only slightly exerted distally, longer than hind tibia (0.29 mm) and mid tibia (0.21 mm); third valvulae (0.12 mm) one-third the length of entire ovipositor. Male: Length (0.88-0.92 mm): Similar to female except central portion of femur II and tibia II slightly infuscate; radicle, scape and pedicel brown (Fig. 6); scape with 3 round sensoria and club with 2 linear sen- sillae; abdomen tapering distally; aedeagus as shown in Fig. 7. Material examined.—Holotype female, Fushan City, Guangdong Province, P.R. China, VII 1992, ex Aphis spiraecola, R. Yokomi, deposited in the Institute of Zo- ology, Academy of Sinica, Being, P.R. China. Paratypes (19 29, 11 46) with same data as holotype. Paratypes deposited in: U.S. National Museum of Natural History, Washington, D.C.; Florida State Collection of Arthropods, Gainesville, Florida; The Natural History Museum, London, En- 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gland; Canadian National Collection, Ot- tawa and the Collection of M. Hayat, Ali- garh, India. Known distribution.— Guangdong Prov- ince, P.R. China. Comments.—Aphelinus spiraecolae is placed in the Aphelinus subgenus of the ge- nus Aphelinus as defined by Hayat (1990). This species is similar in body color and fore wing ciliation to species placed by Ze- havi and Rosen (1988) in the Aphelinus mali group (Aphelinus campestris Jasnosh, A. gossypii Timberlake, A. mali (Haldeman), A. paramali Zehavi & Rosen, and A. pro- ciphili Carver). However, it differs from the species in this group by the color of the legs. The femora of the middle legs of all of the mali group species are dark brown. The legs of the A. spiraecolae female are entirely yel- low except for its dark brown coxae and hind tibia. While differences in body color may be useful in discriminating different Aphelinus species, these differences alone may not always provide conclusive evi- dence to distinguish different species. Jans- sen (1961) and Michel (1969) reported con- siderable intraspecific variation in the color of the body (and legs) of Aphelinus asychis Walker (= semiflavus (Howard)) and A. chaonia Walker, respectively. Given this variation and the fact that species groups are not well justified nor widely used in this genus, we are not assigning 4. spiraecolae to a species group at this time. Of the species mentioned above, Aphe- linus spiraecolae is most similar to A. gos- sypil (Timberlake) in coloration, shape and ciliation of the fore wing, and antennal structure. However, in A. gossypii the fem- ora and tibiae of the fore and middle legs and basitarsi of the hind legs of females are dark brown. The MC setae are elongate, reaching the placoid sensillae (shorter and not reaching placoids in spiraecolae); and the setae along the marginal vein are about 2 times as long as the width of the marginal vein (setae 1.5 times longer than width of marginal vein in spiraecolae). Aphelinus spiraecolae is also similar to Aphelinus chaonia Walker. This species has a wide distribution and host range. It was introduced into California by Flanders and Fisher (1959) from South China for the con- trol of the black citrus aphid, Toxoptera au- rantil, and propagated in the laboratory on the spirea aphid. These two species can usu- ally be easily distinguished from each other by the color of the legs. Normally, the fem- ora of the fore and middle legs of A. chaonia are dark brown. However, the legs are yel- lowish in the light form of 4. chaonia. Nev- ertheless, A. chaonia can be distinguished by the greater number of setae (more than 20) along the proximal border of the linea calva of the fore wing (less than 20 setae in spiraecolae) and the dark base of the gaster (base of gaster yellow in spiraecolae). ACKNOWLEDGMENTS We thank Q. Tang (USDA, ARS, Hor- ticultural Research Laboratory, Orlando, Florida; Visiting Scientist from the Biolog- ical Control Research Institute, Fujian Ag- ricultural College, Fuzhou, Fujian Province, 350002, P.R. China) for technical assistance and who first noted these specific morpho- logical characters during his studies on the biology of this species. We are indebted to Dr. Limhout Nong (Florida Dept. Agric. and Cons. Serv., Div. Plant Ind., Gainesville, Florida) for maintaining the parasitoid col- ony in acontainment facility. We gratefully acknowledge the Ministry of Agriculture, Beying, P.R. China, for kind support and local arrangements that allowed us to con- duct the survey in China for natural enemies of citrus aphids; the Florida Citrus Produc- tion Research Council for its financial sup- port and the Office of International Coop- eration and Development, Research and Scientific Exchange Division, USDA, Washington, D.C. for kind support and co- operation. We thank Drs. M. Hayat (Ali- garh Muslim University), and H. Browning (University of Florida, Citrus Research and Education Center, Lake Alfred, Florida) for VOLUME 97, NUMBER 1 their review of and comments regarding this manuscript. LITERATURE CITED Adlerz, W.C. 1987. Cucurbit potyvirus transmission by alate aphids (Homoptera: Aphididae) trapped alive. Journal of Economic Entomology 80: 87- 92. Cole, F. R. 1925. The natural enemies of the citrus aphids, Aphis spiraecola (Patch). Journal of Eco- nomic Entomology 18: 219-223. Flanders, S. E.and T. W. Fisher. 1959. The economic effect of aphidophagous insects on citrus in South China. Journal of Economic Entomology 52(3): 536-537. Hayat, M. 1990. Taxonomic studies on Aphelinus (Hymenoptera: Aphelinidae) 2. A new subgenus from India, with comments on Mesidia and Mes- idiopsis. Oriental Insects 24: 253-257. Janssen, M. 1961. Pigmentmodifikation und neuer Fundort von Aphelinus semiflavus How. Beitrage zur Entomologie, Band 11: 671-678. Michel, M. 1969. Contribution a l’etude des Aphel- inidae aphidiphages et de leurs hotes en France (Hym. Chalcidoidea). Entomophaga 14(4): 439- 446. Miller, R. L. 1929. A contribution to the biology and control of the green citrus aphid, Aphis spiraecola 21 Patch. Florida Agricultural Experiment Station Bulletin 203: 431476. Tang, Y. Q., R. K. Yokomi, and R. J. Gagné. 1994. Life history and description of Endaphis maculans (Diptera: Cecidomyiidae), an endoparasitoid of aphids in Florida and the Caribbean Basin. Annals of the Entomological Society of America: In press. (Accepted May 5, 1994). Yokomi, R. K. and S. M. Garnsey. 1987. Transmis- sion of citrus tristeza virus Aphis gossypii and Aphis citricola in Florida. Phytophylactica 19: 169-172. Yokomi, R. K., R. Lastra, M. B. Stoetzel, V. D. Dam- steegt, R. F. Lee, S. M. Garnsey, T. R. Gottwald, M. A. Rocha-Pena, and C. L. Niblett. 1994. Es- tablishment of the brown citrus aphid (Homop- tera: Aphididae) in Central America and the Ca- ribbean Basin and transmission of citrus tristeza virus. Journal of Economic Entomology 87: 1078- 1085. Yokomi, R. K., Y. Q. Tang, L. Nong, and M. L. Kok- Yokomi. 1993. Potential mitigation of the threat of the brown citrus aphid, Toxoptera citricida (Kir- kaldy), by integrated pest management. Proceed- ings of the Florida State Horticultural Society 106: 81-85. Zehavi, A. and D. Rosen. 1988. A new species of Aphelinus (Hymenoptera: Aphelinidae) from Is- rael, with notes on the mali group. Israel Journal of Entomology 22: 101-108. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 22-34 A REVIEW OF THE MYDID GENUS PSEUDONOMONEURA BEQUAERT (DIPTERA: MYDIDAE), WITH THE DESCRIPTION OF TWO NEW SPECIES Scott J. FirZGERALD AND Boris C. KONDRATIEFF Colorado State University, Department of Entomology, Fort Collins, Colorado 80523. Abstract.—The mydid genus Pseudonomoneura, with six species is reviewed: P. cali- fornica (Hardy), P. bajaensis, n. sp., P. hirta (Coquillett), P. micheneri (James), P. nelsoni, n. sp. and P. tinkhami (Hardy). A lectotype is designated for Leptomydas concinnus Coquillett, a previously recognized junior subjective synonym of P. hirta. The male and female genitalia of all species are illustrated and a key to the species is presented. Key Words: The genus Pseudonomoneura was pro- posed by Bequaert (1961) for a small group of Nearctic species of flies originally placed in the genus Nomoneura (Hardy 1950). In 1971 Wilcox and Papavero provided the first adequate generic description of Pseu- donomoneura in their review of the Amer- ican genera of Mydidae. The genus Pseu- donomoneura is most similar and appears related to the Nearctic genus Nemomydas, and species of both genera are sympatric. Males of Pseudonomoneura are easily sep- arated from males of Nemomydas by the ventral process of the gonocoxite either pro- duced singly (P. hirta (Coquillett), P. mich- eneri (James)), reduced (P. californica (Har- dy)), or absent (P. bajaensis, n. sp., P. nelsoni, n. sp.). In all species of Nemomydas, the gonocoxites have a dorsal and ventral dig- itate process (Kondratieff and Welch 1990). Additionally, the apex of the aedeagus is an erect slender tube or tongue in Nemomydas, and a thick ventral bulb in Pseudonomo- neura. Females of Pseudonomoneura may be separated from Nemomydas by the an- tennal club being shorter than or subequal in length to the basal flagellomere (Wilcox 1981). In their key to the American genera Diptera, Mydidae, Pseudonomoneura, mydid flies, Nearctic of Mydidae, Artigas and Papavero (1990) added that females of Pseudonomoneura have the “spermathecae with 3 capsules,” and Nemomydas with “‘two capsules.” The number of capsules actually range from two to three in Pseudonomoneura. Hardy (1950) previously characterized Pseudonomoneura as having the proboscis ““developed well beyond the oral margin.” However, both P. bajaensis, n. sp. and P. nelsoni, n. sp. have vestigial mouthparts. Wilcox and Papavero (1971) indicated that the geographic range of Pseudonomo- neura included Utah and northern Mexico states of Sonora, Chihuahua and Coahuila. No specimens from these areas were avail- able for examination. Morphology and terminology follows Wilcox (1981). Abbreviations for deposi- tories of specimens are: California Academy of Sciences (CAS); Arizona State University (ASU); C. P. Gillette Museum of Arthropod Diversity, Colorado State University (CSU); Florida State Collection of Arthropods (FSCA); Los Angeles County Museum of Natural History (LACMNH); University of Arizona (UA); University of California, Da- vis (UCD); National Museum of Natural VOLUME 97, NUMBER 1 History, Smithsonian Institution (USNM); Utah State University (USU); Washington State University (WSU); W. F. Barr Ento- mological Collection, University of Idaho (WFBM). KEY TO THE SPECIES OF PSEUDONOMONEURA (female unknown for P. bajaensis, new species) 1. Mouthparts vestigial Mouthparts well-developed 2. Male terminalia with upper forceps of epan- drium simple (Figs. 20, 21). Female with ab- domen globose, wider than thorax (Fig. 24); terminalia as Fig. 23 .... P. nelsoni, new species Male terminalia with upper forceps of epan- drium bifurcate (Figs. 17 and 18) ss A pee Rites SEE ONE Sa P. bajaensis, new species 3. Proboscis long, extending to second flagello- mere (club) Proboscis short, extending only to or middle Omirst hagellomenrcia ean ea P. tinkhami 4. Male terminalia with upper forceps of epan- drium with subapical digitiform inward pro- jecting process (Figs. 1, 2). Male and female antennal flagellomeres black. Female termin- mltatas ies ea) Fahne Oe P. californica Male terminalia with upper forceps of epan- drium without a subapical digitiform process (Figs. 6, 10). Second antennal flagellomere tinged with red or yellow. Female terminalia ASHRICSMSL ANG lOlea ence es ery re ook 5 5. Male with upper inner flange of upper forceps of epandrium weakly developed in lateral view, apex narrowly rounded (Fig. 5). Female ter- minalia as Fig. 8. Abdominal setae usually dense yellow,-sometimes pray”... 4......-5--- = P. hirta Male with upper inner flange of upper forceps of epandrium strongly developed in lateral view, apex bluntly truncate (Fig. 9). Female termin- alia as in Fig. 12. Abdominal setae usually white ET ea eo ce P. micheneri Pseudonomoneura californica (Hardy) Figs. 1-4 Nomoneura californica Hardy 1950: 11. Holotype male (CAS), USA: California: Riverside County, Blythe, 15 July 1947, J. M. MacSwain, 7amarix; examined. Diagnosis.—The red tergites will distin- guish both sexes from all other species, ex- cept red variants of P. micheneri (James). 23 In the male of P. californica, the upper for- ceps of the epandrium have a subapical dig- itiform process which projects inward in dorsal view (Fig. 2), and posteriorly in lat- eral view (Fig. 1), whereas the upper forceps of P. micheneri lack this process (Figs. 9, 10). Females of P. californica can be sepa- rated from females of P. micheneri by the distinctive shape of the furca, and the more heavily sclerotized edge of tergite ten (Fig. 4). Additionally, the antennae of P. califor- nica are entirely black, whereas the second flagellomere of P. micheneri are usually or- ange-yellow. Discussion. — Hardy (1950) indicated that P. californica was related to P. micheneri. However, P. californica appears to be much more similar to P. bajaensis, n. sp. (Figs. 17-19), and P. nelsoni, n. sp. (Figs. 20-22). All three species lack the flanges of the upper forceps of the epandrium and the ventral processes of the gonocoxites are reduced or absent. Distribution. — Arizona and California to Baja California, Mexico. Specimens examined.— USA: Arizona: La Paz Co: Ehrenberg. 216. June 1938, F. H. Parker, 1 female (USNM); Maricopa Co.: 1.6 mi. E. Barnes Butte nr. Papago Park, 1250’, 26 May 1975, M. Kolner, 1 male, 2 females (1 male, | female in copula) (ASU); California: Imperial Co.: Walters Camp Road, 5-6 May 1985, R. Parks, 1 male (CSU); 15 mi. E. Calexico, UV light, 5 June 1961, G. H. Nelson, 1 male (FSCA); Riv- erside: Gos Blythe; 10° July 1947, J> M. MacSwain, | male (USNM); Whitewater, 3 July 1967, W. F. Barr, 1 male (WFBM); MEXICO: Baja California: 13 mi. N. San Felipe, 9 June 1968, low sandy hills, sand burrs, Prosopis, Lophocerus, and Franseria, N. Leppla, J. Bigelow, M. Cazier, J. David- son, | male (ASU). Pseudonomoneura hirta (Coquillett) Figs. 5-8 Leptomydas hirta Coquillett 1905: 39. Ho- lotype male (USNM), USA: California: 24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 10tg | Me 4 Figs. 1-4. Pseudonomoneura californica. 1, Male terminalia, lateral view. 2, dorsal view. 3, ventral view. 4, Female terminalia, ventral view. (Abbreviations: f, furca; tg, tergite; uf, upper forceps of the epandrium.) Los Angeles County, Claremont, Baker, June, Coquillett collection. Washington, Type No. 7731; examined. D.C.; examined. Leptomydas concinnus Coquillett 1905: 39. | Nomoneura hirta: Hardy 1950: 15. (Synonymized by Hardy 1950: 15.) Lec- totype (here designated) male (USNM), Diagnosis.—Pseudonomoneura hirta is USA: California: Los Angeles County, closely related to P. micheneri and P. tink- VOLUME 97, NUMBER 1 Figs. 5-8. terminalia, ventral view. hami based upon the similarity of male gen- italia. In dorsal view the upper forceps of the epandrium of P. hirta, P. micheneri and P. tinkhami are very similar (Figs. 6, 10, 14). However, in lateral view the upper for- ceps of both P. micheneri and P. tinkhami have the upper inner flange strongly pro- Pseudonomoneura hirta. 5, Male terminalia, lateral view. 6, dorsal view. 7, ventral view. 8, Female duced and the apex bluntly truncate (Figs. 9, 13), whereas the upper inner flange of P. hirta is weakly developed and the apex nar- rowly rounded (Fig. 5). In ventral view the gonocoxites of P. hirta are cleft about one- fifth their length with the ventral processes strongly produced into two outwardly and 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ventrally projecting lobes (Fig. 7). Females of P. hirta are best separated by the presence of usually dense yellow pile on abdominal tergites, and the shape of the furca (Fig. 8). Discussion.— Hardy (1950) provides an adequate description of both the male and female, including the occurrence of two morphs, a more common one with con- spicuous yellow abdominal pile and popu- lations with gray abdominal pile. Separa- tion of unassociated females of P. hirta and P. micheneri can be difficult due to the sim- ilarity of terminalia and the variable col- oration of these two species. Through the kindness of Norman E. Woodley, Smithsonian Institution, syn- types of Leptomydas concinnus, a previ- ously recognized junior subjective synonym (Hardy 1950), were made available for study. These specimens are typical P. hirta. Pseu- donomoneura hirta is apparently common in areas of southern California. This species is very similar in general habitus to the sym- patric Nemomydas pantherinus (Gerstaeck- er) (Kondratieff and Welch 1990). Distribution.—California to Baja Cali- fornia. Material examined.— USA: California: Claremont Co.: “Topotype,” 1 male, 1 fe- male (USNM); 2 males, 3 females (USNM); Kern Co.: Short Cyn., 11 April 1976, J. B. Johnson, | male, 1 female (WFBM); Los Angeles Co.: Charlton Flat, San Gabriel Mts., 29 June 1975, Menke and Pulawski, 1 male (USNM); July, Coquillett coll., “Syntype,” 2 females (USNM); Glendora Ridge | mi. above Mt. Baldy Village, 17 June 1987, 1 female (LACMNH); Pine Can- yon, 6 July 1954, 2 males (LACMNH); Pearblossom, 2 mi. S., 3500’, 1-2 May 1977, R. R. Snelling, | male (LACMNH); Crystal Lake, 9 July 1952, D. Shepherd, 2 males, 1 female (LACMNH); Tanbark Flat, 20 July 1952, D. Shepherd, | female (LACMNH); Tanbark Flat, 12 July 1952, D. Shepherd, 1 female (LACMNH); Tanbark Flat, 25 June 1952, D. Shepherd, | female (LACMNH); Tanbark Flat, 21 June 1950, J. K. Windsor, 1 female (LACMNH); Tanbark Flat, 27 June 1950, J. K. Windsor, | female (LACMNH); Tanbark Flat, 29 June 1952, B. Tinglof, 1 male, | female (LACMNH); San Gabriel Can., 3200’, 28 June 1975, R. L. Westcott, 1 male (WFBM); Mono Co.: Sherwin Sum- mit, Hwy. 395, 7000’, 8 Sept. 1975, R. R. Snelling, on Chrysothamnus, 1 female (LACMNH); Orange Co.: Anaheim 7551 Vista Del Sol, 26 July 1977, ca. 180 m, P. H. Arnaud Jr., 1 male (CAS); Riverside Co.: Pinon Flat, San Jacnito Mts., 30 May 1939, P. D. Gerhardt, | male, 1 female (UA); San Bernardino Co.: Phelan, Sheep Crk. Can., 9 May 1949, A. L. Melander, 1 male (USNM); 1 female (USNM); vic. Cajon Summitt, 1250 m, 3 July 1980, R. L. West- cott, 1 male (CSU); Lytle Creek Road, 2700’, 4 mi. NW Nealey’s Corner, 5 June 1976, R. R. Snelling and C. D. George, 1 female (LACMNH); Camp Baldy, 8 July 1952, D. Shepherd, | female (LACMNH); same lo- cality but, B. Tinglof, 2 males, 2 females (LACMNH); same locality but, 26 June 1950, J. K. Windsor, 1 male (LACMNH); Lonepine Can., 5 mi. SE Wrightwood, 1375 m, 2 July 1980, R. L. Westcott, 1 male (WFBM); San Gabriel Mts., 2 mi. E Wright- wood, | June 1986, G. H. Nelson, | male, 1 female (FSCA); San Diego Co.: Pala, 6 June 1945, A. L. Melander, | male (USNM); Rutherford Ranch, San Feipe Valley, 8 May 1993, R. Parks, 1 male (CSU); Point Loma, 28 May 1978, L. Guidry, 1 male (CSU); Point Loma, 28 June 1977, L. Guidry, 1 male (CSU); Co.?: Mt. Lowe, 3 July 1917, J. M. Aldrich, 6 males, 4 females (USNM); Forest Home, 2 June 1934, C. Dammers, | female (USNM); San Gabriel Mts., 4000 ft., Long canon, 3 July 1910, F. Grinnell Jr., 1 male, 1 female (USNM); San Gabriel Mts., 1-3 July 1914, F. Grinnell Jr., 1 female (USNM); MEXICO: Baja California: Norte Sierra Juarez, 6 mi. N. of Laguna Hanson, 3 June 1982, Faulkner and Brown, | male (CSU); same locality but, 22 June 1980, 1 male (CSU); Ensenada, 31 May 1954, 1 male (LACMNH). VOLUME 97, NUMBER 1 Pseudonomoneura micheneri (James) Figs. 9-12 Nomoneura micheneri James 1938: 63. Ho- lotype male (WSU), California: Riverside County, 7 mi. S. White Water, 13 April 1935, C. D. Michener, in copulation; ex- amined. Nomoneura micheneri: Hardy 1950: 17. Diagnosis.— The long proboscis, extend- ing beyond the apex of the first flagellomere and the second flagellomere orange or tinged with red-orange will separate this species from the very closely related P. tinkhami. Additionally, males of P. micheneri can be distinguished by the more strongly pro- duced upper inner flange of the forceps of the epandrium (Fig. 9). The male genitalia of P. micheneri are also similar in structure to P. hirta, but in lateral view the upper inner flange of the upper forceps of P. micheneri is more strongly developed with the apex bluntly truncate (Fig. 9) in comparison to the weak- ly developed upper inner flange of P. hirta, with the apex narrowly rounded (Fig. 5). Additionally, in ventral view, the gonocox- ites of P. micheneri have the ventral pro- cesses straight (Fig. 11) rather than diver- gent (Fig. 7), as in P. hirta. Discussion. — Pseudonomoneura miche- neri is very similar to P. tinkhami, and not closely related to P. californica as previously indicated by Hardy (1950). Apparently, Hardy based this relationship on the simi- larity of the red abdominal tergites of some P. micheneri populations. Records of this species include a good series from the Ne- vada Test Site. This unique area has been well described by Knight (1968). Distribution.— Arizona, Nevada to Cal- ifornia. Material examined.—USA: Arizona: Mojave Co.: Chloride 4 mi. W., 28 April 1972, G. Bohart, P. Torchio and F. Parker, 1 male (USU); California: San Bernardino @or;_MorongosV;" 16-Apmil’ 1957; R: +R. Snelling, 2 males, 1 female (LACMNH); 11 2H] mi. N. Needles, 29 April 1982, Eriogonum, P. F. Torchio, 8 males, | female, | ? (missing tip of abdomen) (USU); 15 mi. N. Needles, 7 May 1982, P. F. Torchio, 1 male (USU); Cottonwood Wash, 2520’, Sec. 3 TON R12E, 10 May 1982, T. Griswold, 1 male (USU); Winston Wash, 2500’, Sec. 19 T109 R13E, 10 May 1982, T. Griswold, | male (USU); Kelso Dunes, 2600’, Sec. 30 TION R13E, 10 May 1982, T. Griswold, 1 female (USU); Lake Havasu, 15 April 1966, D. L. Coates, 2 males, 1 female (WFBM); San Diego Co.: 1 mi. S. Ocotillo Wells, 26 March 1977, R. W. Brooks, 1 male (UCD); Nevada: Clark Co.: Mercury, 28 April 1962, [BYU AEC NTS], 8 males, 11 females (USNM); same locality but 6 May 1961, 1 male, | female (USNM); same locality but 7 May 1961, 1 female (USNM); same locality but 12 May 1962, 1 female (USNM); Juanita Spr. Ranch S. of Riverside, 11-21 May 1983, F. D. and J. H. Parker, 3 males (USU); Riverside, 1 1- 21) Mayo1983:; FP oD.sandels Es-Parker,, | female (USU); Lincoln Co.: Alamo, 28 April 1973, F. Parker and P. Torchio, | female (USU). Pseudonomoneura tinkhami (Hardy) Figs. 13-16 Nomoneura tinkhami Hardy 1950: 18. Ho- lotype male (CAS) (lost), USA: Califor- nia: Inyo County, above Glacier Lodge, 21 August 1938, E. R. Tinkham. Diagnosis.— This species is very similar to P. micheneri but can be separated by the shorter proboscis, with the apex not ex- tending beyond middle of the first flagel- lomere, black antennae and shape of the upper forceps of the epandrium (Fig. 13). The male genitalia of P. tinkhami are also similar in structure to P. hirta, but in lateral view the upper inner flange of the upper forceps of P. tinkhami is more strongly de- veloped with the apex bluntly truncate (Fig. 13) in comparison to the weakly developed upper inner flange of P. hirta, with the apex narrowly rounded (Fig. 5). Additionally, in 28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9-12. 12, Female terminalia, ventral view. ventral view, the gonocoxite of P. hirta has the ventral process strongly developed into divergent lobes (Fig. 7). Females of P. tink- hami can distinguished most reliably from P. hirta by the shape of the furca (Fig. 16). Discussion.—The original description of 12 _ |i Pseudonomoneura micheneri. 9, Male terminalia, lateral view. 10, dorsal view. 11, ventral view. P. tinkhami was based ona single male from ‘above Glacier Lodge, California.” The ho- lotype was returned to the E. R. Tinkham Collection (Hardy 1950), which was later donated to the CAS. Unfortunately, this specimen is apparently lost (Dr. Paul H. Ar- VOLUME 97, NUMBER 1 Figs. 13-16. Pseudonomoneura tinkhami. 13, Male terminalia, lateral view. 14, dorsal view. 15, ventral view. 16, Female terminalia, ventral view. naud, Jr., personal communication). Ad- ditional inquiries were made to Mark O’Brien, Museum of Zoology, University of Michigan, where some of Tinkham’s ma- terial is deposited. However, no Pseudon- omoneura specimens were located. A male specimen in the CAS, labeled as a “‘topo- type” [of P. tinkhami] by the late Joseph Wilcox, is identical to Hardy’s (1950) orig- inal description and has the distinctive short proboscis as figured for P. tinkhami (Hardy 1950; Fig. 4a), and we base our concept of the species on this specimen. This late sea- son species is apparently rare and has a lim- ited distribution in California. Distribution. — California. Material examined.—California: Inyo Co.: Sherwin Summit, 14 Sept. 1963, J. Wil- 30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cox, | female (CAS); Big Pine, Glacier Lodge, 8-—9000', 9 Aug. 1957, Dorothy Mar- tin, 1 male (labeled as Topotype of P. tink- hami, Det. J. Wilcox) (CAS); Big Pine, 12 Aug. 1957, J. Wilcox, | male (CAS); Big Pine, 6500’, 7 Aug. 1966, J. Wilcox, 2 males, 1 female (CAS); Toms Place, 8 Aug. 1962, L. A. Stange, 1 male (CAS); Symmes Creek, 6900’, 18 Aug. 1982, A. S. Menke, | female (USNM); Mono Co.: 6.5 mi. NW Benton Hot Springs, 2000 m, 3 Aug. 1980, R. L. Westcott, 2 males, 2 females (WFBM). Pseudonomoneura bajaensis Fitzgerald and Kondratieff, NEw SPECIES Figs. 17-19 Holotype: Male (CAS), MEXICO: Baja California Sur, 10 June 1975, Pto. Chale, 60 km S. San Carlos, Howard E. Evans, Biological Note No. 2443. Paratypes: Male, MEXICO: Baja Cali- fornia Sur.: 7-9 June 1975, San Carlos, Howard E. Evans, W. Rubink, and D. Gwyne (CSU). There was no biological note attached to this specimen. Description.— Male: Frons densely sil- ver-white tomentose with dense white pile. Occiput, middle of face, and vertex black; vertex with white-yellow pile on either side of occiput. Antenna short, stout, black, with long black setae on pedicel and scape. Mouthparts vestigial. Thorax: Mesonotum opaque black in ground color with four broad densely tomentose, silver-white vittae, which are narrowly joined on the hind part of the thorax just before the scutellum. Pleu- ra shining dark brown. Halter yellow. Scu- tellum dark brown with thin white pile me- dially. Legs: Entirely brown. Hind leg with brown hair, femur swollen, and black flexor spines moderately developed. Pro- and me- sothoracic legs clothed with longer white se- tae. Wings: Hyaline 6.5—7 mm. Venation as in other Pseudonomoneura. Abdomen: Ter- gite | brown-black, tergites 2-6 dark brown- black with yellow-white posterior margins. Tergites 1-3 with long white pile, tergite | tomentose throughout, tergite 2 laterally and sparsely medially, and tergite 3 only anter- omedially. Remaining tergites with short black setae and some short yellow setae pos- terolaterally. Sternites brown. Bullae larger and black. Genitalia: Upper forceps of the epandrium bifurcate, in dorsal view inner lobe longer and square tipped, outer lobe shorter and acute (Fig. 18). In lateral view both lobes appear acute, the lower outer lobe shorter (Fig. 17). Gonocoxites with U-shaped cleft extending about one-fourth their length, ventral processes of the gono- coxites absent (Fig. 19). Female: Unknown. Etymology.—The epithet is derived from the Baja Peninsula. Diagnosis.—The male of P. bajaensis is easily distinguished from other Pseudono- moneura by the vestigial mouthparts and from P. nelsoni, n. sp. by the bifurcate upper forceps of the epandrium (Figs. 17, 18). The female is unknown. Discussion.— The holotype male was ex- cavated by H. E. Evans from cells ofa ground nest of Bembix rugosa Parker (Hymenop- tera: Sphecidae). This specimen was still fresh when collected and along with two specimens of Ablautus flavipes Coquillett (Asilidae), were provisions for a nearly full grown wasp larva (Evans 1976). The ter- minalia of the paratype are missing. Pseudonomoneura nelsoni Fitzgerald and Kondratieff, NEw SPECIES Figs. 20-24 Holotype: Male (CAS), USA: California: Riverside Co., 7 mi. S. W. Palm Springs, 7 September 1971, N. Papavero. Paratypes: USA: California, San Ber- nardino Co., Cottonwood Wash, S3T9NR- 12E, 2520’, 23 August 1981, T. Griswold, 1 male (USU); same data as holotype, 1 male, 19 females (CAS); Coachella Dunes (near Washington Street), 9 April 1988, R. Rogers, 3 males (LACMNH). Description.— Male: Frons densely sil- VOLUME 97, NUMBER 1 31 Figs. 17-19. Pseudonomoneura bajaensis. 17, Male terminalia, lateral view. 18, dorsal view. 19, ventral view. ver-white tomentose. Occiput black, middle of face black-brown. Antenna short, stout, light brown with short white setae on ped- icel and scape. Mouthparts vestigial. Tho- rax: Mesonotum densely tomentose opaque gray in ground color with four broad densely tomentose silver-white vittae, which are narrowly joined on the hind part of the tho- rax just before the scutellum. Pleura shining brown with some areas of thin silver-white tomentum. Halter yellow. Scutellum silver- white tomentose. Legs: Entirely light brown- yellow. Hind leg with brown-yellow hair, femur not swollen, and brown flexor spines weakly developed. Pro- and mesothoracic legs clothed with longer white setae. Wings: Hyaline, 5-7 mm. Venation as in other Pseudonomoneura. Abdomen: Tergites gray- 32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 20-23. Pseudonomoneura nelsoni. 20, Male terminalia, lateral view. 21, dorsal view. 22, ventral view. 23, Female terminalia, ventral view. VOLUME 97, NUMBER 1 33 Fig. 24. Pseudonomoneura nelsoni, female general habitus. brown with yellow-white posterior margins sometimes as wide as the posterior half of the tergite. All tergites with thin silver-white tomentum and long white decumbent pile. Sternites entirely yellow-brown with short yellow-white pile. Bullae dark brown. Gen- italia: In dorsal view the upper forceps of the epandrium slender and bent outward apically (Fig. 21), in lateral view, slightly square tipped, lacking upper and lower flanges (Fig. 20). Gonocoxites cleft only about one-eleventh of their length, with the ventral processes absent (Fig. 22). Female: Head: Yellow-brown with some light brown tomentum. Occiput dark brown shining. Antenna short, stout, yellow, about twice as long as head. Mouthparts vestigial. Thorax: Yellow-brown in ground color with three brown vittae on mesonotum. Meso- notal pattern densely tomentose. Halter yel- low. Pleura mostly shining with some to- mentose areas. Legs: Yellow, hind femur with strong flexor spines. Wings: Hyaline, 9-12 mm. Venation as in other Pseudono- moneura. Abdomen: Yellow-brown with dark brown markings anteromedially and anterolaterally on tergites (Fig. 24). Bullae small and inconspicuous. Abdomen glo- bose, wider than thorax (Fig. 24). Termin- alia as Fig. 23. Etymology.—The patronym honors Dr. C. Riley Nelson, University of Texas. He has provided much kind assistance with the junior author’s studies of Diptera. Diagnosis.— Males of P. nelsoni can be separated from all other species of Pseu- donomoneura with the exception of P. ba- Jaensis by the vestigial mouthparts. Addi- tionally, this species can be easily distinguished from P. bajaensis by the sim- 34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ple rather than bifurcate structure of the up- per forceps of the epandrium. The combi- nation of a globose abdomen (Fig. 24) and vestigial mouthparts will separate P. nelsoni from all other known females of Pseudon- omoneura. Discussion. — The form of the upper for- ceps of the epandrium of the male are unique within this genus, lacking the digitate struc- ture of P. californica, the bifurcate structure of P. bajaensis, or the upper and lower flang- es of P. hirta, P. micheneri and P. tinkhami. ACKNOWLEDGMENTS We thank the following persons who made valuable material available for study: Paul A. Arnaud, Jr., California Academy of Sci- ences; John T. Doyen, University of Cali- fornia, Berkeley; Carl Olson, University of Arizona; Robert Parks, Colorado State Uni- versity; Wilford J. Hanson, Utah State Uni- versity; Roy R. Snelling, Los Angeles Coun- ty Museum of Natural History; Howard V. Weems, Jr., Florida State Collection of Ar- thropods; Norman E. Woodley, Systematic Entomology Laboratory, National Museum of Natural History; and Richard S. Zack, Washington State University. Alison Wil- liams-Anderson provided the illustrations. Keve J. Ribardo, Curatorial Assistant, CAS provided useful data on California loca- tions. Norman E. Woodley provided help- ful comments on an earlier draft of the manuscript. LITERATURE CITED Artigas, J. N.and N. Papavero. 1990. Studies of My- didae (Diptera). V. Phylogenetic and biogeograph- ic notes, key to the American genera and illustra- tions of spermathecae. Gayana Zoologia 54: 87— 116. Bequaert, M. 1961. Contribution a la connaissance morphologique et a la classification des Mydaidae (Diptera). Brussels Institut Royal des Sciences Na- turelles de Belgique Bulletin 37: 1-18. Coquillett, D. W. 1904. Reports on Californian and Nevadan Diptera, I. Jn C. F. Baker. Diptera. In- vertebrata Pacifica 1: 17-39. Evans, H. E. 1976. Bembicini of Baja California Sur: Notes on nests, prey, and distribution. Pan-Pacific Entomologist 52: 314-320. Hardy, D. E. 1950. The Nearctic Nomoneura and Nemomydas (Diptera: Mydaidae). Wasmann Journal of Biology 8: 9-37. James, M. T. 1938. Notes on some North American Mydaidae (Diptera). Entomological News 49: 63- 64. Knight, H. H. 1968. Taxonomic review: Miridae of the Nevada Test Site and the Western United States. Brigham Young University Science Bul- letin, Biological Series 9: 1-282. Kondratieff, B. C. and J. L. Welch. 1990. The Ne- momydas of southwestern United States, Mexico, and Central America (Diptera: Mydidae). Pro- ceedings of the Entomological Society Washington 92: 471-482. Wilcox, J. and N. Papavero. 1971. The American genera of Mydidae (Diptera), with the description of three new genera and two new species. Arquivos de Zoologia, San Paulo 21: 41-119. Wilcox, J. 1981. Mydidae, pp. 533-540. In Mc- Alpine, J. F., B. V. Peterson, E. Shewell, G. E. Teskey, J. R. Vockeroth, and D. M. Wood, eds., Manual of Nearctic Diptera. Vol. 1. Agriculture Canada Monograph 27. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 35-45 HERMESIA LEFEVRE, A RESURRECTED GENUS OF NEOTROPICAL EUMOLPINAE (COLEOPTERA: CHRYSOMELIDAE) R. WILLS FLOWERS Agricultural Research Programs, Florida A&M University, Tallahassee, Florida 32307. Abstract. —Hermesia Lefévre is reinstated as a valid genus name in the Neotropical Eumolpinae. The genus is redefined to include the species H. aurata (Olivier), H. cyanea Bowditch, and H. inermis Bowditch. Characters for separation of Hermesia and Hylax Lefévre, and the disposition of other species formerly placed in Hermesia are given. Taxonomic characters found in the male endophallus are discussed and illustrated. Key Words: In the course of making an inventory of the Eumolpinae of Costa Rica and devel- oping a key to Central American genera of this taxonomically confusing subfamily, I repeatedly encounter instances where in- correct application of genus names causes confusion and needlessly complicates rec- ognizing phylogenetically meaningful groups of species. In the present paper I discuss the correct placement ofa small but showy group of eumolpines that are widespread in Neo- tropical forests and are frequently found in general collections of Chrysomelidae. Lefévre (1877) established the genus Her- mesia to include Colaspis aurata Olivier and two new species, H. purpurea and H. ful- gidicollis, later adding H. janthina Lefevre (1885). By the time of Blackwelder’s check- list (1946), 12 additional species had been described by Bowditch (1921), Jacoby (1882, 1900a, b, 1904), and Weise (1921). In his catalogue of the Neotropical Eumolpinae, Bechyné (1953) synonymized Hermesia with Hylax Lefévre (1884) and transferred Her- mesia aurata (Ol.) and Hermesia cyanea Bowditch to this genus. The remaining spe- cies of Lefévre’s Hermesia were transferred to other genera (see second list below). Lat- er, Bechyné (1954) stated that although the Chrysomelidae, Eumolpinae, Hermesia, Hylax, ovipositor, endophallus modified hind tibiae of the male H. auratus (Fig. 1) were not spinose as in most Hylax (Fig. 2), H. auratus was not otherwise sep- arable from the rest of Hylax. Still later, Bechyné (1955) reiterated his transfer of //. auratus, placed Rhabdopterus violaceus Ja- coby as a subspecies of Hy/lax auratus, and renamed Hylax violaceus (Jacoby) as Hylax pseudoviolaceus. While working with Costa Rican Eumol- pinae I found many specimens of a species lacking any modification of the hind tibiae of the male but agreeing closely with Hylax auratus in all other respects. This form proved to be Hermesia inermis Bowditch listed under Parachalcoplacis by Bechyné (1953) in his catalog. On further study, it became clear that H. auratus and P. inermis are congeneric, that both show substantial differences in structure of the pronotum from all Tylax that I have been able to examine, and that neither species is congeneric with Chalcoplacis (= Parachalcoplacis as syn- onomized by Monros and Bechyné 1956). I have examined a long series of P. inermis in the collections of the National Biodiver- sity Institute of Costa Rica, identified spec- imens of Hermesia and Hylax in the Bow- ditch Collection of the Museum of 36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 97, NUMBER 1 Comparative Zoology, Harvard University, and general collections of Eumolpinae at Cornell University and the California De- partment of Food and Agriculture. As a re- sult of these studies, I regard Hermesia as a valid genus, based on external characters and supported by internal characters of the male endophallus and the female oviposi- tor. Endophallic structures used in this study Terminology used for structures of inter- nal female genitalia follow Askevold and Flowers (1994). There is no accepted ter- minology for the structures of the male en- dophallus of Eumolpinae. Askevold (1988, 1990, 1991) has studied the Donaciinae and identified a system of sclerites of the male endophallus. Of these, the endophallic lat- eral digits (ELD), the basal supporting block (BSB), the median ejaculatory guide (MEG), and the basal setal brush (bb) appear to be analogous to structures visible in the en- dophalli of a number of Eumolpinae I have studied. I am provisionally adopting As- kevold’s terminology in this paper to refer to these structures, which are analogous in position and appearance to those illustrated for the Donaciinae. The most important dif- ference between the Donaciinae and the Eu- molpinae is that in the Donaciinae the en- dophallus is relatively short with the ELD’s in the apical position. In most Neotropical genera of Eumolpinae so far examined, the endophallus is a very long tube that, when retracted, is doubled back upon itself and lies within the basal hood. There appear to be two groups of sclerites that deploy as the endophallus is everted. The apical lateral digits (ELD in Figs. 8-17), analogous to the — Figs. 1-7. 37 endophallic lateral digits of Askevold, are associated with complex basal supporting block (BSB) from which a long flexible scler- ite (MEG) protrudes forward; the ELD’s are thus subapical and the endophallus contin- ues distally beyond them, sometimes for a considerable distance. At the base of the endophallus there are additional sclerites, presumably derived from the dorsal and ba- solateral sac supporting sclerites of Aske- vold. Among these is another pair of less sclerotized but movable basal lateral digits (BLD) which apparently have no homolog in Donaciinae (Askevold, pers. com.). Be- low the two sets of digits, there is often a field of fine setae on the underside of the endophallus (bb). The main difficulty in everting eumolpine endophalli is trying to pull this compact mass of sclerites, which is under tension when retracted, out through the delicate mem- braneous tube, and work it free without tearing the membrane. Failures are fre- quent. Fig. 8 shows the endophallus of H. aurata after an unusually successful prep- aration in which both sets of lateral digits and the MEG are everted. However, even here over half the endophallus is still re- tracted. The ELD’s are shown in apical view in Figs. 10 and 14. Basal digits (BLD) are blunt membraneous lobes with sclerotized distal surfaces (Fig. 13). Distad of the basal block, the endophallus is a simple mem- branous tube with a small internal apical sclerite (Figs. 11, 16) that may be a guide for the flagellum (not illustrated). Hermesia Lefévre 1877: clxxvili resurrected name Type species: Colaspis aurata Olivier 1808, designated by Bechyné (1950a). External characters of Hermesia and Hylax. 1, 2, posterior tibia of male (redrawn from Bechyné 1949). 1, Hermesia aurata. 2, Hylax viani. 3, fore (left) and middle (right) tibiae and tarsi of Hermesia aurata. 4, 5, prothorax of Hermesia aurata. 4, pronotum. 5, lateral view of prothorax. 6, 7, prothorax of Hylax sp. 6, pronotum. 7, lateral view of prothorax. CX1 = insertion of forecoxa. 38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SB Fm STE, Figs. 8-11. 9, apex of median lobe. 10, apical view of ELD’s. 11, apical sclerites of endophallus. Abbreviations: AP = apodeme, bb = basal setal field, BH = basal hood, BLD = basal lateral digit, BS = basal spur, BSB = basal supporting block, ELD = endophallic lateral digit, MEG = median ejaculatory guide, SBF = subbasal fenestra. Body oblong; length 4.8-6.2 mm; head, pronotum, elytra, underside, and legs bright metallic green, gold-green, or cobalt blue. Head with labrum apically emarginate, with 2 submedian setae and short row of lateral setae along outer apical angle. Frons and clypeus coarsely, sparsely punctate; punctures on clypeus separated by distance equal to the diameter of a puncture, and on frons by distance greater than the diameter of a puncture; surface between punctures smooth or with widely scattered punctulae; antennal calli impunctate; genae with mi- croreticulate area anteriorly between eye and base of mandible; frons with longitudinal impressed median line, deep between an- tennal calli, becoming obsolete toward ver- tex. Eyes oval, weakly emarginate at anten- nal insertion. Hermesia aurata, male genitalia. 8, lateral view of median lobe with partially everted endophallus. Antennae with scape oval, pedicel sub- globose, shorter than scape, distinctly short- er than segment 3; scape and pedicel yel- lowish to reddish brown ventrally, metallic green or cobalt blue dorsally; segments 3-6 reddish brown to piceous, usually paler at extreme apex, remaining segments piceous; all segments filiform, each slightly wider at apex, elongate (L/W seg. 7 = 3.5-4); seg- ments 3-6 with scattered adpressed setae, segments 7-11 densely pubescent, with whorl of long erect setae at apex of each segment from 3-10; segment | 1 short, con- ical. Mouthparts reddish brown to piceous; maxillary palp with apical segment spindle- shaped. Mandibles short, broad, strongly angulately curved, with outer surface smooth and shiny; apical teeth broad, acute. VOLUME 97, NUMBER 1 39 16 Figs. 12-16. Hermesia inermis, male genitalia. 12, lateral view of median lobe with partially everted en- dophallus. 13, frontal view of BLD’s. 14, apical view of ELD’s. 15, apex of median lobe. 16, apical sclerites of endophallus. Abbreviations as in Figs. 8-11. Prothorax distinctly wider than long, L/W = (0.48-0.61; disc of pronotum evenly, fine- ly punctate, punctures separated by 1-2 times the diameter of a puncture, surface between punctures smooth, shining, with scattered punctulae. Apical and basal mar- ginal bead narrow; lateral margin evenly rounded, with broad distinct flange, broad- est at middle, and with fine transverse striae on dorsal surface of flange (Fig. 4). Anterior and posterior angles distinct, outwardly di- rected, formed from projections of lateral flange, each angle with single long erect seta in large puncture. Proepisternum with an- terior margin straight, surface alutaceous to wrinkled. Proepimeron coarsely punctate, punctures separated by distance greater than the diameter of a puncture, surface smooth, shiny. Prosternum weakly declivous ante- rior to procoxae (Fig. 5), surface with long erect yellow setae, intercoxal process broad, 1.1-1.6 x diameter of procoxa, widened be- hind procoxae, longer than mesosternum, subequal to metasternum. Mesosternum broad, subequal in width to prosternum, strongly convex between coxae, width 1.4 x width of mesocoxa, sur- face smooth, slightly wrinkled with sparse short yellow setae. Metasternum smooth, slightly swollen anterior to metacoxae, with sparse short yel- low setae; metepisternum gradually nar- rowed posteriorly, surface alutaceous. Legs sparsely covered with short prostrate setae, all surfaces alutaceous. Femora strongly swollen in middle; tibiae gradually widening toward apex, multicarinate, slightly to moderately sulcate between ca- rinae, with setae linearly arranged in sulci and increasing in length toward apex of tib- 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 17. Abbreviations as in Figs. 8-11. iae. Hind tibiae of male with broad low in- ternal carina at middle (Fig. 1), or at apical fourth, or tibiae unmodified. Tarsi densely and uniformly pilose beneath, basal tarso- mere of fore- and middle legs expanded (Fig. 3), distinctly longer than wide; second tar- somere broadly triangular, with acute api- colateral angles; third tarsomere longer than second, deeply bilobed; terminal tarsomere distinctly surpassing apex of 3rd tarsomere; claws divergent, appendiculate. Elytra moderately punctate, punctures ar- ranged in 13-14 irregular rows separated by two or more times the diameter of a punc- ture; apical third of elytra with punctures in four regular rows which follow sutural and lateral margins; surface between punctures smooth, shining, with sparse punctulae; hu- meri prominent, rounded; basal calli mod- erately developed; postbasal depression present, deeper laterally. Sides subparallel, convergent; apices conjointly broadly rounded. Basal margin moderately costate, costa obliterated toward scutellum. Epi- pleuron narrow, acutely raised, slanted, ta- pering evenly from base to apex. Hylax sp., median lobe with partially everted endophallus (left), frontal view of BLD’s (nght). Scutellum triangular, base subequal to length; surface smooth, with few punctulae. Abdomen with all segments subequal in length, with sparse short yellow setae; with long yellow setae in central transverse row on four basal segments; surface between punctures finely alutaceous. Male sternum VI with shallow lateral depressions, ster- num VII with flattened smooth median area, coarsely punctate laterally, apical emargi- nation broad and shallow. Female sternum VI weakly crenulate on apical half, sternum VII flat, alutaceous, with strongly crenulate lateral margins, apical emargination nar- rower than in male, with median denticle (Fig. 24). Pygidium with longitudinal median groove broad, shallow, extending to apical margin (Fig. 23); lateral margin of groove slightly acutely projecting inward, demar- cated by subbasal bisinuate and apical ar- cuate carinae; surface alutaceous, with scat- tered coarse setiferous punctures; apical and lateral margins crenulate. Male genitalia.—Median lobe strongly curved (in lateral view) to sharply bent (Figs. VOLUME 97, NUMBER 1 41 25 49 Fig. 18-25. Female characters of Hermesia aurata and Hylax sp. 18-19, ovipositor of Hermesia aurata: 18, apex, lateral view; 19, ventral view. 20, Hy/ax sp., base of ovipositor tube (sternum VIII), ventral view. 21, 22, spermatheca: 21, Hermesia aurata; 22, Hylax sp. 23, pygidium of Hermesia aurata. 24, 25, female sternum VII: 24, Hermesia aurata; 25, Hylax sp. Abbreviations: ALA = apicolateral arms of sternum VIII, As = apodeme of sternum VIII, B = baculum, CS = coxostyli, DS = dorsal sclerites of segment VIII, GC = gonocoxae, HS = hemisternites of segment XI, PP = paraprocts, SD = spermathecal duct, SG = spermathecal gland. 42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 8, 12), apex pointed (Figs. 9, 15); basal hood (BH) long, lightly sclerotized, apodemes (AP) distinct at lateral margins of hood; subbasal fenestra (SBF) present; basal spurs (BS) prominent; tegmen slender. Endophallus elongate, with two pairs of lateral digits, the basal pair (BLD) simple, lobe shaped (Figs. 12, 13), the apical pair (ELD’s) bifurcate, articulating with a basal supporting block (BSB); a ventral basal brush (bb) between the two pairs of lateral digits; with thin me- dian ejaculatory guide (MEG) projecting forward when endophallus is everted (Figs. 8, 10); with small, internal, complex scerite at tip of endophallus (Figs. 11, 16); flagel- lum extremely long. Female genitalia. —Segments VIII-XI forming elongate ovipositor (Fig. 19). Ster- num VIII with long rod-like basal apodeme (A8) and weakly sclerotized apicolateral arms (ALA) with several setae; dorsal scle- rites (DS) weak, rod-like. Segment IX cov- ered with minute setae in basal half; hem- isternites (HS) with long basal rods, poorly sclerotized apically; paraprocts (PP) sepa- rated into pair of slender dorsal rods, api- cally forming hood-like projection above genital orifice (Fig. 18); baculum (B) dis- tinct, apical, subequal in length to gono- coxae (GC). Gonocoxae narrow, elongate, with long setae in apical half; coxostyli (CS) distinct, with several long apical setae. Sper- matheca (Fig. 21) with fine annuli, nar- rowed at ramus; duct short, straight, trans- parent, with sclerotized outlet into bursa. Species included Hermesia aurata (Olivier) Colaspis aurata Olivier 1808: 882 (original description) Chalcophana nitidissimus Erichson 1847: 162 (original description); Bechyné 1953: 165 (catalogue) Hylax auratus, Bechyné 1953: 165 (catalogue) Hermesia cyanea Bowditch 1921: 193 (original description) Hermesia inermis Bowditch 1921: 194 (original description) Parachalcoplacis inermis; Bechyné 1953: 170 (catalogue) Rhabdopterus violaceus Jacoby 1882: 151 (original description), new synonymy Hylax auratus violaceus; Bechyné 1953: 165 (catalogue) The remaining species listed as Hermesia in Blackwelder (1946) were placed by Be- chyné (1953) and Bechyné and Bechyné (1961) in the following genera. I can make no judgements at this time as to the cor- rectness of these generic placements; I can only confirm that they do not belong in Her- mesia as defined herein. Corysthea Baly: 1865: 336 gregalis (Weise) 1921: 49 rufa (Weise) 1921: 49 Hermesilla Bechyné: 1954: 216 fulgidicollis (Lefévre) 1877: clxxix f. lampros (Jacoby) 1900a: 352 Janthina (Lefévre) 1885: 39 similis (Bowditch) 1921: 193 Allocolaspis Bechyné: 1950b: 81 brunnea (Jacoby) 1900b: 489 confusa (Bowditch) 1921: 194 Jacobyi (Bowditch) 1921: 193 Lyraletes Bechyné: 1952: 15 purpurea (Lefévre) 1877: clxx1x varicolor (Jacoby) 1904: 514 Ledesmodina Bechyné: 1951: 263 erosula aenea (Jacoby) 1900a: 351 KEY TO MALE HERMESIA (Female Hermesia cannot be distin- guished at present except by association with males.) 1. Inner margin of hind tibia straight, lacking api- caléemarginateraneal- tae eee H. inermis Inner margin of hind tibia expanded, with api- cal emarginate area lined with long setae (Fig. |) Renee et Mera otek at oats ba emia 2 . Hind tibia expanded at middle, apical emar- ginate area almost one-half the length of tibia Rad gE Ee eh ig See Ite H. aurata N VOLUME 97, NUMBER 1 Hind tibia expanded in apical third, emargin- ate area no more than one-fourth the length of [OVE vile Bias has Geog eed es eae eee ere H. cyanea DISCUSSION The single male of Rhabdopterus viola- ceus mentioned by Jacoby and deposited in the Bowditch Collection is a Hermesia lack- ing tibial modifications, hence this form be- longs under H. inermis. Both H. aurata and H. inermis are found in bright metallic green and cobalt blue color forms (all specimens of H. cyanea J have seen are metallic green). Males of Hermesia I have seen from Central America have all been H. inermis; this spe- cies also extends into Colombia (Bowditch Collection); H. aurata and H. cyanea are apparently limited to South America. In the field, the brightly colored Hermesia species are likely to be confused only with members of a group of Colaspis that also are bright metallic green or cobalt blue (the ‘‘bridarollei” group of Bechyné). These Co- laspis, however, have clear yellow to rufo- testaceous legs (in Hermesia the legs are me- tallic, always the same color as the elytra) and much more densely and/or coarsely punctate elytra and pronotum. Host plant data for these species are (as usual for the Eumolpinae) scarce but I have collected H. aurata from leaves of Psychotria (Rubi- aceae) in Rondonia, Brazil. Hermesia, as here delineated, is most similar to Lyraletes Bechyné but can be dis- tinguished by the following combination of characters (based on specimens referable to Lyraletes in the Bowditch Collection): the lateral flange of the pronotum is distinctly sinuate in Lyraletes, evenly rounded in Her- mesia; and the elytra of Lyraletes are widest in their apical third (in Hermesia the sides are subparallel). Aside from differences in the hind tibiae of the males (discussed above), Hermesia can be clearly differentiated from Hy/ax on the following characters. In Hermesia the prosternum (Fig. 5) is gradually declivous 43 anteriorly and its anterior margin meets the gula well behind the mouthparts. This con- dition is widespread in the Eumolpinae and can be seen in the familiar North American Colaspis and Brachypnoea (= Nodonota, see Flowers et al. 1994) species. In Hylax, on the other hand, the anterior margin of the prosternum is somewhat to distinctly con- cave for reception of the postgenal area of the head. Viewed in profile (Fig. 7) the pro- sternum appears to meet the gula almost perpendicularly. When the head is in the resting position, the prosternum rests against the gula close to the base of the mouthparts. This condition is less common in the Eu- molpinae but is found in other Neotropical genera such as Eumolpus Weber and Le- pronota Chapuis. The pronotal lateral marginal bead of Hy- lax is narrow, evenly rounded, lacks stria- tions and is slightly thickened as it passes around the apex of the apical angles (Fig. 6). The anterior apical angles are very large, directed forward and formed from the pro- notum itself. This is quite different from the wide striate bead of Hermesia (Fig. 4) which ends in a small beveled angle at the anterior apical angle. The apical abdominal sternite of the fe- male is quite different between the two gen- era for all specimens examined. In Her- mesia, the lateral margins are strongly crenulate and the apical notch is narrow, with a median tooth (Fig. 24). In Hylax, this sternite is smooth on the margins and the apex has a broad bisinuate emargination (Fig. 25). There are some differences between the genitalia of Hermesia and Hylax (based on dissection ofa series of H. aurata from Peru, H. inermis from Panama and Costa Rica, and specimens of Hy/ax nr. chalybaeus Lef. from Costa Rica). In females, segment VIII is more extensively sclerotized in Hylax, the basal apodeme has a more marked apical expansion (Fig. 20); the spermatheca has a different shape (Fig. 22), and the sperma- 44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON thecal duct is very long and convoluted. In males, the general morphology of the Hylax endophallus is like that of Hermesia but shape of both pairs of lateral digits (ELD and BLD) is different in the two genera. In Hylax the basal setal brush is much more extensive than in Hermesia, the setae are arranged in definite rows and there are rows of setae on the dorsal surface behind the basal digits (Fig. 17). The median lobe apex also differs slightly but this character varies within genera and is not of generic value. It is difficult to say if differences in these struc- tures identified here are phylogenetically useful at the genus level. They do, however, weigh in favor of restoring Hermesia. ACKNOWLEDGMENTS This research was begun during a six- month sabbatical with the National Biodi- versity Institute of Costa Rica (INBio) in 1991. I thank Dr. Rodrigo Gamez, MSc. Angel Solis and the INBio staff for their many kindnesses during that time. I also thank Dr. David G. Furth, Museum of Comparative Zoology (MCZ), Harvard University, for his invaluable assistance in providing specimens from the Bowditch Collection for this project. I also thank Dr. Michael C. Thomas, Florida State Collec- tion of Arthropods (FSCA), Gainesville, Florida, and Dr. Fredrick G. Andrews, Cal- ifornia Department of Food and Agricul- ture, for the loan of additional specimens; and Dr. Ingolf S. Askevold for useful com- ments on the manuscript. This research was funded in part by a grant (FLAX 91005) and in part by a National Science Foun- dation Mid-Career Fellowship (BSR- 9003898). LITERATURE CITED Askevold, I. S. 1988. The genus Neohaemonia Szké- kessy in North America (Coleoptera: Chrysomel- idae: Donaciinae): Systematics, reconstructed phylogeny, and geographic history. Transactions of the American Entomological Society 113: 361- 430. 1990. Reconstructioned phylogeny and re- classification of the genera of Donaciinae (Cole- optera: Chrysomelidae). Quaestiones Entomolgi- cae 26: 601-664. 1991. Classification, reconstructed phyloge- ny, and geographic history of the New World members of Plateumaris Thomson, 1859 (Cole- optera: Chrysomelidae: Donaciinae). Memoirs of the Entomological Society of Canada 157: 1-175. Askevold, I. S. and R. W. Flowers. 1994. Glyptos- celoides dentatus, a genus and species of Eumol- pinae new to Chile (Coleoptera: Chrysomelidae). Revista Chilena de Entomologia. (In press.) Baly, J.S. 1865. Descriptions of new genera and spe- cies of Phytophaga. Transactions of the Entomo- logical Society of London, series 3 2: 333-357. Bechyné, J. 1949. List provisoire des Eumolpides de la République Argentine [et observations diverses sur les Eumolpides de l’Amerique du Sud] (Co- leoptera, Chrysomeloidea). Acta Zoologica Lil- loana 8: 457-535. 1950a. Les générotypes des eumolpides de Amérique du Sud et du Centre avec les diagnoses des formes nouvelles. (Coleoptera, Phytophaga, Chrysomeloidea). Mittilungen Miinchen Ento- mologische Gesellshaft 40: 264-292. 1950b. Notes sur les Eumolpides de l’Amé- rique du Sud, (Col. Phytophaga). Annals and Mag- azine of Natural History. Series 12 3: 75-85. 1951. Liste provisoire des Eumolpides de Bolivie et observations diverses sur les espéces de Amérique du Sud. (Col. Phytophaga). Entomo- logischen Arbeiten aus dem Museum Georg Frey 2: 227-352. . 1952. Duxiéme note sur les Eumolpides néo- tropicaux des collections de I’Institut royal des Sci- ences naturelles de Belgique (1). Institut royal des Sciences naturelles de Belgique. Bulletin 28: 1-20. 1953. Katalog der neotropischen Eumolpi- den (Coleoptera Phytophaga Chrysomeloidea). Entomologischen Arbeiten aus dem Museum Georg Frey 4: 26-304. . 1954. La liste des eumolpides de Rio Grande do Sul (Bresil) et observations diverses sur les es- péces néotropicales. Arquivos do Museu Paran- aense 10: 141-230. 1955. Reise des Herrn G. Frey in Siidamer- ika: Eumolpidae (Col. Phytophaga). Entomolo- gischen Arbeiten aus dem Museum Georg Frey 6: 569-657. Bechyné, J. and B. Springlova de Bechyné. 1961. In- secta amapaensia: Chrysomeloidea (Col. Phy- toph.). Studia Entomologica 4: 409-428. Blackwelder, R. E. 1946. Checklist of the coleopter- ous insects of Mexico, Central America, the West Indies, and South America. Part 4. United States National Museum Bulletin 185: 551-763. VOLUME 97, NUMBER 1 Bowditch, F.C. 1921. South American Eumolpidae, mostly of the group Colaspini. Entomologist 54: 25-30, 72-76, 169-172, 190-195, 214-216, 234— 236, 253-256. Erichson, W. F. 1847. Conspectus insectorum co- leopterorum quae in Republica Peruana observata sunt. Archiv fiit Naturgeschichte 13: 67-185. Flowers, R. W., D.G. Furth,and M.C. Thomas. 1994. Notes on the distribution and biology of some Florida leaf beetles (Coleoptera: Chrysomelidae). The Coleopterists’ Bulletin 48: 79-89. Jacoby, M. 1882. Insecta, Coleoptera, Eumolpidae, Chrysomelidae. 6: 145-224. In Godman, F. D. and O. Salvin, eds., 1879-1901. Biologia Centrali- Americana. Dulach & Co., London. 1900a. Descriptions of two new species of Hermesia (Chrysomelidae, fam. Eumolpidae). Annali del Museo Civico di Storia Naturale di Genova 40: 351-353. . 1900b. Descriptions of new species and a new genus of South American Eumolpidae, with re- marks on some of the genera. Transactions of the Entomological Society of London 1900: 453-510. 1904. Descriptions of some new species of phytophagous Coleoptera obtained by Baron E. Nordenskiéld in Bolivia and the Argentine Re- public. Arkiv for Zoologi 1: 513-524. 45 Lefévre, E. 1877. (La description suivante d’un genre nouveau de coléoptéres, appartenant a la famille des eumolpides.) Bulletin de la Société Entomo- logique de France 1877: clxxvili-clxxix. 1884. (Description d’un nouveau genre de la famille des emolpides.) Bulletin de la Société En- tomologique de France 1884: xliv—xlv. 1885. Eumolpidarum hucusque cognitarum catalogus, sectionum conspectu systematico, ge- nerum sicut et specierum nonnullarum novarum descriptionibus adjunctis. Mémoires de la Société Royale des Sciences de Liége, serie 2 11: 1-172. Monros, F. and J. Bechyné. 1956. Uber einige ver- kannte Chrysomeliden-Namen. Entomologischen Arbeiten aus dem Museum Georg Frey 7: 1118- SHE Olivier, A. G. 1808. Entomologie, ou histoire natu- relle des insectes, avec leurs caractéres génériques et spécifiques, leur description, leur synonymie, et leur figure enluminée. Coléoptéres 6: 613-1104. Paris. Weise, J. 1921. Wissenschaftliche Ergebnisse der schwedischen entomologischen Reise des Herrn Dr. A. Roman in Amazonas 1914-1915. 6. Chrys- omelidae. Arkiv for Zoologi 14: 1-205. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 46-49 THE LARVA AND PUPA OF THE CADDISFLY SPECIES, HELICOPSYCHE PARALIMNELLA HAMILTON (TRICHOPTERA: HELICOPSYCHIDAE) MICHAEL A. FLoyp! Clemson University, Department of Entomology, Long Hall, Box 340365, Clemson, South Carolina 29634-0365. Abstract. — Descriptions and illustrations of the larva and pupa of the caddisfly species, Helicopsyche paralimnella, are presented. Although the larva of this species most closely resembles that of H. borealis, diagnostic characteristics are noted. This species possesses a case typical of all other North American species. The known distribution of this species is limited to three localities in southwestern North Carolina and northwestern South Carolina. Key Words: Trichoptera The caddisfly genus Helicopsyche is rep- resented by six species in America north of Mexico. These include Helicopsyche bo- realis (Hagen) (widespread and common throughout much of North America), H. limnella Ross (Arkansas and Oklahoma), H. mexicana Banks (Arizona, California, Texas, and Utah), H. paralimnella Hamil- ton (North and South Carolina), H. piroa Ross (Kansas, Louisiana, and Texas), and H. sinuata Denning and Blickle (California) (Hamilton and Holzenthal 1984). Although only the immature stages of H. borealis have been described and illustrated (Vorhies 1909, Elkins 1936, Ross 1944, and Wiggins 1977), larval characters (1.e. color patterns of the head capsules) have been found that will readily separate three additional spe- cies: H. limnella, H. mexicana, and H. piroa (Hamilton and Holzenthal 1984, S. W. Hamilton, pers. comm.). ' Present address: Commonwealth Technology, Inc., 2520 Regency Road, Lexington, Kentucky 40503-2921. Caddisfly, Helicopsyche, Helicopsychidae, North Carolina, South Carolina, Collections that I made in the Blue Ridge Mountains of North and South Carolina, as well as examination of additional material (Morse et al. 1989) housed in the Clemson University Arthropod Collection (CUAC), have resulted in discovery of the previously unknown larva and pupa of H. paralim- nella. Association of immatures and adults was made by using metamorphotypes (Milne 1934, Wiggins 1977). Morphological ter- minology follows that of Wiggins (1977). Representative specimens have been de- posited in the Entomology Collection of the Royal Ontario Museum (ROM), the Illinois Natural History Survey (INHS), the U.S. National Museum of Natural History (USNM), and the CUAC. Helicopsyche paralimnella Hamilton 1989 (Figs. 1-7) Helicopsyche paralimnella Hamilton, in Morse et al. 1989: 30. Material examined. —NORTH CARO- LINA: Jackson/Transylvania County, VOLUME 97, NUMBER 1 47 0.5mm 6 0.25mm 0.25mm Figs. 1-7. Helicopsyche paralimnella larva, larval case, and pupa. 1-3, larva: 1, head, pronotum, and meso- notum, dorsal view; 2, abdominal segment IX, right posterolateral view. 3, anal claw, lateral view. 4, 5, larval case: 4, lateral view; 5, dorsal view. 6, 7, pupa: 6, right mandible, ventral view; 7, abdominal segments VIII and IX with anal rods, dorsal view. 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Whitewater River at Route 281, 27 June 1991, M. Floyd, 3 larvae, 2 pupae (ROM); 3 larvae, 2 pupae (INHS); 3 larvae, 2 pupae (USNM); 4 larvae, | pupa (CUAC); same locality, 25 Oct. 1989, E. Chen, 1 larva (CUAC); same locality, 7 May 1991, K. Hoffman, 5 larvae (ROM), same locality, 16 March 1989, E. Chen, 3 larvae (INHS), same locality, 7 May 1991, J. Morse, 15 larvae (USNM); M. Floyd, 59 larvae (CUAC); SOUTH CAROLINA: Oconee County, Thompson River at North Caro- lina border, 18 May 1987, S. Hamilton and K. Hoffman, 2 larvae (CUAC); Pickens County, Wildcat Creek, Clemson Univer- sity Experimental Forest, 28 August 1986, J. Wilson, 2 larvae (CUAC). Mature larva. — Figs. 1-3. Length 4-5 mm. Head dark brown except for lightly pig- mented labrum, periocular areas, and por- tion of genae anterior to antennae (Fig. 1). Pronounced carinae running posteriad from anterolateral corners of frontoclypeal apo- tome near antennae, mesad to eyes, then forking to posterolateral corners of fronto- clypeal apotome and to coronal suture. An- terior half of frontoclypeal apotome twice as wide as posterior half, the apotome lyre- shaped, with sinuous lateral margins. Pro- notum dark brown, heavily setose; anterior margin with regular row of about 40 to 50 short, stout spines; few dark muscle scars present on posterior half. Foretrochantin long, with apical seta. Mesonotum brown, with contrasting darker muscle scars and dark, irregular bands running from meson to mesolateral and anterolateral areas; pale, circular area present at midlength of mid- dorsal ecdysial line, just anterior to pair of dark muscle scars. Lateral sclerite of ab- dominal segment I roughly triangular, with many small spines and one seta along pos- terior border (similar to Fig. 5.1A, Wiggins 1977). Abdominal segment VIII with row of lateral tubercles (similar to Fig. 5.1E, Wiggins 1977). Abdominal segment IX with pair of dorsomesal tufts, each with five setae (Fig. 2). Lateral sclerite of anal proleg heavi- ly sclerotized and dark brown, with five to six long, posterior setae. Anal claw comb- like (Fig. 3). Pupa.— Figs. 6, 7. Length 5 mm. Man- dibles long, blade-like, without teeth or ser- rations (Fig. 6). Anal rods of abdomen nar- row, finger-like, with posterior setae nearly twice as long as rods (Fig. 7). Case.— Figs. 4, 5. Length 5 mm. Snail- shaped, composed of fine sand grains. Dor- sal lip of anterior opening extending hood- like over ventral lip. Discussion: The larva of H. paralimnella most closely resembles H. borealis. It can be differentiated from H. borealis by the lack of muscle scars on the head, the pres- ence of darkly pigmented lateral sclerites on the anal proleg, and its small size (mature larva <5 mm). These two species have not been found to occur in the same streams. The Ozarkian species, H. limnella, the probable sister species of H. paralimnella (Morse et al. 1989), is much less darkly pig- mented and has large, pale muscle scars on the head. Except for its small size, the pupa of H. paralimnella does not seem to differ from those of other North American Heli- copsyche species. In addition, no distin- guishing characters have been found for pu- pae of H. borealis, H. limnella, H. mexicana, or H. piroa (Hamilton and Holzenthal 1984). At present, H. paralimnella is known from only three streams in North and South Car- olina. Thompson River (Oconee County, South Carolina) and Whitewater River (Jackson/Transylvania County, North Car- olina) are clear, fourth order, mountain streams (439 m altitude), which have mod- erate to fast current and substrates com- posed of bedrock, cobble, and smooth boul- ders. Riparian vegetation is composed primarily of rhododendron and mixed hardwoods. Wildcat Creek is a clear, third order, upper Piedmont stream (240 m al- titude) with a sand and cobble substrate. Riparian vegetation consists of mixed hard- woods. Larvae and pupae were found either in bedrock areas among patches of river VOLUME 97, NUMBER 1 weed, Podostemum sp., or the underside of cobble and boulders. Morse et al. (1989) reported adult collection dates of 15 June to 21 July. ACKNOWLEDGMENTS I would like to thank Drs. Steve Hamil- ton, Austin Peay State University, and Ralph Holzenthal, University of Minnesota, for making available their illustrations and notes concerning Helicopsyche larvae. I also thank Dr. Steven Moulton, II, University of North Texas, for providing invaluable information concerning Ozarkian species. Helpful reviews of the manuscript were pro- vided by Drs. Peter Adler, Kevin Hoffman, and John Morse (Clemson University). This is technical contribution no. 3532 of the South Carolina Agricultural Experiment Station, Clemson University. LITERATURE CITED Elkins, W. A. 1936. The immature stages of some Minnesota Trichoptera. Annals of the Entomo- logical Society of America 29: 656-681. 49 Hamilton, S. H. and R. W. Holzenthal. 1984. The genus Helicopsyche in America North of Mexico (Trichoptera: Helicopsychidae), p. 16. Jn Morse, J. C., ed., Proceedings of the Fourth International Symposium on Trichoptera, Clemson, South Car- olina. Series Entomologica 30, Dr. W. Junk Pub- lishers, The Hague. 486 pp. Milne, L. J. 1934. Studies in North American Tr- choptera, Part |. Privately published, Cambridge, Massachusetts. 19 pp. Morse, J. C., S. W. Hamilton, and K. M. Hoffman. 1989. Aquatic insects of Lake Jocassee Catch- ment in North and South Carolina, with descrip- tions of four new species of caddisflies (Trichop- tera). Journal of the Elisha Mitchell Scientific Society 105: 14-33. Ross, H. H. 1944. The caddis flies, or Trichoptera, of Illinois. Bulletin of the Illinois Natural History Survey 23: 1-326. Vorhies, C. T. 1909. Studies on the Trichoptera of Wisconsin. Transactions of the Wisconsin Acad- emy of Sciences, Arts, and Letters 16: 647-738. Wiggins, G. B. 1977. Larvae of the North American Caddisfly Genera (Trichoptera). University of To- ronto Press. 401 pp. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 50-62 FOUR NEW SAWFLIES FROM EASTERN NORTH AMERICA, THREE SPECIES OF TENTHREDO AND ONE OF DOLERUS (HYMENOPTERA: TENTHREDINIDAE) HENRI GOULET AND DAVID R. SMITH (HG) Biological Resources Division/CLBRR, Agriculture Canada, K. W. Neatby Build- ing, Ottawa, Ontario, Canada, K1A 0C6; (DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, Washington, D.C. 20560, U.S.A. Abstract.—Three new species of Tenthredo and one new Dolerus from eastern North America are described and illustrated. Tenthredo appalachia and T. masneri occur from southern Canada to Great Smoky Mountains National Park in North Carolina and Ten- nessee. Tenthredo fernowi is more restricted and is known only from northern Virginia and West Virginia to Great Smoky Mountains National Park. Though rarely previously collected, adults of these three species are common in collections from a broadleaf forest in West Virginia. Dolerus klokeorum was discovered in collections from three localities in the Virginia Piedmont. Flight records and habitat data are given for each species. Key Words: Four sawflies are described here so that names will be available for future papers on sawfly studies in the mid-Atlantic states. The three species of Tenthredo are relatively common in collections from within a broad- leaf forest of the central Appalachians of West Virginia. The Dolerus is from Virgin- ia, and it is an addition to the revision of nearctic Dolerus (Goulet 1986). Most specimens of the three Tenthredo species were collected during a study of non- target insects and diflubenzuron (E. M. Bar- rows, Georgetown University) in a broad- leaf forest in the central Appalachians, the Fernow Experimental Forest, Monongahela National Forest, about three miles south of Parsons, Tucker Co., West Virginia (39°3'N, 79°40'W; map in Griffith and Perry 1992). The forest is dominated by oaks, intermixed primarily with beech, sweet birch, maple, yellow poplar (tulip tree), black cherry, pin cherry, white ash, basswood, rhododen- Dolerus, North America, sawflies, Tenthredinidae, Tenthredo dron, and black locust (Anonymous 1987). Elevations of the collection sites vary be- tween 2300 to 2600 feet. Twenty Townes- style Malaise traps were utilized, five in each of four watersheds, for the non-target study, and all were in operation from mid-April to the end of September each year. In each watershed, one trap was adjacent to a stream, two traps were about 20 m up each slope, and another 2 traps were about 40 m up each slope. All were within dense forest. The Malaise trap numbers in the specimen data sections indicate the watershed-trap num- ber, e.g. 4-1 1s watershed 4, trap 1, etc. Trap 1 was the top trap on the northerly-facing slope, trap 2 the middle trap on this slope, trap 3 near a stream, trap 4 the middle trap on the southerly-facing slope, and trap 5 the top trap on this slope. Trap numbers of 6 or higher are Cornell-style Malaise traps set up at forest edges. Much of the terminology, especially for VOLUME 97, NUMBER 1 sculpturation, is based on Goulet (1986). Format and terminology are consistent with that being used for ongoing revisionary studies of Nearctic Tenthredo by the senior author. The character choice in descriptions is a function of species closely related to the one being described. Tenthredo appalachia, n. sp., and 7. masneri, n. sp., are closely related, and their descriptions are similar. Tenthredo fernowi, n. sp., is related to T. rufopecta (Norton), 7. nimbipennis Cres- son, and 7. mellicoxa Provancher, and its description stresses characters significant among these species. Tenthredo appalachia Goulet and Smith, NEw SPeEcIES (Figs. 1, 4, 9) Diagnosis.—Adults are distinguished from those of other Nearctic species of 7en- thredo with minute pulvilli (length of pul- villus of first metatarsomere 0.15-0.20 x length of second metatarsomere) and with microsculpture on the mesonotal median lobe by the following character combina- tion: ventral portion of metasternum com- pletely orange, clypeus white (except on bas- al one-fourth to one-third), and tergites 2- 8 not appreciably paler along posterolateral margin. Female.— Length, 10.5—12.0 mm. Anten- na and head black, with apical *4 clypeus, mandible, labium, and maxillary and labial palpi white; minute white spot present on lower outer orbit (may be lacking) and one on upper inner orbit. Thorax black with teg- ula, spot on lower posterior margin of me- sepisternum near mesocoxa, and metepis- ternum white; posteromedial portion of pectus orange. Abdomen black with lateral margin of tergite 1 white and basal sternite and sometimes part of sternite 2 orange. Legs orange with apical *4 of metatibia and metatarsus black. Wings hyaline; veins and stigma black. Head in dorsal view slightly narrower be- hind eyes (maximal distance between outer 51 margins of eyes 1.05 maximal distance between outer margins of gena). Antennal flagellum long: sixth flagellomere 4 x longer than wide and first flagellomere 1.10-1.20 x as long as second flagellomere. Head and body mostly shining and impunctate, with thorax more dulled by surface sculpture than head. Malar area near mandible with con- vex sculpticells and fine punctures (10-15 um in diameter); central portion of gena with lightly imprinted meshes of microsculpture; remainder of head smooth. Mesepisternal spine obtusely angular (about 130°, Fig. 9). Most of thorax with fine surface micro- sculpture, with rugose microsculpture on posterior half of mesepisternum, and with- out microsculpture on medial portion of posttergite, on lateral lobe of mesoscutum, and anterior half of mesoscutellum. Punc- tures on pectus and on mesoscutum fine (10- 15 wm in diameter), 20-30 um apart on me- dian lobe and pectus, and 10-40 um apart on lateral lobe; punctures of mesoscutellum larger (30 wm in diameter) and 30-60 um apart. Abdomen shining but with fine sur- face microsculpture, meshes of microsculp- ture about 10 um in length by 15-60 um in width, posterior margin of sculpticells clear- ly elevated and scale-like. Setae lacking at base of tergite 1. Pubescence developed over half of tergite 7, and dense on tergites 8-9, 40-50 wm apart. Tarsal pulvilli minute (length of pulvillus of first metatarsomere 0.15-0.18x length of second metatarso- mere) and narrow (width of pulvillus of first metatarsomere about '3 apical width of sec- ond metatarsomere) (Fig. 4). Serrulae of lancet as in Fig. 1. Male.— Length, 10.0 mm. Similar in col- oration to female except for the following: gena with large white spot on lower third; pronotum in lateral view with white spot on ventral surface; mesepisternum black with white spot extended from mesocoxa forward beyond middle, ventral portion of pectus black anteriorly but gradually be- coming orange in posterior half; tergites 1- 8 dark brown but white at side of tergite | >2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and whitish on medial half of tergites 2-5; sternites 2—5 whitish orange. Head in dorsal view clearly narrower be- hind eyes (maximal distance between outer margins of eyes 1.14 maximal distance between outer margins of gena). Setae at base of tergite 1 present only at side and medially. Metatarsus in dorsal view narrow as in female; metatarsomere 2 about 6x longer than wide. Genitalia as in Fig. 6; pe- nis valve with apical spine. Holotype.— Female, labeled “West Vir- ginia: Tucker Co., Fernow Expt. Forest, 31- V-9-VI-91, E.M. Barrows,” “‘Malaise trap 7-2.” Deposited in the National Museum of Natural History, Washington, D.C. Paratypes (14 F, 1 M).—CANADA: QUEBEC: Knowlton, 1-VII-36 (1M). U.S.A.: NEW HAMPSHIRE: Lancaster, Mount Prospect, 19-VI-82 (1F). NORTH CAROLINA/TENNESSEE: Great Smoky Mts. Natl. Park, 8-VII-57 (2F). WEST VIR- GINIA: Same data as holotype except for dates and trap numbers, 10-19-VI-91, trap 1-2 (1F); 20-29-VI-91, trap 1-3 (1F); 21- 30-V-91, trap 4-1 (1F); 10-19-VI-91, trap 4-2 (1F), trap 4-6 (1F), trap 7-6 (1F); 10- 19-VII-91, trap 13-3 (1F); 20-29-V-92, trap 1-2 (1F), trap 7-6 (1F), trap 13-6 (1F); 19-V- 28-VI-92, trap 7-6 (1F). Deposited with the holotype and in the Canadian National Col- lection, Ottawa. Distribution. — This species occurs in the Appalachian Mountains from Great Smoky Mountains National Park northwards to southernmost Quebec. Etymology.—The specific epithet is a noun in apposition and refers to the Ap- palachian Mountain range distribution of this species. Discussion.—Females of 7. appalachia have a mostly black body with orange legs except for the black apical one- to two-thirds of the metatibia and the metatarsus. This color pattern is very similar to those of T. leucostoma Kirby and other similarly col- ored species with an angular mesepisternal spine. The male is rather different in color pattern from those of 7. /eucostoma and similarly colored species. Females of T. /eu- costoma and similarly colored species are differentiated from 7. appalachia as fol- lows: pulvilli very long and wide (length of pulvillus of first metatarsomere 0.40-0.70 x length of second metatarsomere; width of pulvillus of first metatarsomere about two- thirds that of second metatarsomere) (Fig. 5); ventral portion of metepisternum black or black with white on posterior half; pos- terior margin of pronotum white or black, apical one- to two-thirds of metatibia black; and head slightly duller because of surface sculpture, especially on hindorbits. The male of T. leucostoma and of species alluded to above have black stripes or spots on the coxae, trochanters, and femora. From 7. masneri, its closest species, 7. appalachia is distinguished by the black posterior mar- gin of the pronotum in dorsal view, black posterolateral margin of tergites 2-8, black posterior margin of tergite 1, wider black base of clypeus, longer antennal flagellum (sixth flagellomere 4x longer than wide), smaller pulvilli, and lack of setae over at least portion of base of tergite 1. This species is part of the lineage to which most North American Tenthredo species belong. The following species groups pro- posed by Ross (1951) and followed by Smith (1979) belong in this lineage: basilaris, pec- toralis, rufopecta (only T. rurigena Mac- Gillivray), angulifera, leucostoma (exclud- ing nimbipennis), occidentalis, verticalis, semirufa, secunda, xantha, and begimina. This lineage is characterized as follows: me- tatibial spurs less sharp than those of me- sotibiae; penis valve with short, straight, ventroapical spur; and tergites 7-10 or 8- 10 of female completely or almost com- pletely pubescent (except 7. fernaldii MacGillivray, fully pubescent from tergite 2). Except for species of the pectoralis group, species in other groups are often difficult to associate together in groups because char- acter states (other than color pattern) have not been discovered. However, adults of 7. VOLUME 97, NUMBER 1 53 ee oper ll eae ok i Nae 12 Figs. 1-12. Central serrulae of lancet of 1, Tenthredo appalachia; 2, T. masneri; 3. T. fernowi. Metatarsal pulvilli of 4, 7. appalachia; 5, T. leucostoma. Male genitalia of 6, 7. masneri; 7, T. fernowi (ventral view of half of genital capsule on right, lateral view of penis valve on left). Mesepisternal angle (posterior view) of 8, 7. masneri (showing head and thorax in anterior view); 9, T. appalachia; 10, T. fernowi; 11, T. mellicoxa; 12, T. rufopecta. 54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON appalachia and T. masneri are the only ones with an orange metepisternum on the ven- tral surface and with minute pulvilli. The color pattern of the legs and body of the male (coxae, trochanters, basal two-thirds of femora orange or whitish orange, tergites brown except medially on tergites 3-5) sup- ports this proposed relationship. Large pul- villi are seen in the earliest lineages of Ten- thredo as well as in other genera of the Tenthredinini; therefore, small pulvilli are a departure from the ground plan, which probably evolved several times in Tenthre- do, especially in species of most Eurasian groups. Elongation of the flagellum is usu- ally associated with reduction in size of the pulvilli. The lineage to which 7. appalachia belongs seems to be very rare in Eurasia. Only 7. procera Klug with a short spur on the penis valve seems to be related. Oth- erwise, the putative sister group is the atra group (piceocincta group in Ross 1951) in which the spur on the penis valve is long and curved up (similar to Fig. 6). Ecology.—The host is unknown. Adults have been recorded from about 500’ (Knowlton, Quebec) to 5200’ (Great Smoky Mts. Natl. Park, N.C./Tenn.) in deciduous forests. Flight dates are from May 21 to July 19. Twelve specimens were collected in the Fernow Experimental Forest in West Vir- ginia from May 21 to July 19 with most collected in May and June. Specimens were rather uniformly distributed throughout the four study watersheds, with 3 specimens from watershed 1, 3 from watershed 4, 4 from watershed 7, and 2 from watershed 13. Five were from forest-edge traps. The seven specimens from within the forest were all from traps in valleys and northerly-fac- ing slopes: traps 1, 1 specimen; traps 2, 4 specimens, and traps 3, 2 specimens. Tenthredo masneri Goulet and Smith, New SPECIES (Figs. 2, 6, 8) Diagnosis.— Adults are distinguished from those of other Nearctic species of Ten- thredo with minute pulvilli (length of pul- villus on first metatarsomere, 0.20-0.30 x length of second metatarsomere) and with microsculpture on the mesonotal median lobe by the following character combina- tion: ventral portion of metasternum com- pletely orange, clypeus white, and tergites 2-8 white along posterolateral margin. Female.— Length, 1 1.5—12.5 mm. Anten- na and head black; clypeus, labrum, basal 7, mandible, labium, and maxillary and la- bial palpi white; apical '3 mandible reddish; usually minute but occasionally large white spot on lower outer orbit and with minute white to brown spot on upper inner orbit. Thorax black with posterior corners and (in most specimens) ventral margin of prono- tum, tegula, spot (may be as large as pale portion of mesepisternum above mesepis- ternal spine) on lower posterior corner of mesepisternum, and metepisternum white; mesosternum and most of mesepisternum orange (white or whitish orange in some specimens) except for broad black stripe on upper '4 or less of posterior margin of me- sepisternum. Abdomen black; small white spot on lateral margin of tergite 1 extended along posterior margin, and tergites 2-8 each with narrow white band visible at least pos- terolaterally. Legs orange with extreme tip of meso- and metafemora, apical :—'2 me- tatibia, and metatarsus, black. Wings hya- line; veins and stigma black. Head in dorsal view narrower behind eyes (maximal distance between outer margins of eyes 1.10 maximal distance between outer margins of gena). Antennal crest mod- erately elevated with weakly developed an- gular projection behind it. Antennal flagel- lum moderately long: sixth flagellomere about 3 x longer than wide and first flagel- lomere 1.15—1.25 x as long as second fla- gellomere. Malar area near mandible with convex sculpticells and fine punctures (10- 15 wm in diameter); central portion of gena without microsculpture; remainder of head smooth. Mesepisternal spine obtusely an- gular (about 115°, Fig. 8). Most of thorax VOLUME 97, NUMBER 1 with meshes of microsculpture and sculp- ticells flat, but central area in upper half of mesepisternum, mesoscutellum anterome- dially, medial region of posttergite and mesonotal lateral lobe without microsculp- ture. Punctures on pectus and on meso- scutum fine (10-15 um in diameter), 25-30 um apart on median lobe and pectus, and 20-40 um apart on lateral lobe; punctures of mesoscutellum larger (25-30 um in di- ameter) and 25-50 um apart. Abdomen shining but with fine surface sculpture: sculpticells about 15 wm in length by 15-60 um in width, posterior margin of sculpticells clearly elevated and scale-like. Setae visible over most of tergite 1. Pubescence devel- oped over all of tergite 7 and dense on ter- gites 8-9, 40-SO um apart. Tarsal pulvilli minute (length of pulvillus of first metatar- somere 0.20—-0.28 x length of second me- tatarsomere) and narrow (width of pulvillus of first metatarsomere about '4 apical width of second metatarsomere) (as in Fig. 4). Ser- rulae of lancet as in Fig. 2. Male.—Length, 9.0 mm. Antenna and head black; clypeus and mouthparts white with apex of mandible reddish; lower '4 of outer orbit from mandible and minute spot on upper inner orbit whitish. Thorax mostly black dorsally, with large spot on lower pro- notum, posterior corner of pronotum, teg- ula, cervical sclerite, prosternum, central stripe on mesepisternum dorsal to angle, and metepisternum white; mesosternum to an- gle of mesepisternum pale orange. Abdo- men mostly black above with tergites 2-5 to 2-7 yellowish orange at side gradually becoming darker toward dorsal surface; lat- eral margin of tergite 1 narrowly white; pos- terior margin of each segment narrowly whitish; mostly orange ventrally with apical 2-3 sternites black. Legs pale orange to yel- low, fore- and midlegs more yellowish and hindleg more orange; upper surface of tip of each femur with black spot (may be lack- ing on profemur); metatibia brownish to dark orange; metatarsus black; outer surface of mesotibia and mesotarsus darker orange 55 than inner surface. Wings hyaline; veins and stigma black with vein R apical to stigma, amber. Head in dorsal view clearly narrower be- hind eyes (maximal distance between outer margins of eyes 1.20 maximal distance between outer margins of gena). Setae vis- ible over most of tergite 1. Metatarsus in dorsal view wider than in female: metatar- somere 2 about 4x longer than wide. Gen- italia as in Fig. 6; penis valve with !ong apical spine. Holotype. — Female, labeled ““QUE. Gat- ineau Pk, Luskville Falls, 5—22-VII-1988, J. Denis 300 m.” Deposited in the Canadian National Collection, Ottawa, Ontario, Can- ada. Paratypes (63 F, 5 M).—CANADA: NOVA SCOTIA: Cape Breton Highlands Natl. Park, 60°41’W-46°48'’N (1F); Cape Breton Highlands Natl. Park, 60°44’W-— 46°48'N (1F). ONTARIO: 7 mi. E Griffith (1F); Finland (1F). QUEBEC: Mont Albert (1F); Park Reserve (1F); Portneuf Co., St.- Augustin (1F); Berthierville (1M); St. Hi- laire (1F); Knowlton (1F); Ste. Agathe des Monts (1F); Mont Pinacle (1F); Montford (1F); Ste. Anne de Bellevue (1M, 3F); Ri- gaud (4F); King Mountain (summit) near Old Chelsea (2F); Gatineau Prov. Park, Luskville Falls (1F); Lac Roddick (1F). U.S.A.: MICHIGAN: Cheboygan Co. (1M). NEW HAMPSHIRE: Coos Co., First Con- necticut Lake (1F). NEW YORK: Ithaca (1F); Ludlowville (1M). NORTH CARO- LINA/TENNESSEE: Great Smoky Mts. Natl. Park, Indian Gap (1F). WEST VIR- GINIA: Tucker Co., Fernow Expt. Forest, E. M. Barrows, 11—20-V-91, trap 7-3 (1F); 21-30-V-91, trap 4-3 (1F), trap 7-3 (2F), trap 13-2 (1F); 31-V—9-VI-91, trap 4-5 (1F), trap 13-4 (1F); 10-19-VI-91, trap 4-2 (1F), trap 4-6 (1M), trap 7-4 (1F), trap 13-1 (1F), trap 13-3 (2F); 20-29-VI-91, trap 1-1 (1F), trap 1-3 (1F), trap 4-1 (1F), trap 4-2 (1F), trap 7-3 (1F), trap 13-1 (1F), trap 13-3 (1F); 30-VI-9-VII-91, trap 1-1 (1F), trap 1-5 (3F), trap 13-4 (1F); 10-19-VII-91, trap 1-5 (1F), 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON trap 7-1 (1F); 30-VII-8-VIII-91, trap 7-1 (1F); 30-V-8-VI-92, trap 13-3 (1F); 9-18- VI-92, trap 4-5 (1F); 19-28-VI-92, trap 13-5 (1F); 8-VII-92, on leaf of maple near trap 7-4 (1F); 9-18-VII-91, trap 4-5 (1F); 19- 28-VII-92, trap 4-3 (1F), trap 13-2 (1F); 29- VII-7-VIII-91, trap 13-2 (1F); 8-17-VIII- 92, trap 1-5 (1F); 18-27-VIII-92, trap 13-5 (1F); 24 km E Richwood, (1F). Deposited with the holotype; National Museum of Natural History, Washington, D.C.; Uni- versity of Montreal; Lyman Entomological Museum; and Carnegie Museum. Distribution.—This species is recorded from northwestern Ontario near the Man- itoba border to the Atlantic coast in Nova Scotia, and south along the Appalachian Mountains to Great Smoky Mountains Na- tional Park on the Tennessee-North Caro- lina border. Etymology.— We name this elegant spe- cies in honor of Dr. Lubomir Masner, a dear colleague, who, through his unabated en- thusiasm for field work, has provided us with numerous specimens and unusual spe- cies of sawflies. Dr. Masner also collected the northernmost record at the fringe of the boreal forest (Roddick Lake, Quebec). Discussion.—Adults of 7. masneri ap- pear to resemble those of 7. rufopecta and T. mellicoxa in the rufopecta group. How- ever, 7. masneri does not belong to this group, but is part of a very large lineage to which most North American species belong (see discussion under 7. appalachia). Fe- males of 7. masneri are near perfect color matches of 7. rufopecta. However, both sex- es of 7. rufopecta are differentiated from those of 7. masneri by the following fea- tures: antennae partly brownish (usually the undersurface); clypeus entirely or partly black (at least with some black and not en- tirely white); approximately upper half of mesepisternum black; minute white spots on lower outer eye orbit and upper inner orbit lacking; pro- and mesotarsal segments ringed with black at their apices; and me- sepisternal spine more obtuse (angle about 135°, Fig. 12). Both sexes of 7. mellicoxa are differentiated from 7. masneri by: a sharply defined white lateral stripe on the abdomen; posterior margins of abdominal segments narrowly but distinctly white; minute white spots on the upper inner orbit lacking; and the mesepisternal spine hardly suggested (rather flatly rounded without trace of an angle, Fig. 11). From 7. appa- lachia, probably its nearest species, 7. mas- neri 1s differentiated by the white posterior margin of the pronotum in dorsal view; white posterolateral margin of tergites 2-8; white posterior margin of tergite 1; lack of black at base of clypeus (except at side in some specimens); shorter flagellum (sixth flagellomere about 3x longer than wide); and presence of setae over at least base of tergite |. Ecology.—The host is unknown. Adults of the species have been recorded from sea level (Montreal, Quebec, region) to 5200’ (Great Smoky Mts. Natl. Park, N.C./Tenn.) mainly in deciduous forest habitats. Flight is recorded from May 11 to September 10. In the North (Ontario, Quebec, Nova Scotia and New England) most specimens were collected from mid-July to early August, but in the South (West Virginia) the peak flight is from mid-June to mid-July. Thirty-eight specimens were collected in the Fernow Ex- perimental Forest in West Virginia. They appeared to be rather uniformly distributed throughout the study watersheds. Traps in watershed | yielded 8 specimens; watershed 4, 8 specimens; watershed 7, 7 specimens; and watershed 13, 13 specimens. Most spec- imens were collected from traps near streams and on the southerly-facing slopes. Traps 1 yielded 7 specimens; traps 2, 5 specimens; traps 3, 11 specimens; traps 4, 3 specimens; and traps 5, 10 specimens. In the two years of the study, the flight period was rather long, beginning May 11 and ending August 27. Most specimens were collected June 10- 29, with numbers gradually decreasing to August. VOLUME 97, NUMBER 1 Tenthredo fernowi Goulet and Smith, NEw SPECIES (Figs. 3, 7, 10) Diagnosis.— Adults are distinguished from those of other Nearctic species of Ten- thredo with very small metatarsal pulvilli (length of pulvillus on first metatarsomere 0.15-—0.20x length of second metatarso- mere), and without meshes of microsculp- ture on dorsal surface of head, mesonotal median lobe, and mesepisternum (including pectus) by the following character combi- nation: clypeus black with sublateral white spots in female; tergites 2 and 3 orange and remaining tergites black; and mesepister- num (including pectus) black with white spot near mesocoxa or with stripe from meso- coxa to about middle of mesepisternum an- teriorly. Female.— Length, 10.5—12.0 mm. Anten- na and head black; clypeus mostly white laterally, black centrally; labrum, basal 7 of mandible, labium, and maxillary and labial palpi white; apical '4 of mandible reddish. Thorax black with posterior corner of pro- notum, tegula, short stripe or spot on lower posterior corner of mesepisternum, and me- tepisternum white. Abdomen black with small white spot on lateral margin of tergite 1 and 2nd and 3rd segments and anterior 3) of 4th sternite orange. Legs orange with extreme tip of meso- and metafemora, ex- treme apex of mesotibia, apical 4 of me- tatibia, and metatarsus black; apical 4 of mesotarsal segments usually blackish; basal Y4—'; of first metatarsomere orange. Wings hyaline; veins and stigma black with costa and vein R apical to stigma, amber. Head except near mandible, mesoscu- tum, mesoscutellum (except at side), me- sepisternum, pectus (except on outer half in some specimens), mesoscutellar appendage (except at side) and metascutellum without meshes of microsculpture, thus surface of these structures very bright. Gena near mandible with deeply impressed meshes of microsculpture and convex sculpticells, and 37) with some very fine punctures (about 15 um in diameter). Punctures on mesoscutum 10- 15 um in diameter; slightly denser on me- dian lobe (SO—75 um apart) than on lateral lobe (40-100 um apart); on mesoscutellum coarse (20-25 um in diameter) and 50-100 um apart. Mesoscutellar appendage with fewer than 8 punctures. Mesepisternum slightly extended outward and rounded without distinct medial angle (Fig. 10). Me- tascutellum with fewer than 10 setae. Ter- gites 2-8 with meshes of microsculpture variably transverse (length about 10 um and width 15-60 um); sculpticells flat and very slightly raised along posterior margin and scale like. Tarsal pulvilli minute, short (length of pulvillus of first metatarsomere 0.18-0.20x length of second metatarso- mere) and narrow (width of pulvillus of first metatarsomere about ' apical width of sec- ond metatarsomere) (as in Fig. 4). Serrulae of lancet as in Fig. 3. Male.—Length, 9.0-9.5 mm. Similar in coloration to female except clypeus white or with slight indication of black medially, no white spot laterally on tergite 1, 2nd and 3rd abdominal segments sometimes black- ish laterally, and apical 3 or more of me- tatibia black. Second metatarsomere in dor- sal view not widened, similar to female, about 5-6 x longer than wide. Genitalia as in Fig. 7; penis valve without apical spine. Holotype.—Female, labeled ““WEST VIRGINIA: Tucker Co., Fernow Expt. For- est, 31-V—9-VI-1991, E.M. Barrows” “Mal- aise trap 4-2.’ Deposited in the National Museum of Natural History, Washington, DG Paratypes (97 F, 5 M).—U.S.A.: MARY- LAND: Prince Georges Co., Patuxent Wild- life Center, 6—10-VII-90 (1F); Montgomery Co., Plummers Island, 24-V-75 (1F). NORTH CAROLINA: Great Smoky Mts. Natl. Park, 4-VII-62 (1F). VIRGINIA: Fairfax Co., near Annandale, Malaise trap, 27-V-3-VI-84 (1F); Falls Church, 24-VI-16 (1F), 1-VI (1M); Giles Co., Cold Spring, Va. Hwy 700 14-VI-75 (1F); Loudoun Co., 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bluemont, 6-V-13 (1F); Clarke Co., Univ. Va. Blandy Exp. Farm, 2 mi S Boyce, 19- 30-IV-90, trap 8 (2F), 1-13-V-90, traps 8, 9 (3F), 14-24-V-90, trap 8 (2F); Shenan- doah Co., Mount Jackson, 25-V-62 (IF); Shenandoah Co., Shenandoah Natl. Park, Compton Gap, 22-V-2-VI, 87 (1F); Shen- andoah Co., Shenandoah Natl. Park, Big Meadows, 14-VI-82 (1F). WEST VIRGIN- IA: Same data as holotype, except for dates and trap numbers: 21-30-V-91, trap 1-4 (1F), trap 4-1 (1F), trap 4-4 (1F), trap 7-1 (3F), trap 7-2 (1F), trap 7-3 (1F), trap 7-4 (2F), trap 13-3 (2F), trap 13-4 (1F); 31-V- 9-VI-91, trap 1-4 (1F), trap 1-5 (1F), trap 4-3 (1F), trap 4-4 (1F), trap 4-6 (1F), trap 7-2 (1F), trap 7-3 (2F), trap 7-6 (1F), trap 13-3 (1F), trap 13-4 (2F); 10-19-VI-91, trap 1-2 (2F), trap 4-1 (1F), trap 4-3 (1F), trap 4-6 (1F), trap 7-3 (1F), trap 7-6 (1F); 20- 29-VI-91, trap 1-1 (IF), trap 4-2 (1F), trap 4-3 (1F), trap 7-3 (1F), trap 13-4 (1F); 30- VI-9-VII-91, trap 1-1 (1F), trap 4-3 (1F); 10-19-V-92, trap 1-1 (1F); 20-29-V-92, trap 1-2 (1F), trap 4-1 (1F), trap 4-2 (1F, 1M), trap 4-3 (2F), trap 7-3 (1F, 2M), trap 13-2 (1F), trap 13-4 (1M), trap 13-6 (1F); 30-V- 8-VI-92, trap 4-3 (2F), trap 7-2 (1F); 9-18- VI-92, trap 1-1 (1F), trap 1-2 (1F), trap 4-3 (1F), trap 4-4 (1F), trap 7-3 (1F), trap 7-5 (1F), trap 13-3 (1F); 19-28-VI-92, trap 1-3 (1F), trap 4-2 (2F), trap 4-3 (2F), trap 13-4 (1F); 29-VI-8-VII-92, trap 4-1 (3F), trap 4-3 (1F), trap 7-1 (1F), trap 13-4 (1F); 9- 18-VII-92, trap 1-1 (1F), trap 4-1 (2F), trap 4-2 (1F), trap 7-1 (1F), trap 7-4 (1F), trap 13-3 (1F), trap 13-4(1 F); Cranberry Glades, 3-4-VI-55 (2F). Deposited with the holo- type; Museum of Comparative Zoology; Florida State Collection of Arthropods; Snow Entomological Museum; and the Ca- nadian National Collection, Ottawa. Distribution.—This species is recorded from Maryland, Virginia, West Virginia, and North Carolina. Etymology.—This species is named for Bernhard E. Fernow, a German-born for- ester who pioneered scientific forestry in the United States (Anonymous 1987). The Fer- now Experimental Forest, the type locality and area from which most of the type ma- terial was collected, was named in his hon- or. Discussion. — Tenthredo fernowi is a slen- der species with very small pulvilli (similar to Fig. 4). The orange color of the second and third abdominal segments, white spot laterally on the clypeus of the female, and mostly orange legs is a color combination not known in other North American Ten- thredo. The clypeus of the male may be mostly white, but lack of meshes of mi- crosculpture on the head, mesepisternum, pectus, and mesonotal median lobe will aid in its separation. Coloration is rather stable, but a few mi- nor variations occur. The white area at the lower posterior portion of the mesepister- num may vary from a small spot to a stripe extending anteriorly for about half the length of the mesepisternum; the third abdominal segment may be all orange or slightly black- ish posteriorly; the fourth sternite may be all black to mostly orange; and the lateral white spots on the clypeus may be large or small in the female and the clypeus mostly or all white sometimes with a medial black- ish area in the male. Based on Ross’ (1951) classification, 7. fernowi would belong in the rufopecta group. Ross created groups for the Nearctic species of the genus in the 1951 Hymenoptera Cat- alog, but, to our knowledge, did not char- acterize them elsewhere. A study in progress by the senior author based on numerous structures in both sexes does not support some of the species associated with the ru- fopecta group. Tenthredo repleta Mac- Gillivray (type not studied yet) is, if cor- rectly interpreted, a member of the grandis group (median lobe of mesoscutum clearly with microsculpture, mesepisternum dense- ly punctate, antennal flagellum flattened and apical joints white), and 7. rurigena is more closely allied to Ross’ secunda or semirufa groups (metatibial spurs less sharp than me- VOLUME 97, NUMBER 1 sotibial ones, male penis valve with ventro- apical short and straight spine). This leaves T. rufopecta and T. mellicoxa in the rufo- pecta group. In North America, the rufo- pecta group is nearest to species related to T. grandis (Norton) (not Ross’ grandis group, excluding 7. colon Klug) and 7. ruma MacGillivray. However, the rufopecta group appears nearest to Euroasiatic lineages. The rufopecta group is characterized as follows: microsculpture lacking (no traces of mesh- es) on dorsal surface of head and mesonotal median lobe; pulvilli very small (length of pulvillus of first metatarsomere 0.15-0.20 length of second metatarsomere) and nar- row (width of pulvillus on first metatarso- mere about one-third that of second me- tatarsomere); at least tergites 6-10 completely or almost completely pubescent in female; penis valve without ventro-apical spine; metatibial spurs as sharp as those of mesotibia; antennal crest developed but low, hardly or not angularly produced between level of median ocellus and antennal socket; and at least mesotarsomeres 1-4 darkened on apical 0.20-0.40 (thus ringed with brown or black at apex). Except for color characters the above def- inition applies closely to some species in Eurasia such as 7. hilalis Smith. In North America, the group as here defined consists of above three species and 7. nimbipennis. Within this group. 7. fernowi is nearest to T. rufopecta. Adults of T. fernowi share the following derived character states with 7. rufopecta: first flagellomere barely longer than second; mesepisternum and pectus smooth (shared also with 7. nimbipennis); clypeus with white spots sublaterally (shared also with 7. nimbipennis); and complete lack of angular projection between antennal crest and level of median ocellus. Adults of 7. fernowi are unusual in that the mesepister- nal spine is hardly suggested and that the first metatarsomere of the male is not wid- ened. In these two features, adults of 7. fer- nowi match those of the Eurasian species, T. hilalis. 39 Ecology.—The host is unknown. Adults of this species have been recorded from sea level (Patuxent, Maryland) to 4200’ (Great Smokies Mts. Natl. Park, N.C./Tenn.) in broadleaf forests. Flight is recorded from April 19 to July 18. Most specimens were collected from mid-May to mid-June and were still common until mid-July. Eighty- three specimens were taken in traps in the Fernow Experimental Forest in West Vir- ginia in 1991 and 1992. This was the most commonly collected species of Tenthredo in the forest. The species appeared to be uni- formly distributed throughout the study wa- tersheds. Five of the specimens were taken from forest-edge traps. Traps in watershed 1 yielded 13 specimens; watershed 4, 30 specimens; watershed 7, 21 specimens; and watershed 13, 14 specimens. Among the traps within the watersheds, traps | yielded 18 specimens; traps 2, 15 specimens; traps 3, 27 specimens; traps 4, 16 specimens; and traps 5, 2 specimens. Adults appeared to be most common in the valleys and on the northerly-facing or shaded slopes. They were least common in the high traps on the south- facing slopes. Flight period for the two years combined was from May 10 to July 19 (Fig. 13). Most specimens were collected May 21- 30 with numbers steadily decreasing to July. Specimens collected at the University of Virginia Blandy Experimental Farm, Clarke Co., Va., were all from one trap which was within a dense 85-year-old, oak-elm-hick- ory woodlot. After three years of collecting in the same general area in other habitats, no additional specimens were obtained. Thus, this species appears to be associated with the dense broadleaf forests. Dolerus (Achaetoprion) klokeorum Goulet and Smith, NEw Species (Fig. 14) Diagnosis.— Females are distinguished from other Nearctic species of Dolerus by the following combination of character states: punctures of mesepisternum large (100-150 um in diameter); mesonotum 60 25 20 15 of specimens 10 number PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0 Ds an fo. i=) an fo. n n a an on Uf a mi en = a ih ve ) ~ > =) So = a °o = FF Q ~ Sia) GG Ss =| Q ss > > oO > > ro) > a collection date Fig. 13. Flight period for Tenthredo fernowi, Fernow Experimental Forest, West Virginia, 1991-1992. Num- ber of specimens captured per collection date. black only on lateral lobes; abdomen and base of sheath (valvifer 1 and 2) completely orange; and wings uniformly lightly infus- cated. Female.— Length, 7.5—8.5 mm. Antenna, head, and mouthparts black. Thorax orange with mesosternum, lower '4—'3 of mesepis- ternum, and most of mesonotal lateral lobes except posterior corners black. Abdomen orange with cercus black and apical segment (valvula 3) of sheath black laterally. Legs black. Wings moderately, uniformly infus- cated; veins and stigma black. Surface microsculpture generally lacking except on outer half of pectus, head (be- tween punctures dorsally), side of tergites 7-9, last segment of sheath (valvula 3), cox- ae, and femora where flat sculpticells are outlined. Clypeal emargination deep, sub- equal to medial length of clypeus. Punctures on postocular area on both sides of post- ocellar region as far as upper '4 of gena densely punctate (a little less so near post- ocellar region) and about 80 um in diameter. Punctures on mesepisternum large (100-150 um in diameter) and on pectus very small (20 um in diameter) and 30-80 um apart except on oblique furrow (about 50 um in diameter). Mesonotal lateral lobe without large punctures laterally. Mesoscutellar ap- pendage about 175 um in length medially. Distance between cenchri about 1.3 x width of acenchrus. Setae of mesepisternum about 100 um in length. Cercus about 3 x wider than long. Sheath in posterior view widely concave, in dorsal view with longest setae regularly curved, forming an angle of about 10°; ventral margin of last segment of sheath (valvula 3) regularly curved and forming an angle of about 45° with dorsal margin. Lan- cet as in Fig. 14; annuli each with one small spine and without seta-like sculpticells; ser- rulae except basal two, each with 4 or 5 coarse posterior subbasal teeth. Male. — Unknown. Holotype.— Female, labeled ““VIRGIN- IA: Louisa Co., 4 mi. S. Cuckoo, 12-25-IV- 88, J. Kloke & D.R. Smith, Malaise trap.” VOLUME 97, NUMBER 1 61 Fig. 14. Female lancet of Dolerus klokeorum. Deposited in the National Museum of Nat- ural History, Washington, D.C. Paratypes (4 F).—U.S.A.: VIRGINIA: Same locality as holotype, 29-III-11-IV-88 (1F), 5-25-IV-89 (1F); Essex Co., 1 mi SE Dunnsville, 12—29-IV-91, David R. Smith, Malaise trap 10 (1F); Fairfax Co., Spring- field, 29-IV-1973 (1F). Deposited with the holotype and in the Canadian National Col- lection, Ottawa. Distribution.— Known only from Virgin- la. Etymology.—This species is named for Jack and Beth Kloke, who have kindly al- lowed the junior author access to their prop- erties in Virginia for field work. Discussion.— Females of this species re- semble those species of Achaetoprion with large punctures on mesepisternum (couplets 1-4 of Goulet’s 1986 key), especially those of the Dolerus abdominalis (Norton) and D. eurybis Ross lineage defined by the follow- ing shared derived character states: clypeus deeply emarginate (shallowly emarginate in other Achaetoprion and most Dolerini) and punctures large on mesepisternum (small in other Achaetoprion except D. versus Nor- ton). Females of D. eurybis are especially similar to D. klokeorum in color, but D. eurybis has the basal half of the wings deeply infuscated and the apical half hyaline; the longest setae of the sheath in dorsal view regularly curved and directed posteriorly (at about 10°); and broad, laterally projecting winglike processes on the annuli of the api- cal half of the lancet. Dolerus abdominalis has the thorax entirely black, except some- times the metapleuron, the sheath entirely black with apex convex, and broad, laterally projecting winglike processes on the annuli of the apical half of the lancet. Despite sim- ilarities of the above external character states with other species, there is a major differ- ence in the ovipositor structure. In D. klo- keorum each annulus of the lancet has a spurlike process, while in D. eurybis and D. abdominalis it consists of a winglike process on the apical annuli. If this very unusual winglike process on the apical annuli of the lancet is derived (see Goulet 1986, figures 47 and 321, notes on affinities, p. 88), then D. abdominalis and D. eurybis would not be closely related to D. klokeorum which has only a spurlike process on the annuli as in D. nortoni Ross, D. mimus Goulet, D. tacoma Goulet, and D. neoagcistus Mac- Gillivray (see Goulet 1986, figures 310, 312, 314 and 317). The spurlike process repre- sents an earlier stage leading to the evolu- tion of the winglike process. The relation- ship of species with a spurlike process on the annuli is unresolved. It is unlikely that winglike processes of the type described in Goulet (1986) would evolve independently. Thus, the similarities in external characters noted above between D. klokeorum and D. eurybis and D. abdominalis may be con- vergent. 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ecology.—Specimens were from traps in lowlands next to streams or drainages, areas frequently flooded during heavy rainfall. Trap 10 in Essex Co. traversed a small in- termittent stream in a woodlot; the stream bed was moist in early spring but dry most of the summer. This species may feed on grasses or sedges in such stream beds or other similar seepage areas. Equisetum, a common host for other groups of Dolerus, was not present in these habitats. The five females were collected from March 29 to April 29. ACKNOWLEDGMENTS This study was supported in part by Co- operative Agreement 42-646, U.S. Forest Service (to E. M. Barrows) and Georgetown University. We also thank Mr. and Mrs. J. G. Kloke for allowing field work on their property in Louisa and Essex counties, Vir- ginia, and Edward F. Connor, Michael A. Bowers, and Christopher F. Sacchi, for al- lowing field work at the University of Vir- ginia Blandy Experimental Farm and State Arboretum of Virginia in Clarke Co., Vir- ginia. Robert Overholser, art intern with the Systematic Entomology Laboratory, pre- pared Fig. 14. Some of the study specimens were loaned by officials of the University of Montreal, Montreal, Quebec; Lyman En- tomological Museum, MacDonald College, Ste. Anne de Bellevue, Quebec; Carnegie Museum, Pittsburgh, Pennsylvania; Muse- um of Comparative Zoology, Harvard Uni- versity, Cambridge, Massachusetts; Florida State Collection of Arthropods, Gainesville, Florida; and the Snow Entomological Mu- seum, University of Kansas, Lawrence. We thank the following for review of the manu- script: Edward M. Barrows, Georgetown University, Washington, D.C.; Gary A. P. Gibson, Biological Resources Division, Ag- riculture Canada, Ottawa; and Raymond J. Gagné and David A. Nickle, Systematic En- tomology Laboratory, U.S.D.A., Washing- ton, D.C. LITERATURE CITED Anonymous. 1987. Forest Research: Fernow Exper- imental Forest. United States Department of Ag- riculture, Forest Service, Northeastern Forest Ex- periment Station NE-INF-75-87, 12 pp. Goulet, H. 1986. The genera and species of the Ne- arctic Dolerini (Symphyta: Tenthredinidae: Selan- driinae): Classification and phylogeny. Memoirs of the Entomological Society of Canada, No. 135, 208 pp. Griffith, M. B. and S. A. Perry. 1992. Plecoptera of headwater catchments in the Fernow Experimen- tal Forest, Monongahela National Forest, West Virginia. Proceedings of the Entomological Soci- ety of Washington 94: 282-287. Ross, H. H. 1951. Suborder Symphyta (= Chalas- togastra), pp. 4-89. Jn Muesebeck, C. F. W., K. V. Krombein, and H. K. Townes, eds., Hyme- noptera of America North of Mexico, Synoptic Catalog. United States Department of Agriculture, Agriculture Monograph No. 2, 1420 pp. Smith, D. R. 1979. Suborder Symphyta, pp. 3-137. In Krombein, K. V., et al., eds., Catalog of Hy- menoptera in America North of Mexico, Volume 1, pp. 1-1198. Smithsonian Institution Press, Washington, D.C. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 63-70 MIRABILIS-FEEDING HELIODINES (LEPIDOPTERA: HELIODINIDAE) IN CENTRAL ILLINOIS, WITH DESCRIPTION OF A NEW SPECIES T. HARRISON AND S. PASSOA (TH) Department of Entomology, University of Illinois, Urbana, Illinois; (SP) USDA/ APHIS/PPQ, Reynoldsburg, Ohio. Abstract. —We report finding in central Illinois an example of simultaneous, sympatric use of Mirabilis nyctaginea (Michx.) Sweet (Nyctaginaceae) by larvae of four species of Heliodinidae. The insects involved are Heliodines tripunctella W\sm., H. nyctaginella Gibson, H. ionis Clarke, and an undescribed Heliodines species. We provide a description of the new moth and keys to adults of the four Illinois species. Key Words: Members of the genus Heliodines are very small, primarily diurnal moths; most have golden-orange forewings adorned with raised patches of metallic lead-gray scales. The 22 described species are predominantly Nearc- tic or Neotropical, the exceptions being sin- gle European and Australian representa- tives. Larval host plants have been published for five species: Heliodines roesella (L.) on Chenopodiaceae (e.g. Stainton 1854, Em- met 1985); Heliodines quinqueguttata Wal- singham (1897) on Portulacaceae; Heliod- ines extraneella Walsingham (1881) on Onagraceae (Braun 1925); and two species, Heliodines nyctaginella Gibson (1914) and Heliodines ionis Clarke (1952), on Nycta- ginaceae. Heppner and Duckworth (1983) listed 1 1 North American species of Heliodines north of Mexico, and Hodges (1983) noted that the moths are seldom collected, probably because they are diurnal. A documented characteristic of heliodinid larval biology 1s the occurrence of more than one species in close proximity to each other (Wester 1956), sometimes with larvae feeding simulta- neously on the same individual plant. We Heliodinidae, Heliodines, new species, Nyctaginaceae, Mirabilis report finding in central Illinois a striking example of sympatric, simultaneous use of Mirabilis nyctaginea (Michx.) Sweet (Nyc- taginaceae) by four heliodinid species: He- liodines tripunctella Walsingham (1892), a leaf miner; H/. nyctaginella, a flower feeder and upperside leaf skeletonizer; H. ionis, a stem borer; and an undescribed Heliodines species (underside leaf skeletonizer). As a preliminary step in our study of this insect- plant interaction, we provide a description of the new moth and keys for identifying adults of the four Illinois species. Heliodines cliffordi Harrison & Passoa, NEw SPECIES (Figs. 1-3, 6-8, 12-13, 16) Ovum (Fig. 1): Whitish, somewhat flat- tened. Laid on the underside of a leaf. After hatching, the larva does not eat the remains of the egg. Larva (Fig. 2): On Mirabilis nyc- taginea in Illinois; also known from Mirab- ilis longiflora L. in Arizona (our comments refer to Illinois specimens). One to several larvae on a single leaf, usually on the lower leaves of the plant. There are four larval instars. Upon hatching, the larva moves a 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. larva, 3, Larval damage to Mirabilis nyctaginea, with several larvae on underside of leaf. Heliodines cliffordi. \, Ovum. 2, Mature small distance from the remains of its egg and burrows into the leaf. First two instars spent as a leaf miner, third and fourth in- stars as a leaf skeletonizer, feeding beneath a flat sheet of fine white silk on the underside of the leaf, eating all but the upper epidermis in an irregular blotch pattern (rarely, feeding on upperside of leaf). The “window” of damage (Fig. 3) is conspicuously visible on the upper surface of the leaf. Frass is de- posited outside the feeding area and persis- tently adheres to the external surface of the silken sheet. General color of mature larva pale green, with no contrasting pinacula or other external markings; cuticle translucent, the aorta faintly visible as a dark dorsal midline; head blackish brown throughout larval life; prothoracic shield concolorous with body during first three instars, blackish brown and divided longitudinally at the midline during fourth instar. A single SV seta on segment AY in allinstars. Pupa: Ob- tect; flattened dorsally, keeled laterally. No dorsal transverse rows of spines. No co- coon, pupation occurring among a slight, loose mass of silk from which the pupa is not protruded at eclosion. Adult (male and female): //ead: Labial palpus: First and second segments dull yellow, the second slightly darker dorsally toward apex; third segment dull yellow suffused with light brownish gray. Antenna: Shining dark gray with violet reflections; apical five segments white. Face and head capsule: Smoothly scaled, shining gray; reflections brassy, greenish, or violet with differing angles of incident light; ocellus large and prominent; occipital scale band contrastingly dull yel- low. Thorax: Shining gray as for head. Forewing: Length 4.5—-5.0 mm. Ground col- or bright golden orange. Five costal, raised, metallic lead-gray spots, designated Cl (basal) to C5 (apical), at approximately 0.07, 0.19, 0.35, 0.50, and 0.67 (numbers repre- sent respective average distances from wing base to centers of spots, expressed as per- centages of total wing length, n = 5); spots C2 and C3 extend farther toward posterior margin of wing than do the other costal spots; C5 appearing triangular due to a shining- gray patch that narrows as it extends along costa from apical edge of the spot. Three dorsal, raised, metallic lead-gray spots, des- ignated D1 (basal) to D3 (apical), at ap- proximately 0.27, 0.44, and 0.60; D1 and D3 extend farther toward costal margin of wing than does D2. Anterior margin from wing base to spot C2, and posterior margin from wing base to D1, broadly margined with black; between these anterior and pos- terior areas, a discal, longitudinally orient- ed, yellowish-orange patch extends from near wing base to a point between C2 and D1, where it blends into the golden-orange VOLUME 97, NUMBER 1 Figs. 4-9. Male genitalia. 4, Heliodines tripunctella (ventral aspect, spread and flattened, aedeagus removed). 5, H. nyctaginella (lateral aspect, aedeagus removed). 6, H. cliffordi, valve (lateral aspect). 7, 1. cliffordi, aedeagus (lateral aspect). 8, 1. cliffordi (lateral aspect, aedeagus removed). 9. H. ionis (lateral aspect, aedeagus removed). 66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 14° Figs. 10-14. Female genitalia. 10, Heliodines tripunctella (ventral aspect). 11, H. ionis (ventral aspect). 12% H. cliffordi, detail of signum. 13, H. cliffordi (ventral aspect). 14, H. nyctaginella (ventral aspect). VOLUME 97, NUMBER 1 ground; amount of black between spots C2 and C3 variable, always with at least a wide black anterior margin but in some speci- mens filled with black to the height of the two spots. Other spots margined with black to a varying degree, in most specimens at least finely so. Orange area extending farther apically on anterior margin than on poste- rior; costoapical extremity of orange area often with a small yellowish-orange patch. Wing distad of orange area very dark shin- ing gray, with one small black patch in basal region of this area near costa and one from apical margin of spot D3. A discal, metallic lead-gray spot, rather small and only slightly raised, at approximately 0.77; the apical half of this spot lies in the outer gray area of the wing, the basal half interrupts the distal edge of the orange area. Fringe dark gray. All veins separate, including R, and R,. Hind- wing: Length 4.0-4.5 mm. Lanceolate, very dark gray; fringe dark gray. Rs and M, sep- arate, CuA, and CuA, present; remnants of two anal veins persistent. Legs: Foreleg with coxa shining gray, cream colored at apex; femur shining gray on outer surface from body, cream colored on inner; tibia shining gray laterally, cream colored medially; tar- sus shining gray. Midleg colored as foreleg but tibia ringed with cream color at apex. Hindleg colored as midleg but apex of femur cream colored, and tibia with two dark-gray bands, one from tibial base to first pair of spurs, the other from two-thirds tibial length to edge of apical cream-colored band; spurs shining gray on outer surfaces from tibia, cream colored on inner; tibia smooth, not spined or otherwise modified. Abdomen: Shining gray, somewhat darker than thorax. Genitalia: As illustrated and described be- low in the key. Voltinism: Probably four complete gen- erations per year; mature larvae appear in central Illinois from late May to mid-June, again from mid- to late July, and at all times from early August through mid-September. Generations become asynchronous so that by August and afterward larvae of all instars 67 Figs. 15-18. tripunctella. 16, H. cliffordi. 17, H. nyctaginella. 18, H. ionis. Wings of adult moths. 15, Heliodines can be found at once. Viable pupae from mature larvae collected 17 September 1993 had not produced adults by mid-November 1993; this suggests that the insect overwin- ters as a pupa. Distribution: In addition to the type lo- 68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cality, Charleston, Coles County, Illinois, this species is represented by specimens from Putnam, Mason, and Champaign counties in Illinois [in the collection of the Illinois Natural History Survey, Champaign (INHS)]; and from the states of New Mexico (INHS), Iowa, Kansas [in the collection of the United States National Museum of Nat- ural History, Washington, D.C. (USNM)], and Arizona (reared from Mirabilis longi- flora by R. Wielgus; in the collection of D. Wagner, Storrs, Connecticut). Data for all Illinois specimens are entered into the IIli- nois State Lepidoptera Data Base, Spring- field. Holotype male: Collected as larva on Mi- rabilis nyctaginea, USA: Illinois, Coles County, Charleston, T12N, R9E, NW 4 Sec. 11, 14-VI-1992, T. Harrison. Iss. 30-VI- 1992 (USNM). Allotype female: Same data as for holotype except collected 8-VI-1990, iss. 22-VI-1990 (USNM). Paratypes: De- posited in the following collections: USNM; INHS; Illinois State Museum, Springfield; collection of S. Passoa, Reynoldsburg, Ohio; University of California, Berkeley; Cornell University, Ithaca, New York; American Museum of Natural History, New York, New York; Florida State Collection of Ar- thropods, Gainesville; Academy of Natural Sciences, Philadelphia, Pennsylvania; Ca- nadian National Collection, Ottawa; The Natural History Museum, London, En- gland. Etymology: This species is named for the senior author’s son, Clifford R. Harrison. Discussion: Heliodines cliffordi is similar to H. nyctaginella and H. ionis but displays darker and more extensive gray and black scaling than do those two species. The wing venation is the same among the three. Gen- italia differ as specified below in the key. In all three species, there is one SV seta on larval segment A9. There are four larval instars in H. cliffordi, in contrast to five in- stars in H. nyctaginella and non-overwin- tering H. ionis, and six instars in overwin- tering H. ionis (Wester 1956). Heliodines tripunctella differs from H. nyctaginella, H. ionis, and H. cliffordi in that veins R, and R, in the forewing and Rs and M, in the hindwing are stalked; CuA, is absent in the hindwing; and there are two SV setae on larval segment A9. Other characters that distinguish H/. tripunctella from the other three Illinois species are mentioned in the keys. For field identification of mature larvae of the two leaf-skeletonizing species, H. clif- fordi is pale green with a blackish-brown head and is found on the underside of the leaf, while H. nyctaginella is deep reddish brown with a tan head, and it skeletonizes the upper surface of the leaf. As for mature larvae of the other two Illinois species, H. ionis is yellowish white and is a stem borer; H. tripunctella is dull green with dark tho- racic markings and is a leaf miner through- out larval life. Key TO ADULTS OF ILLINOIS MIRABILIS-FEEDING HELIODINES, BASED ON GENITALIA Ler IMales = soasys he tS aeet aycaes ae eon 2 WWepEemalesa tc. ne eee nae 5 2. Each lobe of socius bifid; saccus and aedeagus relatively short, saccus less than three times as long as tegumen (Fig. 4) ........ tripunctella 2'. Each lobe of socius single, not bifid; saccus and aedeagus relatively elongate, saccus great- er than three times as long as tegumen ..... 3 3. Each lobe of socius at least five times as long as its maximum width, acuminate apically; tegumen small (Fig. 5)) ............ nyctaginella 3’. Each lobe of socius much less than five times as long as wide, bluntly rounded apically; te- gumenlarge 4s. o-oo cnc eee ce 4 4. Valve composed of broad, triangular basal re- gion and narrow, slightly curved apical region (FI8S:105: 8) ita: an ee Ce eee cliffordi 4'. Valve rather uniformly narrow along entire length, expanding slightly in width from base to apex (Fig. 9) ionis 5. Ductus bursae much less than 20 times as long as wide, its junction with corpus bursae oc- curring posteriad of anterior margin of seg- ment A7; signum a uniformly sclerotized patch with one inwardly protruding spinelike pro- cess (Fig. 10) tripunctella 5’. Ductus bursae at least 20 times as long as its VOLUME 97, NUMBER 1 tie I width at narrowest point, its junction with cor- pus bursae occurring well anteriad of anterior margin of segment A7; signum not as above.. 6 Signum divided into two sclerotized plates that lie on opposite surfaces of corpus bursae; duc- tus seminalis arising from ductus bursae, at a point posteriad of corpus bursae (Fig. 11) ... ionis . Signum a single elongate-rhomboid plate lying lengthwise along side of corpus bursae (Fig. 12); ductus seminalis and ductus bursae orig- inating very near each other, both from pos- terior end of corpus bursae Ductus bursae uniformly narrow from ostium to corpus bursae; a prominent accessory cor- pus bursae arising from near anterior end of corpusibursae (E1913) maaan eae eee cliffordi Ductus bursae widened near ostium (Fig. 14); no accessory corpus bursae ........ nyctaginella Key To ADULTS OF ILLINOIS MIRABILIS-FEEDING HELIODINES, BASED ON EXTERNAL CHARACTERS Dorsum of abdomen predominantly orange; at rest, living moth with metathoracic legs held elevated above body; forewing with a nearly uniformly wide, shining-gray border extending unbroken around wing margin from distal edge of spot C2 to a point slightly distad of spot D1; each forewing with three costal and one dorsal raised, lead-gray spots (Fig. 15) ..... tripunctella Dorsum of abdomen dark gray or black, not orange; metathoracic legs not held elevated; no continuous shining-gray border around margin of forewing; forewing always with more than four lead-gray spots Occipital scale band dull yellow, contrasting with shining-gray head and thorax; forewing with five costal, raised, lead-gray spots; distal margin of orange area of forewing interrupted by discal lead-gray spot; anterior margin of forewing from base to spot C2 and posterior margin from base to spot D1 broadly mar- gined with black, the discal longitudinal patch between these black areas yellowish orange (Fig. NG) FARE AN ents Aer ees Perched towne Th cliffordi . Occipital scale band concolorous with thorax or contrastingly black; six costal, lead-gray spots on forewing; no discal lead-gray spot, distal margin of orange area on forewing an unbro- ken curve; basal area of forewing narrowly or not at all margined with black; orange ground unicolorous, no contrastingly yellowish areas RIED YH Patch clon y/o at Be “AVS YS GTA ok Hyped 3 Posterior margin of forewing distinctly lined with black from base to basal edge of spot D1 69 (Fig. 17); occipital scale band shining gray, concolorous with head and thorax; antennae Wunitve (uhopol Seong mcoceneseued soon nyctaginella 3’. Posterior margin of forewing not lined with black at base (Fig. 18); occipital scale band black, contrasting with head and thorax; an- fennacientirelysGark£ ax, socasc/aten 5 nie ionis ACKNOWLEDGMENTS We would like to thank the following peo- ple for providing assistance with this pro- ject: for scanning electron micrographs of the ovum of H. cliffordi, Kathy Wolken and John Mitchell, The Ohio State University, Columbus; for comments on the manu- script, Dr. M. R. Berenbaum, Department of Entomology, University of Illinois, Ur- bana, and two anonymous reviewers; for the loan or gift of photo slides and/or specimens of Heliodinidae, and for helpful and infor- mative correspondence regarding these in- sects, all of the following: Dr. R. W. Hodges, USDA, ARS/SEL, Washington, D.C.; Dr. Y. Arita, Meijo University, Nagoya, Japan; Dr. C. W. Baker, Boise State University, Boise, Idaho; Ms. K. Methven, Illinois Nat- ural History Survey, Champaign; Dr. J. A. Powell, University of California, Berkeley; M. Schaffer, The Natural History Museum, London, England; Dr. D. Wagner, Univer- sity of Connecticut, Storrs; Dr. L. Wolfe, Hebrew University, Jerusalem, Israel; and, finally, we owe special thanks to Mr. Y.-F. Hsu, University of California, Berkeley. LITERATURE CITED Braun, A. F. 1925. Microlepidoptera of northern Utah. Transactions of the American Entomological So- ciety 51: 183-226. Clarke, J. F.G. 1952. A new heliodinid from Illinois (Lepidoptera, Heliodinidae). Proceedings of the Entomological Society of Washington 54: 138-139. Emmet, A. M. 1985. Heliodinidae. Jn Heath, J. and A. M. Emmet, eds., The Moths and Butterflies of Great Britain and Ireland, Volume 2. Cossidae- Heliodinidae. Harley Books, 460 pp., 14 pls. Gibson, A. 1914. A new elachistid from Manitoba. The Canadian Entomologist 46: 423-424. Heppner, J. B. and W. D. Duckworth. 1983. Heliod- inidae. Jn Hodges, R. W., T. Dominick, D. R. Davis, D. C. Ferguson, J. G. Franclemont, E. G. 70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Munroe, and J. A. Powell, eds., Check list of the Lepidoptera of America North of Mexico. Wedge Entomological Research Foundation, 284 pp. Hodges, R. W. 1983. In Hodges, R. W., T. Dominick, D. R. Davis, D. C. Ferguson, J. G. Franclemont, E. G. Munroe, and J. A. Powell, eds., Check list of the Lepidoptera of America North of Mexico. Wedge Entomological Research Foundation, 284 pp. Stainton, H. T. 1854. Insecta Britannica. Lepidop- tera: Tineina. John Edward Taylor, 313 pp., 10 pls. Walsingham, Lord (Thomas de Grey). 1881. On some North American Tineidae. Proceedings of the Zoological Society of London: 301-324, 2 pls. 1892. Steps towards a revision of Chambers’s index, with notes and description of new species. Insect Life 4: 384-385. 1897. Revision of the West-Indian Micro- lepidoptera, with descriptions of new species. Pro- ceedings of the Zoological Society of London: 54— 183. Wester, C. 1956. Comparative bionomics of two spe- cies of Heliodines on Mirabilis (Lepidoptera, He- liodinidae). Proceedings of the Entomological So- ciety of Washington 58: 43-46. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 71-76 ATACTOCORIS PERNERI, N. SP., A NEW APTEROUS CARVENTINAE FROM JAMAICA (HETEROPTERA: ARADIDAE) ERNST HEIss Entomology Research Group, Tiroler Landesmuseum, 2A J. Schraffl-Strasse, A-6020 Innsbruck, Austria. Abstract. —The first record of the monobasic genus Atactocoris since its description 30 years ago is reported and the new species A. perneri, is described. A habitus of both sexes and the male genital structures of 4. perneri are figured. Additional data are given for the female holotype and only known specimen of the type species, A. farri Kormilev, 1964. Key Words: The genus Afactocoris was created by Kormilev (1964) to contain the species, A. farri Kormilev from Jamaica. The descrip- tion was based on a single female and no further specimens have been reported since. I have recently had the opportunity to col- lect additional material that proved to be- long to a new species which is described and figured below. The holotype of A. farri, preserved in the collection of the Institute of Jamaica in Kingston is redescribed. Measurements are given in millimeters. Atactocoris perneri Heiss NEw SPECIES Figs. 1-5, 7-12 Diagnosis: Distinguished from the only known species of this genus, 4. farri, by smaller size, shorter antennae and antennal segment I distinctly shorter than width of head across eyes (longer in A. farri). In the female it is further separated by shorter pos- terolateral projections of tergite VII (Figs. 4, 6). Description: Male. Apterous; body elon- gate, smooth and shiny beneath incrusta- tion that usually conceals the taxonomically Heteroptera, Aradidae, Carventinae, Atactocoris, new species, Jamaica important cuticular structures. Head, an- tennal segments I to III, and legs with long erect bristles, lateral borders of body beset with shorter ones which are partly present also on dorsal and ventral surface. Head.—Wider than long (1.31:115) (length measured from apex of genae to transverse furrow delimiting the vertex pos- teriorly); anterior process of genae slightly produced over clypeus, its apex rounded. Antenniferous tubercles cylindrical with pointed anterolateral process. Eyes small, stalked. Postocular portion of head strongly converging; neck with 2 (1 + 1) transverse bladelike elevations that bear a tuft of long bristles. Vertex with a median elevation that is separated by deep curved sulci from ru- gose lateral portions. Antennae about twice as long as width of head across eyes (2.70: 1.39), relative length of antennal segments T::00:1V = 1.15:0.45:0.72:0.37. Antennal segment I distinctly shorter than width of head (1.15:131 = 0.88). Rostrum short, not reaching closed elevated border of wide ros- tral groove, rostral atrium slitlike. Thorax.—Pronotum about 4.5 x as wide as long with an ill defined collar. Lateral margins sinuate, thickened, elevated and 12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. Atactocoris perneri, n. sp., holotype male. 1, dorsal view; 2, ventral view, pilosity omitted; 3, lateral view of terminal segments. Scale 1 mm. granular, posterolaterally produced and converging to angular anterolateral corners. Disc with smooth oblique elevations sepa- rated by deep furrows, medially with a small longitudinal ridge that is narrowed anteri- orly and transversely depressed marking the posterior limit of pronotum. Metanotum wider than pronotum with sinuate lateral margins that are also pro- jecting, thickened and elevated. Separated from pronotum and metanotum by deep transverse furrows laterad of median ridge that extends posteriorly over fused meta- notum and tergites I + II. Disc with smooth longitudinal elevations, deeply excavated laterad of the median ridge. Metanotum wider than mesonotum, completely fused with tergites I + II, its lateral margins similar to those of meso- notum, converging anteriorly. Fused me- dian ridge enlarging posteriorly where it is incised. Disc with irregular smooth longi- tudinal ridges that show a certain variability within the examined type series as well as VOLUME 97, NUMBER 1 73 Figs. 4-6. 4, 5. Atactocoris perneri, n. sp., female; 4, dorsal view; 5, ventral view of terminal segments. 6. Atactocoris farri, holotype female dorsal view. Arrows indicate distinguishing characters. Scale 1 mm. 74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the position and extension of the sulcus of the median ridge that may reach the line of the anterior border of metanotum. Abdomen.— Tergal plate formed by fused mediotergites III to VI with deep depressed glabrous areas and a flat median ridge bear- ing the dorsal abdominal scent gland open- ings and smaller depressions of different size. Mediotergites and dorsal laterotergites (Dltg) separated from each other and from thorax by a suture. Surface of Dltg II to VII rugose with deep apodemal impressions, their lat- eral margin is formed by the reflexed ventral laterotergites which are partly projecting. Ditg I + II separated by a suture. Dltg VII raised medially for the reception of the py- gophore, paratergites VIII small, rounded. Ventral side.—Pro-, meso-, and meta- sternum and sternites I + II fused but marked by deep transverse furrows. Pro- sternum with a deep median impression; mesosternum medially with paired deep impressions, the ridge between 1s partly lowered; metasternum shows similar pits as mesosternum with additional rugosities la- terad of them. Sternite III only medially fused to I + II, IV to VII completely sep- arated. Surface of sternites III to VII shiny, deeply excavated around apodemal im- pressions with a mat median subtriangular area that is also present on fused medio- sternites (Mst) I + II, Mst VII (in male) and metasternum. Spiracles II to IV ventral, close to margin, V to VII lateral and visible from above, VIII terminal. Male genitalic structures (Figs. 7-12).— Pygophore with a median dorsal sulcus flanked by irregular curved ridges covering the exposed part. Upper margin formed by an elevated curved ridge that bears long se- tae. Parameres long and slender, bladelike, projecting from dorsal opening at rest. Legs long and slender, trochanters fused with femora, claws with thin curved pul- villi. Female.—Similar to male but larger and abdomen more rounded laterally. Ditg VII medially with an elevated transverse ridge bearing dense pilosity. Paratergites VIII short and rounded, reaching apex of tergite 1D. Coloration.— Uniformly dark brown, ap- pendages yellowish brown. Holotype.— Male, Jamaica, Portland, Fern Hill near Frenchman Cove, in rain for- est under bark of rotten log 1.3.1993 leg. F. Perner and E. Heiss. The holotype is pre- sented to the Tiroler Landesmuseum Inns- bruck and deposited as permanent loan in the collection of the author. Paratypes.—24 males and 6 females col- lected with holotype. 2 males and 2 females are from Jamaica, Portland, Sherwood for- est near Dragon Bay on dead log of Akee tree (Blighia sapida Konig, Sapindaceae) 24.2.1993 leg. E. Heiss. Paratypes also will be deposited at National Museum of Nat- ural History, Washington, D.C., and Insti- tute of Jamaica, Kingston, and the collec- tion of E. Heiss. Measurements. — Holotype male: Length 6.3 mm; thorax length 2.2 mm; max. width of pronotum 2.1 mm, mesonotum 2.3 mm, metanotum 2.65 mm. Width of abdomen across tergite V 3.0 mm, VI 2.9 mm, VII 2.35 mm. Ratio length of antennae/width of head across eyes 2.05, ratio antennal seg- ment I/width of head 0.88. Male paratypes range in size from 6.2—6.6 mm, females from 7.6-8.2 mm. Etymology.—Named in honour of my friend F. Perner on the occasion of his 60th birthday and in recognition of his efforts and enthusiasm when searching for apter- ous Aradidae. Atactocoris farri Kormilev Fig. 6 Atactocoris farri Kormilev 1964:115. The female holotype has been cleaned from incrustation for comparison of cutic- ular structures with those found in A. per- neri, n. sp. As the description given by Kor- milev is sufficient, only some additional information is given. The characteristic depressions on pro-, meso- and metasternum are also present and VOLUME 97, NUMBER 1 Figs. 7-12. Atactocoris perneri, n. sp., male genital structures; 7, pygophore caudal view; 8, lateral view; 9, dorsal view. 10-12, left paramere in different positions. Scale Figs. 7-9, 0.5 mm, 10-12, 0.1 mm. might be of generic importance, as a differ- ent extension and position of such pits were observed in other apterous Carventinae genera. Contrary to the description and fig- ure 4 given by Kormilev (1964) there are no postocular tubercles present. Measurements of the cleaned speci- men.—Length 9.1 mm; thorax length 2.95 mm, width of pronotum 2.92 mm, meso- notum 3.3 mm, metanotum 3.85 mm; width of abdomen across tergite V 4.2 mm, VI 4.05 mm, VII 3.45 mm. Head width/length 63/56, antennal segments I:II:HI:1V = 67: 28:39:19. Ratio length of antennae/width of head 2.43, antennal segment I/width of head 1.06. Discussion: From the Neotropical Re- gion 27 genera of Carventinae are known, of which 24 are apterous. Kormilev and Van Doesburg (1977) provided a key for the then known 11 apterous genera (12 catalogued by Kormilev and Froeschner 1987), which was also the basis for a new key given by Grillo Ravelo (1988) where he included his 12 new genera from Cuba. The genus Apterocoris belongs to a group of three genera characterized by closed la- bial atrium, tergites I + II fused with meta- notum and Ditg II + III separated by a suture, including also Peggicoris Drake (1956) and Rhysocoris Usinger and Mat- suda (1959). Peggicoris is distinguished by its uninter- rupted median longitudinal ridge, which ex- tends from pronotum to anterior margin of abdominal disk; Rhysocoris by narrow and long head without stalked eyes and the po- sition of spiracles which are all lateral and visible from above. The 12 genera described from Cuba by Grillo Ravelo are difficult to recognize without respective illustrations and belong to another group of genera with fused Ditg II + III. ACKNOWLEDGMENTS Special thanks are due to Mr. T. Farr, Kingston, Jamaica, for the loan of Aradidae under his care and Mr. Fred Perner, Inns- bruck, for his able assistance during our col- lecting trip to Jamaica. 76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Drake, C. J. 1956. New Neotropical genera and spe- cies of apterous Aradidae. Journal of the Wash- ington Academy of Sciences 46: 322-327. Grillo Ravelo, H. 1988. Los Aradidos (Heteroptera) de Cuba I Subfamilia Carventinae. Universidad Central de Las Villas Cuba. 113 pp. Kormilev, N. A. 1964. Neotropical Aradidae XIII (Heteroptera: Aradidae). Journal of the New York Entomological Society 72: 112-119. Kormilev, N. A. and R. C. Froeschner. 1987. Flat bugs of the world. A synonymic list (Heteroptera: Aradidae). Entomography 5: 1-246. Kormilev, N. A. and P. Van Doesburg. 1977. A new genus and new species of the Carventinae from Surinam (Hemiptera, Aradidae). Zoologische Me- dedeelingen 52(1): 1-6. Usinger, R. L. and R. Matsuda. 1959. Classification of Aradidae. British Museum (Natural History), London. 410 pp. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 77-85 SYSTEMATIC NOTES ON SOME SRI LANKAN SCOLITDAE (HYMENOPTERA: ACULEATA) KARL V. KROMBEIN Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. Abstract. —Campsomeriella litoralis Krombein, new species, is described from both sexes, Scolia (Discolia) lankensis Krombein, new species, is described from a male, and the previously unknown female of Scolia (Discolia) gunawardaneae Krombein is also described. Distributional and systematic notes are added for 20 other species obtained since the family was revised in 1978. Key Words: A revision of 25 taxa of Ceylonese Sco- liidae (Krombein 1978) was based on spec- imens collected during the Smithsonian’s “Ceylon Insect Project,” 1969-1976, aug- mented by material borrowed from other collections. Additional specimens were col- lected from 1977 to 1981 by teams that in- cluded a specialist from the United States and two or more Sri Lankan technicians. Personnel who participated during this last 5-year period of the Smithsonian’s “Ceylon Insect Project” were as follows in alpha- betical order: D. W. Balasooriya, P. Fer- nando, T. Gunawardane, V. Gunawardane, M. D. Hubbard, M. Jayaweera, L. Jaya- wickrama, P. B. Karunaratne, S. Karuna- ratne, M. Kosztarab, K. V. Krombein, P. A. Panawatta, P. Leonage, S. Siriwardane, L. Weeratunge, N. V. T. A. Weragoda, and T. Wijesinhe. Two specimens are also in- cluded from my recent field work in Sri Lan- ka during 1993. Two new species and the previously un- known female of another species were col- lected. Descriptions of these are followed by new distributional data and a few sys- tematic notes on other species arranged in the systematic sequence used in my revi- sion. Wasps, Scoliidae, Sri Lanka, systematics, distribution The abbreviations T 1-7 and S 1-7 are used in the descriptions to denote abdom- inal terga and sterna 1-7. Campsomeriella litoralis Krombein, NEw SPECIES Figs: 1, 3, 5 Campsomeriella collaris collaris Betrem. Krombein, 1978, in part: 18-19 (Ja-Ela specimens only). Etymology.— From the Latin /itoralis, of the seashore. Male.—Length 14.0 mm, forewing 11.5 mm. Black, the following pale yellow: nar- row streak on basal two-thirds of outer mandibular surface, clypeus except narrow- ly at apex and a spot on basal two-thirds, narrow at upper end and expanding into a quadrate spot in middle, small narrow streak along lower inner eye orbit, pronotum with a narrow apical streak interrupted for a short distance toward scapula and a narrow streak adjacent to propleuron, small anterior spot on tegula, tiny anterolateral spot on scutel- lum, streak posteriorly on apical half of fore- and midfemora and on most of hindfemur, outer surface of tibiae except apices, outer surface of forebasitarsus, broad band cov- 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON _ 10mm _, se Figs. 1-4. WIWQ’ | 4. 1-3, Campsomeriella species. 1, C. litoralis, n. sp., genitalia, ventral aspect at left, dorsal aspect in middle, aedeagus, lateral aspect at right; 2, C. c. collaris (Fabricius), genitalia, oriented as in | (Krombein 1978: fig. 15, by M. C. Druckenbrod); 3, C. /itoralis, n. sp., antenna, lateral aspect. 4, Micromeriella m. marginella (Klug), antenna, lateral aspect. ering most of dorsum of T | shallowly emar- ginate anteriorly in middle and narrowing laterally, broad bands on apical three-fourths of T 2-3 emarginate anteriorly in middle and narrowing laterally, band on apical half of T 4, a pair of transverse spots on apical half of T 5 narrowly separated at midline, and a pair of small posterolateral spots on S 2. Erect vestiture abundant, glittering white except black on seventh abdominal seg- ment, S 6 without a median tuft of longer, dense, black setae as in C. c. collaris (Fa- bricius) (cf. Figs. 5, 6); head and thorax with silvery tomentum, especially dense on tho- rax as in C. c. collaris. Wings slightly in- fumated with weak yellowish reflections. Genitalia as figured, paramere broader and volsella not so setose as in C. c. collaris (cf. Figs.ls2). Female.—Length 15.3-18.2 mm, fore- wing 11.3-13.4 mm. Integument black. Vestiture white except becoming infuscated on apical abdominal segments, erect on oc- ciput and dorsum of thorax anteriorly form- ing a distinct dense ruff, sparse and recum- bent on abdomen; temple, vertex posteriorly, pronotal dorsum and anterior third of scutum with dense, short, appressed setae as in C. c. collaris. Wings brown, not so dark as in C. c. collaris, basal two-thirds lighter than apex, with weak bluish reflec- tions. Punctation much as in C. c. collaris, differing as follows: vertex with a few more scattered punctures, although still mostly smooth; and lower metapleural plate with fewer punctures. Holotype.—é, Sri Lanka, Col[ombo] Dist[rict], Pamunugama, seashore, 16 March 1981, K. V. Krombein, T. Wijesinhe, L. Weeratunge (USNM). Paratypes (all USNM).—2 4, 2, same label data as holotype; 4 é, 2 2, same locality, but 16 January 1977, K. V. Krombein, P. Fer- nando, D. W. Balasooriya, V. Gunawar- dane. 2 6, same locality, but 26 July 1993, K. V. Krombein, A. W. Norden, P. B. Ka- runaratne. 4, Col[ombo] Dist[rict], Uswe- takeiyawa, seashore, same date and collec- VOLUME 97, NUMBER 1 79 Figs. 5, 6. Campsomeriella species, apex of abdomen, lateral aspect. 5, C. litoralis, n. sp.; 6, C. c. collaris (Fabricius). tors as holotype. 4, same locality as preceding specimen, but 0-50 ft, 8 May 1976, K. V. Krombein, P. B. Karunaratne, S. Karuna- ratne, D. W. Balasooriya. 25 ¢, Col[ombo] Dis[trict], Ja-Ela, Old Dutch Canal, sea lev- el, 8 May 1976, K. V. Krombein, P. B. Ka- runaratne, S. Karunaratne, D. W. Balasoo- riya. 2 6, Sri Lanka, Ham[bantota] Dist[rict], Palatupana, W[ild] Llife and] N[ature] P[rotection] S[ociety] Bungalow, 0-50 ft, 18- 21 January 1979, K. V. Krombein, P. B. Karunaratne, T. Wijesinhe, S. Sirawardane, T. Gunawardane. A pair of paratypes have been deposited in the National Museum, Colombo, and a male paratype in the Nat- ural History Museum (BMNH), London, and in the Staatliches Museum fiir Natur- kunde, Stuttgart. Variation. — Male paratypes are 10.1—13.9 mm long. Coloration is variable: the least maculated specimen from Pamunugama has reduced pronotal markings, lacks the lateral spots on scutellum, and the bands on T 2- 3 are about half the width of tergum, and spots are lacking on T 5; on other specimens from Pamunugama, the spots on T 5 range from tiny dots to narrowly separated trans- verse bars; a male from Uswetakelyawa has a complete band on T 5; and the two spec- imens from Palatupana have the clypeus yellow except extreme apex and small me- dian and basal spots, scutellar spots are larg- er, almost meeting on midline in one spec- imen which also has a median spot on metanotum, T 5 has an apical band nar- rowly emarginate anteriorly on midline, spots on S 2 are larger, and S 3 has a pair of small posterolateral spots. Discussion. — Campsomeriella litoralis is known from only four localities in Sm Lan- ka, all on the seashore, one in the extreme southeastern part of the country, the other three about a third of the distance north- ward on the west coast. I expect that C. litoralis is more widely distributed in sandy parts of the litoral area in Sri Lanka, and that it may also occur in similar habitats in southern India. It should be noted that C. c. collaris (Fabricius) has not been collected in the litoral area, although it is otherwise widely distributed in Sri Lanka. The female runs to C. c. collaris in my 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON WW" | iS Ss eee ee 1.0mm 8. Figs. 7, 8. Scolia (Diliacos) lankensis, n. sp. 7, genitalia, ventral aspect at left, dorsal aspect in middle, aedeagus, lateral aspect at right; 8, antenna, lateral aspect. key (1978), and is distinguished by having the basal two-thirds of the forewing lighter and with weak bluish reflections, the mostly pale, sparse vestiture of the abdomen, and the smaller average size (14-27 mm in C. c. collaris). The male also runs to C. c. collaris, but with some difficulty because of the frequent maculations on T 5S. It is separated from C. c. collaris by the absence of a relatively long tuft of dense, erect, long, black setae (“‘cop- ulation brush” of Betrem 1967) on S 6 (cf. Figs. 5, 6), and markedly by the confor- mation and vestiture of the male genitalia (cf. Figs. 1, 2). The more abundantly mac- ulated specimens will run to Micromeriella m. marginella (Fabricius). The antennae are shorter in both C. c. collaris and C. litoralis than in M. m. marginella (cf. Figs. 3, 4), and the apical flagellar segments are two-thirds as wide as long rather than half as long. The genitalia of M. m. marginella are also mark- edly different (Krombein 1978: fig. 12) Scolia (Discolia) lankensis Krombein, NEw SPECIES Figs. 7, 8 Etymology.—From Lanka, the ancient Sinhalese name for Sri Lanka. Male.—Length 22.4 mm, forewing 19.3 mm. Black, the following orange: upper half of clypeus, lower half of area frontalis; large area on face including narrow line along lower inner eye orbits, ocular sinus, and front to posterior edge of fore ocellus, small mark behind and between posterior ocellus and upper eye margin narrowing to a streak along VOLUME 97, NUMBER 1 upper two-thirds of outer eye orbit; oblique rectangular mark on scapula extending from anterolateral margin to tegula; a pair of large, transverse, anterolateral marks on T 3; wide bands on T 4-6; a pair of small, anterolat- eral spots and a narrow median band on T 6; and a narrow, median band on T 7. Ves- titure black, reddish on orange areas except scapula, apical fringes of T 2-6 and S 2-6 and base of S 7. Wings dark brown with golden reflections, anterior third of forewing more strongly infuscated, microtrichiae present on wing membrane beyond margin- al and submarginal cells. Antenna long, tapering gradually at apex (Fig. 8); spatium frontale with small, con- tiguous punctures; front with somewhat larger punctures except for a small, smooth area between median patch and punctate ocular sinus, well developed fissura frontalis extending to anterior ocellus; vertex with more scattered, small punctures. Scapula except narrow, micropunctate posterior margin with small, mostly sub- contiguous punctures; scutum with some- what larger, mostly subcontiguous punc- tures except a small, median, posterior area with slightly more separated punctures; scu- tellum similarly punctate but more sparsely along midline and a small, wider posterior area; metanotum similarly punctate except a narrow, median strip; upper metapleural plate with punctures as on scutum except lower anterior area smooth and micro- punctate; punctation of dorsal propodeal surface similar to that on scutum; lateral propodeal surface contiguously punctate on upper third, elsewhere with most punctures separated by about a puncture width. Apical fringes of T 1-6 and S 2-6 with a single row of tiny, contiguous punctures; T 1-2 with small punctures mostly separated by half a puncture width; T 3-6 with small, more dispersed punctures; genitalia (Fig. 7). Female.— Unknown. Holotype.—Sn Lanka, Kan[dy] Dist[rict], Udawattakele Sanct[uary], 510-580 meters, 2-5-XI-1977, K. V. Krombein, P. B. Ka- 81 runaratne, T. Wijesinhe, M. Jayaweera. De- posited in National Museum of Natural History (USNM). Remarks.—This species runs to couplet 21 in my key (1978), but is distinguished from S. picteti Saussure and S. ceylonicola Betrem, both members of the S. erythro- cephala group, by the dark brown rather than yellowish or slightly infumated wings, lack of pale thoracic markings except on scapula, and the orange rather than yellow maculations of the abdomen. The genitalia differ from those of other Scoliinae in lack- ing setae on the dorsal surface of the para- mere. In Betrem’s key (1928: Tabelle III, pp. 256-263) S. lankensis runs to S. pseudosi- nensis Betrem from Sidapur, Coorg, India, a member of the S. pekingensis group. The unique male allotype of S. pseudosinensis in the Agricultural Station, Pusa, India, is not available for study, and I am not certain that S. lankensis belongs to this group of species. Betrem did not figure the genitalia, and his description indicates the following differences in coloration and punctation be- tween S. pseudosinensis and S. lankensis: area frontalis all black in the former species, orange on the lower half in the latter; spa- tium frontale and tegula posteriorly orange in former, black in latter; thorax beneath and legs with reddish brown setae in former, black in latter; pronotum not thickly, finely punctate in former, scapula mostly contig- uously punctate in latter; and metanotum not thickly punctate in former, most of punctures on latter separated by half the diameter of a puncture. Betrem characterized females of the S. pekingensis group as follows: front and ver- tex mostly yellow or red, and mostly or en- tirely smooth with very few punctures; lam- inae frontales short, broader below than above; thorax entirely black; propodeum densely and deeply punctate; abdominal terga often finely punctate posteriorly; and at least T 3 with pale markings. The un- known female of S. /ankensis should agree 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in most of these details if the taxon is, in- deed, a member of the S. pekingensis group. Scolia (Discolia) gunawardaneae Krombein Scolia (Discolia) gunawardaneae Krom- bein, 1978: 23, fig. 30. I described this rare species of the Dry Zone from a single male from Tennamar- avadi, Trincomalee District, on the north- east coast. We collected a second male and three females on the southeast coast. The specimens are unworn and appear to have eclosed recently. They were visiting a flow- ering plant for nectar, and all specimens bear a few to many pollen grains especially be- neath the head. Female.—Length 14.7-19.0 mm, fore- wing 12.8-16.2 mm. Black, the following light red: largest female with a narrow stripe behind eye on vertex, and a diffuse, narrow spot anteriorly on scapula; broad bands on T 3-5, largest female also with a pair of small, rounded, sublateral spots on middle of 2; sterna black in smallest female, S 2 of intermediate female with a diffuse, narrow reddening anteriorly and a pair of small, elliptical spots laterally at midline, S 3 of largest female with broad anterior band and a pair of transverse spots posterolaterally, and S 4 and 5 with successively smaller, paired posterolateral spots. Wings dark brown, forewing with blue reflections, an- terior third darker than rest of membrane, microtrichiae confined to cells except a small patch adjacent to marginal cell. Erect ves- titure black except white on occiput, nar- rowly on anterolateral margin of pronotum and more widely anterodorsally. Anterior margin of clypeus rounded, flat anterior rim with dense, small punctures, raised upper area glossy, with scattered mi- cropunctures and a few larger ones laterally; area frontalis with a few, large punctures laterally; spatium frontale with large, most- ly subcontiguous punctures; front with large punctures separated by less than the width of a puncture narrowly along midline and along upper eye margin, scattered else- where; fissura frontalis lacking; vertex with scattered medium punctures, narrow patch of close punctures behind ocelli extending toward upper inner eye margin; occiput with close, subcontiguous punctures. Scapula mostly subcontiguously punc- tate; scutum with a smooth, more or less U-shaped area on posterior half, anterior third with small, subcontiguous punctures, narrow area inside parapsidal furrows with larger subcontiguous punctures, area laterad of parapsidal furrows with larger scattered punctures, posterior margin with 2—3 rows of close, smaller punctures; scutellum most- ly with larger, subcontiguous punctures, and a few small, scattered, impunctate areas; median area of metanotum similarly punc- tate, lateral areas with smaller, contiguous punctures; mesopleuron with moderate, subcontiguous punctures except anterior half of anterior slope below tubercle smooth with numerous micropunctures, and posterior half of posterior slope and tubercle similarly micropunctate; upper plate of metapleuron smooth with numerous micropunctures ex- cept for a few larger punctures dorsally, low- er plate with larger punctures separated by about half the diameter of a puncture except a narrow, subtriangular posterior area densely micropunctate; median horizontal area of propodeum punctate as on scutellum except a narrow median strip impunctate; lateral horizontal area of propodeum smooth on inner anterior fourth, with somewhat smaller, subcontiguous punctures else- where; lateral surface of propodeum with mostly medium size punctures separated by no more than half a diameter and with scat- tered small impunctate areas; median area of posterior propodeum with medium size, frequently subcontiguous punctures on up- per half, more scattered punctures on lower half; lateral area of posterior propodeum with mostly subcontiguous punctures ex- cept a small, smooth, micropunctate area at upper inner angle. VOLUME 97, NUMBER 1 Dorsal surface of T 1 with a small tuber- cle, small area immediately posterad with scattered punctures of medium size, punc- tures elsewhere subcontiguous to contigu- ous especially laterally. Discussion.—The female of S. gunawar- daneae runs to couplet 28 in my key (1978), sharing the distinction with S. keiseri Krombein and S. karunaratnei Krombein of having the occiput partly or entirely clothed with white setae. It differs from both in having broad, light red bands on T 3-5; neither of the other species has red on T 5, and T 3 has a pair of spots rather than a band. The female of S. keiseri also differs in having abundant white setae on the entire body rather than just on the occiput and neck, the fissura frontalis is present on lower half of front, and the scutellum has most of the surface devoid of large punctures. The female of S. karunaratnei also differs in having a fissura frontalis on the lower part of the front, the upper front virtually devoid of large punctures, the scutum with a large, quadrate, impunctate area posteriorly, and the scutellum also is less punctate posteri- orly. The male agrees well with the holotype in coloration and punctation, and in the characteristic genitalia. Additional locality data.—3 9, 6; Ham- bantota District, Palatupana, 29 Mar—2 Apr 1981, K. V. Krombein, T. Wiesinhe, L. Weeratunge. One female has been placed in the National Museum, Colombo, Sri Lanka. The unique male holotype which was on loan to USNM has now been returned to the National Museum in Colombo. Phalerimeris phalerata turneri (Betrem) Additional localities. —Trincomalee Dis- trict: Tennamaravadi, 3 6, 20 Mar. Anu- radhapura District: Amarivayal, 12 4, 21 Mar; Padaviya, 6 4, 12—22 Mar; and Gal- kadawala, 4, 13 Mar. Kurunegala District: Kurunegala, Badegamuwa Jungle, 3 4, 14— 15 Mar, and 2, 2 4, 20 Sep. Kandy District: 83 Ulhitiya Oya, 15 mi NNE of Mahiyangana, 6, 5-6 Sep; and Thawalamtenne, 3, 16-18 Sep. Ratnapura District: Sinharaja Jungle, 2-3 mi S of Weddagala, 4, 8-12 Feb, and 4 6, 22-23 Sep; Belihul Oya, Rest House, 4, 23 Mar, and 24 6, 9-11 Apr; and Ambame Hena, 8 mi N of Kalawana, é, 4 Apr. Mon- aragala District: Angunakolapelessa, 4, 17- 19 June. Colpacampsomeris indica eliformis (Saussure) Additional localities.—Ratnapura Dis- trict: Gilimale, Induruwa Jungle, ¢, 12-15 Mar; and 2 mi S of Weddagala, Sinharaja Jungle, 6, 8-12 Feb. Sericocampsomeris pseudindica (Betrem) Additional localities.—Kegalla District: Kitulgala, Bandarakele Jungle, 3, 17-18 Mar. Ratnapura District: 2 mi S of Weddagala, Sinharaja Jungle, 3 , 8-12 Feb. Campsomeriella collaris collaris (Fabricius) The paired lateral spots on T 5 of the male may occasionally coalesce medially to form a narrow strip. Additional localities. —Mannar District: Kondachchi, Ma Villu, 2, 6 6, 11-12 Apr, and é, 19 Sep. Trincomalee District: Ten- namaravadi, 9 6, 20 Mar. Colombo District: Katunayaka, 6, 16 June. Kegalla District: Kitulgala, Bandarakele Jungle, 4, 15 Apr, and 4, 25-26 Oct. Ratnapura District: Be- lihul Oya, Rest House, 6, 10-11 Apr. Galle District: Kanneliya section, Sinharaja Jun- gle, 6, 2-5 Oct. Monaragala District: An- gunakolapelessa, 3 6, 17-19 June. Micromeriella marginella marginella (Klug) A large male from Colombo, 12 mm long, has abnormally broad bands on T 1-3, sim- ilar in extent to those of Campsomeriella collaris collaris (Fabricius). Unlike C. c. col- laris, it has the usual apical band on T 5, 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and the genitalia are normal for M. m. mar- ginella. Additional localities. —Mannar District: Kondachchi, Ma Villu, 6, 22—28 Jan, ?, 17- 21 Feb, 2, 13 6, 11-12 Apr, and 9, 16-19 Sep. Colombo District: Pamunugama, sea shore, 6, 16 Jan, and 2°, 16 Mar; and Gam- paha Botanical Garden, 6, 27 Sep. Ham- bantota District: Bundala Sanctuary, Cir- cuit Bungalow, 5—S0 ft, 8 2, 17 6, 22-24 Aug. Megacampsomeris ceylonica ceylonica (Kirby) The male from Thawalamtenne is very small (10.5 mm) and has reduced yellow markings: head with only clypeus mostly yellow, elsewhere black; thorax entirely black; apical bands of T 1-3 narrower, those of 2 and 3 separated in middle; and legs with normally red areas brownish. Additional localities.—Kandy District: Thawalamtenne, ?, 6, 12—13 Mar; and Ad- ams Peak Trail, 4.5 mi W of Maskeliya, 1610-1690 m, 14 4, 20-21 Oct. Kegalla Dis- trict: Kitulgala, Makande Mukalana, 4, 3-4 Feb, and 4, 25-26 Oct. Liacos erythrosoma cruszi Krombein Additional locality.—Trincomalee Dis- trict, Thiriyaya, 4, 14 Mar. Megascolia (Regiscolia) azurea michaae (Betrem) Additional localities.—Anuradhapura District: Galapitawewa, 2 4, 19 Mar; and Amarivayal, 3 6, 21 Mar. Trincomalee Dis- trict: Thiriyaya, 7 6, 14 Mar. Kandy Dis- trict: Thawalamtenne, 3, 7-8 Sep. Colombo District: Handapangoda Timber Reserve, 2, 18 Jan. Microscolia hydrocephala (Micha) Additional localities.—Kandy District: Thawalamtenne, 6, 21 Mar, and 11 3, 7-8 Oct. Monaragala District: Angunakolape- lessa, 2, 17-19 June. Austroscolia ignota (Betrem) Additional localities. —Mannar District: Cashew Corporation, Ma Villu, 2 4, 17-21 Feb. Trincomalee District: Tampalakaman Naval Head Works, 6, 29 Jan; and Ten- namaravadi, 6, 20 Mar. Amparai District: Ekgal Aru Sanctuary Jungle, 2, 9-11 Mar. Galle District: Kanneliya, Sinharaja Jungle, 2 3, 24-26 Jan. Austroscolia ruficeps henryi Krombein This is the second known male of this rare taxon. It agrees very well with the descrip- tion of the allotype, but it is smaller, only 14.5 mm long rather than 20 mm. Additional locality.—Trincomalee Dis- trict: Tampalakaman Naval Head Works, 6, 29 Jan. Scolia (Discolia) cyanipennis Fabricius Additional localities.—Matale District: Sigiriya, ?, Aug. Hambantota District: Pal- atupana, 3 2, 6, 29 Mar—2 Apr. Scolia (Discolia) affinis Guérin-Meéneville Additional localities. —Anuradhapura District: Padaviya Archeological Site, 3, 1 1- 14 Oct. Colombo District: Labugama Res- ervoir, 2, 6, 11 Jul. Ratnapura District: Gil- imale, Induruwa Jungle, 6, 7-8 Mar, and 4, 16-19 Apr. Galle District: Kanneliya sec- tion, Sinharaja Jungle, 6, 2-5 Oct. Scolia (Discolia) trivandrumensis Betrem Additional localities. —Mannar District: Kondachchi, Ma Villu, 3 6, 11-12 Apr; and 0.5 mi NE Kokmotte Bungalow, Wilpattu Natl. Park, 2 3, 22-23 Jan. Trincomalee District: China Bay, Ridge Bungalow, 2 4, 8-11 Oct. Galle District: Kanneliya section, Sinharaja Jungle, 6, 2-5 Oct. Hambantota District: Palatupana, °, 29 Mar—2 Apr. VOLUME 97, NUMBER 1 Scolia (Discolia) aureipenniformis Betrem Additional localities. —Trincomalee Dis- trict: Trincomalee, China Bay, 2 6, 8-11 Oct. Colombo District: Labugama Reservoir, 10 6, 11 Jul. Galle District: Kanneliya section, Sinharaja Jungle, 2 6, 2-5 Oct. Scolia (Discolia) fasciatopunctata Gueérin-Méneville Additional localities. —Mannar District: Kondachchi, Ma Villu, 6, 11-12 Apr. An- uradhapura District: Galkadawala, 6, 13 Mar. Amparai District: Ekgal Aru Sanctu- ary Jungle, 6, 9-11 Mar. Kegalla District: Kitulgala, , 3-5 Feb. Ratnapura District: Gilimale, Induruwa Jungle, 3, 13-15 Mar. Galle District: Hiniduma, 2, 15 Jul. Scolia (Discolia) bipunctata bipunctata Fabricius Additional localities. — Trincomalee Dis- trict: Nilaweli, 9 6, 19-20 Nov; and Ten- namaravadi, 5 6, 20 Mar. Scolia (Discolia) karunaratnei Krombein Additional localities.—Mannar District: Ma Villu, Cashew Corporation, 9, 6, 17-21 Feb. Puttalam District: Pannika Wila, Wil- pattu Natl. Park, 6, 1 Nov. Colombo Dis- trict: Colombo, ¢, 27 June. Hambantota District: Palatupana, WLNPS Bungalow, 3 3, 18-21 Jan. 85 Scolia (Discolia) keiseri Krombein Additional locality.—Mannar District: Kondachchi, Ma Villu, 3, 11-12 Apr. Scolia (Discolia) picteti Saussure Additional locality.— Mannar District: 0.5 mi NE of Kokmotte, Wilpattu Natl. Park, 6, 15-16 Feb. ACKNOWLEDGMENTS I am grateful to George Venable for pre- paring the line drawings (except Fig. 2) il- lustrating this paper, to Beth B. Norden for preparation of specimens for scanning elec- tron microscopy, and to Susann Braden for making the micrographs (all of the Smith- sonian Institution). I thank Till Osten, Staa- tliches Museum fiir Naturkunde, Stuttgart, for helpful discussion of systematic prob- lems. I am grateful to Till Osten, and to Arnold S. Menke, Systematic Entomology Laboratory, USDA, for their helpful review of the manuscript. LITERATURE CITED Betrem, J. G. 1928. Monographie der Indo-Austral- ischen Scoliiden (Hym. Acul.) mit zoogeogra- phischen Betrachtungen. Treubia, 9 (supplemen- tary volume), 398 pp. . 1967. The natural groups of Campsomeriella Betr., 1941 (Hymenoptera Scoliidae). Entomolo- gische Berichten 27: 25-29. Krombein, K. V. 1978. Biosystematic studies of Cey- lonese wasps, II: A monograph of the Scoliidae (Hymenoptera: Scolioidea). Smithsonian Contri- butions to Zoology 283: 1-54. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 86-89 NOTES ON THE BEHAVIOR AND TAXONOMY OF MEGACHILE (XEROMEGACHILE) BRIMLEYI MITCHELL AND ITS PROBABLE CLEPTOPARASITE, COELIOXYS (XEROCOELIOX YS) GALACTIAE MITCHELL (HYMENOPTERA: MEGACHILIDAE) KARL V. KROMBEIN AND BETH B. NORDEN Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. Abstract. —Megachile brimleyi Mitchell and its probable cleptoparasite, Coelioxys gal- actiae Mitchell, were studied in south central Florida during August, 1992. Foraging adults were collected along with other bees and wasps attracted to the flowers of Galactia regularis and Tephrosia chrysophila. Excavation of two active M. brimleyi nests provided details of nest structure and cell construction. Difficulties in keying both female and male C. galactiae are discussed, and revised couplets are provided for females. Key Words: A brief visit to the Archbold Biological Station (ABS), Highlands County, Florida, provided an opportunity to observe and col- lect Megachile brimleyi Mitchell and its probable cleptoparasite, Coelioxys galactiae Mitchell. Bees were collected 14-16 August 1992, and two active nests were dug up on the 15th. Plants visited by the bees were collected and pressed. Voucher insect and plant specimens are deposited in the Na- tional Museum of Natural History, Smith- sonian Institution (USNM) and the Arch- bold Biological Station (ABS). Stupy SITE Bees were studied along a firebreak cut- ting through sand pine scrub near the south- ern end of the Lake Wales Ridge. This nar- row, north-south ridge of sandy uplands is part of the ancient line of dunes that re- mained above water during Pleistocene in- undations. The substrate is composed of loose, small grained, white quartz sand and particles of charred organic material, the residue from repeated scrub fires. The ster- Megachile, nest, Coelioxys, cleptoparasite ile, sandy soil is well drained but moist be- low the surface. MATERIALS AND METHODS Adult bees were collected on plants, and observed as they excavated nests. Plant specimens were put in plastic bags and transferred later to a plant press. The two bee nests were examined after blowing plas- ter-of-Paris into the nest openings to trace their paths, and then carefully removing sand grains from around the tunnels with a tablespoon, penknife, and small paint brush. Nest contents were preserved in Kahle’s so- lution, then examined and measured in the laboratory. The bees were identified using Mitchell (1962, 1973) and by comparison with specimens identified by him in USNM. RESULTS AND DISCUSSION Nest architecture.—Nesting data have been reported for only one other species of the subgenus Xeromegachile, M. rubi Mitchell in North Carolina (Sivik 1954) and Georgia (Eickwort et al. 1981). We found VOLUME 97, NUMBER 1 Table 1. Tiphiidae Tiphia intermedia Malloch: 4. Larridae Trypoxylon johannis Richards: 4. Philanthidae Cerceris tolteca Saussure: 4. Halictidae Nomia maneei Cockerell: 2 without pollen. Augochloropsis sumptuosa (Smith): 2 without pollen. Megachilidae Anthidium maculifrons Smith: 2 without pollen. Anthidiellum notatum rufimaculatum Schwarz: 6. 87 Wasp and bee visitors to flowers of Galactia regularis. Megachile (Litomegachile) brevis pseudobrevis Mitchell: 6, 2 with abundant G. regularis pollen on scopa. Megachile (Litomegachile) mendica mendica Cresson: 3 2, two with abundant G. regu/aris pollen on scopa, one with no pollen. Megachile (Xeromegachile) brimleyi Mitchell: 4, 2 2 without pollen. Megachile (Xeromegachile) integra Cresson: 5 4, 2 2, one female with G. regularis pollen on the scopa, one without. Coelioxys (Xerocoelioxys) galactiae Mitchell: 2. M. brimleyi females digging their own nests in sandy soil exposed to full sunlight, sim- ilarly to M. rubi. M. brimleyi tunnels were in level ground, and penetrated the sub- strate at an angle of 45°. The entrances of the two M. brimleyi nests were 8.5 and 10.0 mm in diameter. The females used their hindlegs to kick the sand grains 3-6 cm from the nest entrances, thus forming a semicir- cular spoil heap. Both tunnels were 10 cm long, and ended in an enlarged chamber, a single cell. It was our impression that these nests, the only two found in the area, were newly initiated, and eventually might have contained more cells. Sivik reported that nests of M. rubi had 1-3 cells in a linear series, while 1+ cells were reported by Eick- wort et al. The cell of one M. brimleyi nest was lined with 11 leaf pieces cut from Galactia re- gularis. Eight of the pieces were ca 9 mm wide and 15.5 mm long with rounded edges. Three subcircular pieces, used to form the cell base, were ca 7 mm wide and 8 mm long. The leaf cuttings were weakly ce- mented together. The cell was ca 17 mm long, and it had not been provisioned or capped. Sivik found that M. rubi formed cells of similar size from leaves of Betula lenta, and capped the cells with 4-5 leaf pieces. Eickwort et al. reported ca 14 leaf pieces used to form a cell, and ca 3 to cap Hite Plant and insect associations.—Two le- guminous plants, the pink-flowered Galac- tia regularis, and the white-flowered, pros- trate Tephrosia chrysophylla, attracted a variety of wasps and bees (Tables 1, 2). Both sexes of Megachile brimleyi visited flowers of both plants, and one nesting female car- ried G. regularis pollen on the scopa. Inter- estingly, females of Coelioxys galactiae also visited both plants. Behavioral data strong- ly suggest that C. galactiae is a cleptopar- asite of M. brimleyi. We watched two fe- males of C. galactiae each following a M. brimleyi female, and then entering her nest after her. We also observed other Coelioxys females, presumably galactiae, following M. brimleyi females as they flew over the sand and among the plants. Megachile integra Cresson is the only oth- 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Bee visitors to flowers of Tephrosia chry- sophylla. (No wasps were noted on this plant.) Halictidae Nomuia maneei Cockerell: 3 2, one with abundant T. chrysophylla pollen on the scopa, others with lesser amounts. Megachilidae Megachile (Xeromegachile) brimleyi Mitchell: 4, 2 without pollen. Coelioxys (Xerocoelioxys) galactiae Mitchell: 2 °. er Xeromegachile that we collected at the study site. Both sexes were collected on Gal- actia flowers, but only one female had nu- merous pollen grains of Ga/actia on the sco- pa. Nothing is known of the nesting habits of M. integra. Coelioxys galactiae Mitchell Fig. 2 The syntype series of C. galactiae is from North Carolina, Illinois, and northern Flor- ida (Clay and Levy counties). Both sexes have light red legs except for the coxae. However, nine females and four males from ABS in south central Florida have the legs predominantly black or dark brown except for two females which have reddish brown tibiae and tarsi. This is a striking reversal of the normal coloration of aculeate Hy- menoptera in which there is an increasing amount of red in more southern popula- tions. Mitchell (1962) separated females of three species of Coelioxys, galactiae Mitchell, so- dalis Cresson and immaculata Cockerell, from the other eastern species of the genus by the similarly shaped sixth sternum. It is not notched laterally but is incurved api- cally to form a short, acute, subtriangular projection. Females of C. galactiae from ABS do not key out properly in Mitchell (1962, couplets 13-14). In a series of nine females from ABS the tibiae and tarsi are predominantly black or dark brown in sev- en, reddish brown in two. We examined the series of these three spe- cies in the National Museum of Natural History (USNM), and the syntypes of C. galactiae in the collection at North Carolina State University. We believe that the fol- lowing key will distinguish the three taxa with greater certainty. It should replace cou- plets 13 and 14 in Mitchell (1962). 13. Carina on pronotal lobe strongly raised, la- mellate; vestiture of head short, appressed, dense on clypeus and lower front; thorax with areas of short, dense, appressed white setae— narrow bands anteriorly and posteriorly on scutum, narrow anterior band on scutellum sometimes interrupted medially and the pos- terior declivous surface, small median patch on metanotum, narrow anterior and posterior bands on mesopleuron; clypeus slightly con- vex, not elevated above level of face Ree eee a ee dom et os C. galactiae Mitchell Carina on pronotal lobe not raised, obscured by vestiture in C. sodalis; vestiture of head appressed and not so dense on clypeus and supraclypeal area only; thorax either without appressed setae, or with such areas greatly re- duced in extent; clypeus slightly convex or elevateds (95.15 tneee or ae eee Meee 14 14. Clypeus slightly convex, not elevated above level of face; vestiture of head, except clypeus and supraclypeal area, and thorax long, erect ANGESIIVeIy, Meee ee ee C. sodalis Cresson Clypeus more strongly convex, elevated above level of face as viewed in profile; vestiture appressed on clypeus and supraclypeal area, suberect laterally along orbit and erect on up- per front, shorter than in C. sodalis; bands of appressed, decumbent hair on scutum absent anteriorly, evanescent laterally and posteri- orly; appressed setae on scutellum forming a narrower anterior band than in C. galactiae, sparser on posterior declivous surface; meta- notum with longer, suberect, evenly distrib- uted setae; mesopleuron with narrower an- terior and posterior bands of longer, suberect SClaclGa tae ener C. immaculata Cockerell Floridian males of C. galactiae key to C. boharti Mitchell (Mitchell 1962) because of their dark legs. We examined the unique holotype of C. boharti (USNM), and find that it is readily distinguished by the abun- dant, broad, short setae on the mesopleural disk, 2-3 times as long as wide, the more VOLUME 97, NUMBER 1 { 0.25mm Figs. 1, 2. Coelioxys species, apical half of para- mere, lateral aspect, ventral margin at bottom. 1, C. boharti Mitchell; 2, C. galactiae Mitchell. infuscated wings, and the paramere (Fig. 1) which in lateral view widens gradually for most of the apical half, and then tapers obliquely to the narrowly rounded apex. In C. galactiae the discal setae of the meso- pleuron are narrower, mostly more than four times as long as wide, and the apical half of the paramere (Fig. 2), in lateral view, widens gradually over most of its length with the apex broadly rounded. The discal vestiture of the mesopleuron is variable in width in specimens from ABS, but it is never as broad and short as in C. boharti. 89 ACKNOWLEDGMENTS We are grateful to personnel at the Arch- bold Biological Station for their hospitality, and for providing laboratory facilities. We are indebted to Alan Herndon, Florida In- ternational University, Miami, for identi- fication of plants during our stay at the Sta- tion. We thank R. L. Blinn, Department of Entomology, North Carolina State Univer- sity, for the loan of the syntypes of Coelioxys galactiae Mitchell in that collection. We are grateful to George Venable, Department of Entomology, SI, for preparing the illustra- tions. We also appreciate the assistance of Arnold W. Norden in the collecting of bees and wasps at ABS. We thank S. W. T. Batra and J. Cane for reviewing the manuscript. LITERATURE CITED Eickwort, G. C., R. W. Matthews, and J. Carpenter. 1981. Observations on the nesting behavior of Megachile rubi and M. texana with a discussion of the significance of soil nesting in the evolution of megachilid bees (Hymenoptera: Megachilidae). Journal of the Kansas Entomological Society 54: 557-570. Mitchell, T. B. 1962. Megachilidae, pp. 5-232. In Bees of the Eastern United States, vol. 2. Technical Bulletin 152, North Carolina Agricultural Exper- iment Station. 1973. A subgeneric revision of the bees of the genus Coelioxys of the Western Hemisphere. Department of Entomology, North Carolina State University, Raleigh, 129 pp. Sivik, F.P. 1954. Ecological notes on the three species of solitary bees. Entomological News 65: 253-256. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 90-111 NATIVE BLACK GRASS BUGS (JRBISIA-LABOPS) ON INTRODUCED WHEATGRASSES: COMMENTARY AND ANNOTATED BIBLIOGRAPHY (HEMIPTERA: HETEROPTERA: MIRIDAE) JOHN D. LATTIN, ANNE CHRISTIE, AND MICHAEL D. SCHWARTZ (JDL) Systematic Entomology Laboratory, Department of Entomology, Oregon State University, Corvallis, Oregon 97331-2907, U.S.A.; (AC) William Jasper Kerr Library, Oregon State University, Corvallis, Oregon 97331-4501, U.S.A.; (MDS) Centre for Land and Biological Resources Research, Biological Resources Division, Research Branch, Agriculture Canada, Ottawa, Ontario, K1A 0C6, Canada. Abstract. —The introduction of crested wheatgrasses (Agropyron spp.) into North Amer- ica to improve the carrying capacity of western rangelands resulted in unanticipated increases in the population levels of native black grass bugs (/rbisia spp. and Labops spp.). The more than two hundred publications, reports, and theses listed in this annotated bibliography contain information on the systematics, biology, and ecology of the bugs and management techniques for controlling and minimizing bug damage to these non- indigenous species of wheatgrasses. Key Words: Black grass bugs, /rbisia, Labops, Hemiptera: Heteroptera, Miridae, plant bugs, bibliography, rangeland, non-indigenous species, wheatgrasses, Agro- Dyron INTRODUCTION Grasslands play an important environ- mental role worldwide. Over 7 billion hec- tares occur, including rangeland and pas- ture, exceeding forests and woodlands by over one-half billion hectares (Lauenroth 1979, Turner and Meyer 1992). Grasslands of all types have always received extraor- dinary use as a resource for grazing by both wild and domestic animals (Crawley 1983). The area occupied by grasslands globally declined from 1700 to 1980, chiefly because the land was being converted to other ag- ricultural uses, largely croplands. Grassland areas have increased since 1950, primarily at the expense of forests and woodlands (Turner and Meyer 1992). Such shifts in global carbon storage patterns are becoming more apparent and the consequences of these conversions are attracting worldwide con- cern. This paper addresses the interaction be- tween several groups of native grass in- sects—the black grass bugs (Hemiptera: Heteroptera: Miridae) and range grasses, chiefly non-indigenous species that have been planted widely throughout western North America for range improvement. Most of the annotated references deal with some aspect of the habits and characteristics of plant bugs of the genera /rbisia Reuter and Labops Burmeister. While no simple solution to the problems caused by these insects emerged from this review, we have documented the diverse interactions between introduced organisms (here, several grass species from Central Asia) and elements of the native insect fau- na, some with close relatives in the original VOLUME 97, NUMBER 1 home of the grasses. While it is more com- mon for crop pests to be non-indigenous themselves, host plant shifts of native in- sects onto introduced plant species do occur (Kogan and Lattin 1993). The basic prob- lem of damage to the planted wheatgrass species results from these host shifts. The bugs continue to feed on their native hosts, of course, but the coevolution of these in- sects with their native hosts generally results in resistance to extensive damage by the bugs. This resistance normally is not present in the introduced grass species. Breeding programs for rangeland grasses have tended to emphasize agronomic improvements rather than resistance to insects (Watts et al. 1982). If, at the time of original introduction of the wheatgrasses, a thorough review of po- tential pests had been made, it would have disclosed that at least one native species of Labops found in Central Asia feeds upon species of Agropyron Gaertn. (Kerzhner 1973). This example illustrates the desira- bility of careful screening of plant materials (and, of course, animals) before their intro- duction to new regions. HISTORICAL BACKGROUND The grasses Dillman (1946) provided a detailed ac- count of the introduction of the crested wheatgrasses into North America. N. E. Hansen of the U.S. Department of Agri- culture first encountered crested wheat- grasses in Asia when he visited the Valuiki Experiment Station about 150 miles north of St. Petersburg in 1897-1898. These grasses (Agropyron spp.) were already being raised and tested for cultivation by Vasili S. Bogdan. Hansen obtained seed samples to be tested upon his return to the United States. Samples were sent to five different states but information about the results of these early plantings is scarce. A second shipment provided by Bogdan arrived in late 1906. This was the one that 91 led to the successful planting and distribu- tion of crested wheatgrass in North Amer- ica. One lot was labeled Agropyron crista- tum (L.) Gaertn. and five lots were labeled A. desertorum (Fisch.) Schult. The second shipment was divided and sent to 15 ex- periment stations from 1907 to 1913. Ac- cording to Dillman (1946) “All evidence thus far collected points to the conclusion that the successful plantings made at the Belle Fourche Station, Newell, S. Dak., from 1908 to 1915, and the experiments begun in 1915 at the Northern Great Plains Field Station, Mandan, N. Dak., were responsible for the early distribution and establishment of crested wheatgrass in the northern Great Plains.’’ Love and Hanson (1932) provided an early treatment on the life history of crested wheatgrass and included a key to identify the species then involved. Dewey (1983) reviewed the taxonomy of wheat- grass and related grasses. Barkworth and Dewey (1985) provided an overview of the perennial Triticeae of North America, in- cluding the crested wheatgrasses. Although Rogler and Lorenz (1983) ex- tolled the virtues of crested wheatgrass as being just about the perfect range grass, they did suggest that some improvements were possible and needed—including insect re- sistance. Most of the references included be- low are related to insect problems of only one type—black grass bugs, mainly as pests of crested wheatgrasses. One wonders how intensive breeding programs using native grasses might have resulted in breeding lines of similar forage desirability. The bugs The insects belong to the family Miridae (Hemiptera: Heteroptera), the largest family of the true bugs, about 2000 species are known from Canada and the United States (Henry and Wheeler 1988). While most of the species are herbivorous, some are pre- daceous and some feed on both plants, in- sects and other arthropods. Several genera have species that feed on grasses. Most of 92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON these genera are found in the tribe Steno- demini, subfamily Mirinae. Some of these genera are mentioned in the papers below, including Leptopterna Fieber, Litomiris Slater, Stenodema Laporte, and Trigono- tylus Fieber. Also mentioned in several pa- pers is Conostethus Fieber, a genus that 1s placed in the tribe Phylini, subfamily Phy- linae. The two genera discussed here as black grass bugs are /rbisia (Mirinae: Mirini) and Labops (Orthotylinae: Halticini). Some of the earlier literature referred to species of the genus 7hyrillus Uhler, but this genus was synonymized with /rbisia many years ago. The species of /rbisia and Labops that feed on crested wheatgrasses are all native species. The only non-indigenous species of bug mentioned in the literature reviewed here is Leptopterna dolabrata (Fallén), the meadow plant bug, a species introduced into North America many years ago from Eu- rope. There are other non-indigenous spe- cies of mirids that are known to feed upon grasses in North America (Wheeler and Henry 1992); they were not included in this annotated bibliography. Irbisia Reuter, 1879, contains 23 species (Schwartz 1984); all of the species are found in North America, mainly in the western provinces and states. The currently accept- ed species are J. bliveni Schwartz, 1984, J. brachycera (Uhler), 1872, I. californica Van Duzee, 1921, I. cascadia Schwartz, 1984, J. castanipes Van Duzee, 1921, J. cuneoma- culata Van Duzee, 1934, I. elongata Knight, 1941, I. fuscipubescens Knight, 1941, I. in- comperta Bliven, 1963, I. knighti Schwartz and Lattin, 1984, J. /imata Bliven, 1963, I. mollipes Van Duzee, 1917, I. nigripes Knight, 1925, J. oreas Bliven, 1963, J. pa- cifica (Uhler), 1872, I. panda Bliven, 1963, I. sericans (Stal), 1858, J. serrata Bliven, 1963, I. setosa Van Duzee, 1921, I. shulli Knight, 1941, 7. si/vosa Knight, 1961, J. sita Van Duzee, 1921, J. solani (Heidemann), 1910. The only species of this genus not restricted to North America is J. sericans whose range extends into eastern Siberia (Kulik 1965, Vinokurov 1979, Kerzhner 1988). Several species of Irbisia have been re- ported from the crested wheatgrasses as well as other graminoid plants and, in fact, from other types of plants besides the grasses. Species of /rbisia often shift from grasses to other plants late in their adult stage, some- times to crops of economic importance. Eggs of some bug species are placed in grass stems while other bug species have been reported to deposit their eggs in non-graminoid plants (Schwartz 1984). Before the genus was re- vised, references were often made only to the genus rather than to a particular species, although J. pacifica was often cited because it was abundant and distinctive in appear- ance. The revision by Schwartz (1984) placed the taxonomy of /rbisia upon stable ground. Labops Burmeister, 1835, contains 12 species and occurs in both the Old and New Worlds (Carvalho 1958, Kerzhner 1988) with one species, L. burmeisteri Stal, present in both regions (Kulik 1965, Vinokurov 1979, Kerzhner 1988). Seven species are found only in North America. The currently accepted species of Labops occurring in North America are: L. brooksi Slater, 1954, L. burmeisteri Stal, 1858, L. chelifer Slater, 1954, L. hesperius Uhler, 1872, L. hirtus Knight, 1922, L. tumidifrons Knight, 1922, L. utahensis Slater, 1954, L. verae Knight, 1929. Of these, only L. hesperius, L. hirtus, and L. utahensis have been reported from crested wheatgrasses. The most economi- cally important species is L. hesperius. The other two species of Labops may be impor- tant in more limited areas, especially L. utahensis. MANAGEMENT CONSIDERATIONS Many papers deal with different methods of coping with the damage caused to the grasses by species of Jrbisia and Labops, depending chiefly on the geographic area. VOLUME 97, NUMBER 1 While insecticide treatments have been used at different times and places in the past, emphasis has shifted towards cultural con- trol, partly because of cost and partly be- cause many compounds used previously are no longer available or registered. Further, the widespread use of pesticides has been deemphasized for environmental reasons. Nontarget organisms are receiving much greater attention, especially in seminatural settings such as rangelands. The main management efforts seem to be in the timing of grazing and in the manip- ulation of the planting mixtures related to the natural vegetation. Pure stands of crest- ed wheatgrass are more likely to sustain greater damage than range that maintains a diversity of species more nearly approxi- mating original conditions (Spangler and MacMahon 1990). Increased efforts to breed resistant strains and to elucidate the nature of this resistance are occurring. Much, how- ever, remains to be done in developing breeding programs for improving native grasses. The original wheatgrasses brought in were first located at a research station in Central Asia where they were being raised for selection for superior varieties in that area. The consequences of introducing several non-indigenous grass species (Agropyron spp.) into western North America have been their widespread use as replacements or augmentations for native grass species, at times approaching monocultures. While this has improved the carrying capacity of many rangelands, it has often been at the expense of the native grasses, species often well adapted to particular localities throughout this vast portion of the continent. The di- versity of native grass species found throughout this region attests to the selec- tion and adaptation that occurred through time, resulting in well-adapted forms. Some of these grasses were unable to withstand the grazing pressures to which they have been subjected, and the introduced grasses seemed to hold the answer to increasing the 93 carrying capacity of these rangelands (Perry 1954). As is often the case, planting a virtual monoculture within a highly diverse flora may create new problems. Expecting one or two species to replace a vast mosaic of grass species adapted over thousands of years to particular conditions may be unrealistic. For example, native insects whose populations were maintained at reasonable levels be- cause of the diversity of grass species nor- mally found in any given area often re- sponded to this newly created monoculture by increasing in density. Natural controls that helped maintain lower population levels of the insects that feed on native grasses may not have been able to sustain those lower levels because of the sudden abundance of new host plants. The increased densities of the insects, of course, reduced the very resources and ben- efits the plantings were supposed to provide. Many of the references included here doc- ument the differences in insect population levels between planted and unplanted rangeland. Some obvious parallels exist between newly created insect problems on modified rangelands and more conventional agricul- tural practices. When native vegetation of any sort 1s converted to croplands, changes in the biota occur (Kogan and Lattin 1993). Monocultures do not resemble most natural vegetational patterns. Usually planted in high density for convenience of harvest, such artificial environments create conditions that often result in rapid build-up of pests, generally at population levels far above what might otherwise occur. Natural control mechanisms in the form of parasites and predators are usually rendered ineffective because the very situation that allowed these controls to function has been disrupted. A long-term consequence of the change to vast monocultures of crops of all types has been a gradual increase in reliance upon a variety of chemicals to provide the levels of control expected. After many years of this depen- 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dency and the development of pesticide re- sistance and the impact of pesticides on the environment, we are again examining al- ternative approaches. The emergence of In- tegrated Pest Management (IPM) is gaining in popularity and in sophistication. It holds real promise in rangeland management where some semblance of native/natural conditions still exists in contrast to a corn- field, for example. With all of the various breeding programs involving the intro- duced crested wheatgrasses, one wonders what might have happened if similar efforts had been applied to our diverse native grass flora. CONCLUSIONS It is unlikely that the black grassbug/ crested wheatgrass problem will ever be re- solved completely. The non-indigenous grasses have been planted so widely that they are unlikely to be replaced or displaced. The native bugs are not likely to disappear either. They have been and continue to be well adapted to the grasslands of western North America. The presence of crested wheatgrasses simply provides additional re- sources and most likely supports higher population levels than existing undisturbed grasslands. Based on the bulk of the inves- tigations reviewed for this publication, it appears that the most likely improvement will occur by moving away from monocul- tures. Bug populations should decrease as habitats become more heterogeneous. In- creased resistance of some varieties of wheatgrasses seems likely to help (e.g. Levin 1973, Ling et al. 1985), as will careful at- tention to grazing pressures—especially when less than ideal climatic conditions fol- low heavy attack by the bugs. Increased at- tention to greater diversity of native grasses already adapted to local conditions pro- vides yet another possibility; the best ap- proach to ameliorating the problem appears to be an ecosystem approach to manage- ment rather than relying on only a few fac- tors. It seems that the use of pesticides is more and more unlikely for a variety of rea- sons. True integrated pest management may be the best approach to this complex prob- lem (Watts et al. 1982). ACKNOWLEDGMENTS We extend our special thanks to George G. Hewitt, U.S. Department of Agriculture (retired) for suggesting this project and for his early efforts on it and to Connie J. L. Foiles, U.S. Department of Agriculture, for her help on the earlier portions of this work. We thank the many authors and librarians who responded to our requests for papers and information and especially Ted Evans, Utah State University; Eric Coombs, Ore- gon Department of Agriculture; Doris Tilles for assistance in obtaining theses on inter- library loan; Olga N. Krankina for assis- tance in Russian translation; and Colleen MacRae for her thorough preparation of the manuscript. We thank the two reviewers for their comments and helpful suggestions. LITERATURE CITED Barkworth, M. E. and D. R. Dewey. 1985. Genom- ically based genera in the perennial Triticeae of North America: Identification and membership. American Journal of Botany 72: 767-776. Carvalho, J.C. M. 1958. Catalogo dos Mirideos do Mundo. Part III. Subfamilia Orthotylinae. Arqui- vos dos Museu Nacional, Rio de Janeiro, Brazil 47: 1-161. Crawley, M. J. 1983. Herbivory: The dynamics of animal-plant interactions. University of Califor- nia Press, Berkeley and Los Angeles. 437 pp. Deitz, L. L. and L. M. Osegueda. 1989. Effectiveness of bibliographic data bases for retrieving ento- mological literature: A lesson based on the Mem- bracoidea (Homoptera). Bulletin of the Entomo- logical Society of America 35: 33-39. Dewey, D.R. 1983. Historical and current taxonomic perspectives of Agropyron, Elymus, and related genera. Crop Science 23: 637-642. Dillman, A. C. 1946. The beginnings of crested wheatgrass in North America. Journal of the American Society of Agronomy 38: 238-250. Henry, T. J. and A. G. Wheeler, Jr. 1988. Miridae, pp. 251-507. In Henry, T. J.and R. C. Froeschner, eds., Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden. 958 pp. VOLUME 97, NUMBER 1 Kerzhner, I. M. 1973. True bugs (Heteroptera) fauna of Tchvinskoy AASR. Fauna of Siberia, Part 2. Academy of Science SSSR, Trudy 16: 78-91 (in Russian). 1988. Miridae, pp. 778-857. In Ler, P. A., ed., Guide to the Insects of the Far East of the USSR. Volume II. Academy of Science of the USSR, Leningrad. 972 pp. (in Russian). Kogan, M. and J. D. Lattin. 1993. Insect conserva- tion and pest management. Biodiversity and Con- servation 2: 242-257. Kulik, S. A. 1965. Blindwansen Ost Sibiriens und des Fernen Ostens. (Heteroptera: Miridae). Acta Faunistica Entomologica Musei Nationalis Pragae 11(98): 39-69 (in Russian with German summa- Ty). Lauenroth, W. K. 1979. Grassland primary produc- tion: North American grassland in perspective, pp. 3-24. InN. R. French, ed., Perspectives in Grass- land Ecology. Springer-Verlag, New York. 204 pp. Levin, D. A. 1973. The role of trichomes in plant defense. Quarterly Review of Biology 48: 3-15. Ling, Y. H., W. F. Campbell, B. A. Haws, and K. H. Asay. 1985. Scanning electron microscope (SEM) studies of morphology of range grasses in relation to feeding by Labops hesperius. Crop Science 25: 327-332. Love, L. D. and H. C. Hanson. 1932. Life history and habits of crested wheatgrass. Journal of Ag- ricultural Research 45: 371-383. Perry, E. 1954. New ranges for old. Western Live- stock Journal 32 (July, 1954): 26-27, 39. Rogler, G. A.and R. J. Lorenz. 1983. Crested wheat- grass—early history in the United States. Journal of Range Management 36: 91-93. Schwartz, M. D. 1984. A revision of the black grass bug genus /rbisia Reuter (Heteroptera: Miridae). Journal of the New York Entomological Society 92: 193-306. Spangler, S. M. and J. A. MacMahon. 1990. Artho- pod faunas of monocultures and polycultures in reseeded rangelands. Environment Entomology 19: 244-250. Turner, B. L. land W. Meyer. 1992. Science capsule. Office for Interdisciplinary Earth Studies, Earth- quest, Summer 1992. 4 pp. Vinokurov, N. N. 1979. Identification manual of the fauna of the USSR. No. 123. Heteroptera of the Yakutia. Zoological Institute of the USSR, Len- ingrad. 232 pp. (in Russian). Watts, J. G., E. W. Huddleston, and J.C. Owens. 1982. Rangeland entomology. Annual Review of Ento- mology 27: 283-311. Wheeler, A. G., Jr.and T.J. Henry. 1992. Asynthesis of Holarctic Miridae (Heteroptera): Distribution, biology, and origin, with emphasis on North America. Entomological Society of America, Tho- mas Say Foundation. No. 15. 306 pp. 95 ANNOTATED BIBLIOGRAPHY This bibliography is an annotated list of publications and reports describing the bi- ology and documenting the occurrence of native black grass bugs on introduced crest- ed wheatgrasses and other plants in North America. The time period covered con- cludes with December, 1993. The list began with an early effort some years ago to bring together the literature on native black grass bugs. This early list was subsequently further developed by a com- prehensive search of several bibliographic sources, either in print, CD-ROM, or online format. These sources included AGRICO- LA, Bibliography of Agriculture, Biological Abstracts, Biological and Agricultural Index (formerly Agricultural Index), CAB Ab- stracts, Dissertation Abstracts, Index to the Literature of American Economic Ento- mology, Review of Agricultural Entomol- ogy (formerly Review of Applied Entomol- ogy Series A. Agricultural), and Zoological Record. All sources were searched from their beginning publication date. The need to search a variety of bibliographic sources to uncover entomological literature has been discussed by Deitz and Osegueda (1989). The list of references cited in each article or report located in the literature search was checked to identify additional publications that were relevant to the scope of the bib- liography. Several theses were located in the process which are included in the bibliog- raphy for completeness, but they have not been annotated. Scientific names which are no longer valid have been corrected to conform with cur- rent taxonomic use. Akingbohungbe, A. E., J. L. Libby, and R. D. Shenefelt. 1972. Miridae of Wisconsin (Hemiptera: Heter- optera). University of Wisconsin, College of Ag- riculture and Life Sciences, Research Division. R. 2396. 24 pp. [Labops hirtus and L. burmeisteri were reported from Wisconsin. The genus Labops was included in a key to the genera of Wisconsin Miridae. } Akingbohungbe, A. E., J. L. Libby, and R. D. Shenefelt. 1973. Nymphs of Wisconsin Miridae, Hemip- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tera: Heteroptera. University of Wisconsin, Col- lege of Agriculture and Life Sciences, Research Division. R. 2561. 25 pp. [The nymph of Labops hirtus 1s characterized. A diagnostic key to the nymphs of Wisconsin Miridae includes Labops. Akingbohungbe, A. E. 1974. Nymphal characters and higher classification analysis in the Miridae (He- miptera: Heteroptera) with a subiamily key based on the nymphs. Canadian Entomologist 106: 687— 694. [The genus Labops was included in a study on dorsal abdominal gland openings in the mirid nymphs. The opening is simple and has a sclero- tized bar, a character shared by most of the Or- thotylinae included in the study.] Anonymous. 1953. Outbreaks and new records. FAO Plant Protection Bulletin 2: 43-45. Rome. [/rbisia brachycera is reported (as I. arcuata) on Agropyron intermedium at the Experimental Station, Leth- bridge, Alberta. ] Anonymous. 1966. Wheatgrass bugs (Labops spp.). United States Department of Agriculture, Agri- cultural Research Service, Plant Protection Quar- antine, Cooperative Economic Insect Report 16: 623-624. [The economic importance of Labops hesperius and L. hirtus in western United States is reviewed. Crested and intermediate wheatgrass- es were injured along with other grasses. Life his- tory, habits, distribution, and a description of L. hesperius are given. A guide to detecting bug dam- age to grasses is included.] Anonymous. 1977. Putting the stops on Labops. Or- egon’s Agricultural Progress. 21: 11-12. [The hab- its and damage caused by black grass bugs (Labops hesperius) in Oregon are reviewed. ] Ansley, R. J., Jr. 1979. The influence of black grass bugs (Labops hesperius) and cattle on the vigor of crested wheatgrass (Agropyron cristatum (L.) Gaertn.). M.S. thesis, Utah State University, Lo- gan. 84 pp. Ansley, R. J.and C. M. McKell. 1982. Crested wheat- grass vigor as affected by black grass bug and cattle grazing. Journal of Range Management 35: 586— 590. [Light to moderate populations (50-200 per m’) of black grass bugs (Labops hesperius) ap- peared to reduce plant vigor in a seeded mono- culture of crested wheatgrass in Utah. Combined grazing by cattle and black grass bugs resulted in greater reduction in vigor than by the insects alone.] Araya, J. E. 1982. Studies of selected predators of black grass bugs (Labops hesperius Uhler and Ir- bisia brachycera Uhler) on ranges of Utah. M.S. thesis, Utah State University, Logan. 102 pp. Araya, J. E. and B. A. Haws. 1988. Arthropod pre- dation of black grass bugs (Hemiptera: Miridae) in Utah ranges. Journal of Range Management 41: 100-103. [Two species of Nabidae, Nabis alter- natus Parshley and N. vanduzeei Kirkaldy, were found to be effective predators on black grass bugs with prey size scaled to predator size. Six species of spiders were also found to feed on the bugs: Xysticus cunctator Thorell, Misumenops lepidus (Thorell), 14. celer (Hentz), Tibellus sp., Casti- aneira sp., and Tetragnatha sp.] Araya, J. E. and B. A. Haws. 1991. Arthropod pop- ulations associated with a grassland infested by black grass bugs, Labops hesperius and Irbisia bra- chycera (Hemiptera: Miridae), in Utah, U.S.A. FAO Plant Protection Bulletin 39: 75-81. [Nabis alternatus and N. vanduzeei were reported to be important predators of Labops hesperius and Ir- bisia brachycera. A number of spiders were also recorded as predators of these two plant bugs in- cluding: Xysticus cunctator, Misumenops lepidus, M. celer, Tibellus sp., Castianeira sp., Tetragnatha sp., and Metepeira foxi Gertsch & Ivie. The value of these generalist predators is discussed. The au- thors suggest diversifying the plant species in the rangeland as a means of reducing the bugs’ im- pact.] Armitage, H.M. 1952. Current insect notes. State of California, Department of Agriculture. Bulletin 41: 180-183. [Labops hesperius is reported on crested wheat and smooth brome grass from Plumus County, California.] Arnott, D. A. and I. Bergis. 1967. Causal agents of silver top and other types of damage to grass seed crops. Canadian Entomologist 99: 660-670. [The damage caused by different organisms that might result in silver top is reviewed. Miridae were in- cluded, but /rbisia and Labops are not mentioned specifically. ] Babcock, J. M., L. K. Tanigoshi, E. A. Myhre, and R. S. Zack. 1993. Arthropods occurring on sweet white lupin and native lupins in southeastern Washington. Pan-Pacific Entomologist 69: 261- 271. [Labops hesperius is reported from the native Lupinus leucophyllus Douglas in southeastern Washington. The bugs were found on the plants towards the end of May and remained until the plants dried out; they were not found on white lupine (Lupinus albus L.), a non-indigenous spe- cies grown as a rotational crop in the area.] Banks, N. 1910. Catalogue of the Nearctic Hemip- tera-Heteroptera. American Entomological Soci- ety, Philadelphia. 103 pp. [The catalog lists early state records for Irbisia brachycera, I. pacifica, and I. sericans, and Labops hesperius. (For more recent information on distribution, see Schwartz 1984, Henry and Wheeler 1988.) (Note: /. gilvipes is now placed in the genus Mecomma Fieber.)] Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America. Nature Publishing Com- pany, Indianapolis. 1116 pp. [Labops hesperius and L. burmeisteri are treated. Each species is de- VOLUME 97, NUMBER 1 scribed briefly and distribution records given. Blatchley states that he considers L. hirtus a syn- onym of L. hesperius.] Blatchley, W. S. 1928. ‘“Quit-Claim” specialist vs. the making of manuals. Bulletin of the Brooklyn Entomological Society 23: 10-18. [The identity of Labops hesperius is noted.] Blatchley, W. S. 1934. Notes on a collection of Het- eroptera taken in winter in the vicinity of Los Angeles, California. Transactions of the American Entomological Society 60: 1-16. [/rbisia cuneo- maculata is described from Sunland, California. Irbisia californica was common in February and March on grasses and herbs.] Bliven, B. P. 1961. New species of /rbisia from Cal- ifornia. Occidental Entomologist 1: 45-49. [Six species of Irbisia are described as new: I. eurekae, I. paenulata, I. vestifica, I. gorgoniensis, I. tejonica, and J. silvosa, all from California. (See Schwartz 1984 for synonymy.)] Bliven, B. P. 1963. New species of /rbisia from Cal- ifornia II. Occidental Entomologist 1: 68-86. [Thirteen species of /rbisia are described as new, 12 from California and one from Arizona. Those described from California include: J. umbratica, I. upupa, I. retrusa, I. incomperta, I. neptis, I. paulula, I. lacertosa, I. panda, I. limata, I. ustri- cula, I. inurbana, and I. serrata. Irbisia oreas is described from Arizona. (See Schwartz 1984 for synonymy.)] Bohning, J. W. and W. F. Currier. 1967. Does your range have wheatgrass bugs? Journal of Range Management 20: 265-267. [Labops hesperius damaged crested wheatgrass throughout much of western North America. Some basic biological in- formation is included. The question was raised as to whether this was a native insect species (it is— J.D.L.). Although 900 acres were sprayed with malathion in the Santa Fe National Forest in 1967, the bugs returned the following year.] Bowers, D. M. 1976. Insect consumption of seeded rangeland herbage in a selected area of Diamond Fork Canyon, Utah County, Utah. M.S. thesis, Utah State University, Logan. 45 pp. Bowers, D. M. and B. A. Haws. 1977. Impact of insect consumption of rangeland forage in Utah. Society of Range Management, 30th Annual Meeting, Abstract, p. 35. [Insects consumed 34% (2.8 AUM) of the total available forage of a study area seeded to crested wheatgrass in Utah County, Utah. Cattle consumed 28% (2.1 AUM) at the same site. These comparative figures were used to calculate losses from insects.] Brewer, P. S., W. F. Campbell, and B A. Haws. 1979. How black grass bugs operate. Utah Science 40: 21-23. [The method of feeding by Labops hesper- ius is described. Photographs and photomicro- oi graphs of the bug and damage are included. In- termediate wheatgrass and Kentucky bluegrass were more attractive to the bugs than crested wheat- grass. ] Burkhardt, C. C. 1974. Grass bug control in grass and wheat. University of Wyoming. Agricultural Experiment Station. Research Journal No. 82. 2 pp. [Damage to crested wheatgrass in eastern Wy- oming by Labops hesperius is reported with bug densities reaching 100 per ft?. A number of dif- ferent compounds were applied to research test plots of grasses and winter wheat. Good control was achieved with a variety of compounds but many did not yet have label approval.] Burkhardt, C. C., J. M. Edwards, and L. E. Bennett. 1986. Chemical control of grass bugs on crested wheatgrass. University of Wyoming, Laramie. College of Agriculture. Annual Research and Ex- tension Centers Progress Report No. 885. 170- 172. [Seven insecticides were tested against La- bops hesperius on crested wheatgrass in Wyoming; most gave 92—100% control on three and seven- day post treatment sampling. ] Burmeister, H. C. C. 1835. Handbuch der Entomo- logie. Tome 2, Abtheil 1: i-xii, 1-400. T. Enslin, Berlin. [This paper contains the original descrip- tion of Labops based upon L. diopsis Burmeister [a junior synonym of Capsus sahlbergi Fallén, monotypic (Henry and Wheeler 1988)].] Campbell, W. F., B. A. Haws, K. H. Asay, and J. D. Hansen. 1984. A review of black grass bug re- sistance in forage grasses. Journal of Range Man- agement 37: 365-369. [Several mechanisms of re- sistance are discussed including trichome length, non-preference, tolerance, and antibiosis. The au- thors conclude that there is potential for breeding resistance to black grass bugs in range grasses. ] Carvalho, J.C. M. 1958. Catalogo dos Mirideos do Mundo. Parte III. Subfamilia Orthotylinae. Ar- quivos do Museu Nacional, Rio de Janeiro, Brazil 47: 1-161. [Eleven species of Labops are reported from throughout the world, included are Labops brooksi, L. burmeisteri, L. chelifer, L. hesperius, L. hirtus, L. nigripes Reuter, L. sahlbergi Fallen, L. setosus Reuter, L. tumidifrons, L. utahensis, and L. verae. Relevant literature is referenced and dis- tribution given.] Carvalho, J.C. M. 1959. Catalogo dos Mirideus do Mundo. Parte IV. Subfamilia Mirinae. Arquivos do Museu Nacional, Rio de Janeiro, Brazil 48: 1- 385. [Fourteen species of /rbisia are reported from North America and the Soviet Far East: /rbisia brachycera, I. californica, I. castanipes, I. cuneo- maculata, I. elongata, I. fuscipubescens, I. mol- lipes, I. nigripes, I. pacifica, I. paeta, and I. seri- cans. (For the most recent documentation, see Schwartz 1984.)] 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Childs, L. 1914. Insect notes. California State Com- mission of Horticulture. Monthly Bulletin 3: 220. [Irbisia brachycera and I. californica (as I. serican, see Schwartz 1984, p. 234) are reported feeding on grain at Stanford University, causing consid- erable damage.] Cockerell, T. D. A. 1893. The entomology of the mid- alpine zone of Custer County, Colorado. Trans- actions of the American Entomological Society 20: 305-370. [The biogeography of the region and the impact of the topography, vegetation, and habitats on the insect fauna are discussed. Irbisia brachy- cera is reported from Westcliff, Colorado (as Cap- sus brachycorus (sic!) Uhler).] Cockerell, T. D. A. 1910. Some insects from Steam- boat Springs, Colo. II. Canadian Entomologist 42: 366-370. [Irbisia brachycera is reported from Steamboat Springs, Colorado.] Coombs, E. M. 1985. Growth and development of the black grass bug (Labops hesperius Uhler) in the State of Utah. M.S. thesis, Utah State University, Logan. 163 pp. Denning, D. G. 1948. The crested wheat bug. Uni- versity of Wyoming, Agricultural Experiment Sta- tion. Circular No. 33. 2 pp. [Labops hesperius is reported damaging crested wheatgrass in Wyo- ming and affecting the palatability of the damaged hay to cattle.] Dickerson, G. W. 1978. Control of black grass bugs (Labops hesperius Uhler) in northern New Mexico. Journal of Range Management 31: 398-399. [Tri- chlorfon, malathion, and methylparathion were used to control Labops hesperius in improved wheatgrass pastures. All provided good control; untreated plots showed up to a 50% reduction in herbage yields.] Dillman, A. C. 1946. The beginnings of crested wheatgrass in North America. Journal of the American Society of Agronomy 38: 237-250. [The introduction of crested wheatgrass, Agropyron cristatum, into North America from Siberia, in- cluding sites of original plantings from the early seed importation, is reviewed in detail.] Downes, W. 1924. New records of Hemiptera from British Columbia. Proceedings of the Entomolog- ical Society of British Columbia 21: 27-33. [La- bops hirtus is reported from Chilcotin, British Co- lumbia.] Downes, W. 1927. A preliminary list of the Heter- optera and Homoptera of British Columbia. Pro- ceedings of the Entomological Society of British Columbia 23: 1-22. [Irbisia nigripes, I. brachycera var. solani (probably J. serrata), and I. sericans, I. pacifica (as Thyrillus pacificus), Labops hirtus, L. tumidifrons, and L. burmeisteri are reported from British Columbia, Canada.]} Drake, C. J. 1922. Heteroptera in the vicinity of Cranberry Lake. Syracuse University, New York State College of Forestry, Technical Publication No. 16: 54-86. [Labops hirtus is reported from New York, including both short- and long-winged forms.] Essig, E.O. 1915. Injurious and beneficial insects of California (Second edition). California State Com- mission of Horticulture. The Monthly Bulletin. Supplement 4. 541 pp. [/rbisia brachycera, re- ported as the black plant-bug, is a common insect in California attacking many cultivated crops and native plants. Irbisia californica (see Schwartz 1989, p. 263), reported as /rbisia sericans, the lesser black plant-bug, damaged barley, oats and wheat as well as other plants. /rbisia pacifica, reported as the pacific plant-bug, fed on grasses and grains, in- cluding barley, oats, and wheat.] Essig, E.O. 1926. Insects of Western North America. Macmillan Company, New York. 1035 pp. [Six species of /rbisia are included: JI. solani, I. bra- chycera, I. californica, I. mollipes, I. setosa, and I. sericans. Irbisia solani was reported as moving into cultivated fields and gardens when the grasses dry up and attacking a wide variety of plants. /r- bisia californica and I. mollipes attacked grasses and grains. Irbisia pacifica was reported from grasses and grain.] Essig, E. O. and W. M. Hoskins. 1944. Insects and other pests attacking agricultural crops. University of California, Agricultural Extension Service. Cir- cular 87 (revised). 197 pp. [The black grass bug, Irbisia solani, and the California plant bug, J. cal- ifornica, attacked grain in California. Irbisia spp. also attacked peaches and nectarines. ] Fisher, E. M. and R. W. Every. 1969. Federal-State Cooperative Economic Insect Report. Oregon (week ending May 23, 1969). 1 p. [Labops hes- perius adults damaged brome grass, intermediate wheatgrass, pubescent wheatgrass, and orchard grass in eastern Oregon. Most of the damaged grass was on soil bank land; some stands were as old as 10 years. ] Foster, R. N., R. T. Staten, E. Miller, J. A. Henderson, J. B. Thernley, D. K. Sato, E. W. Huddleston, and R. G. Bullard. 1981. Malathion for control of black grass bugs. Insecticide and Acaricide Tests 6: 135-136. [Malathion (ULV) was applied at dif- ferent concentrations on replicated 40 acre plots in New Mexico for control of Labops hesperius and L. hirtus on Agropyron desertorum (Fisch.) Schulten 1978. All treatments were highly suc- cessful. ] Fuxa, J. R. 1975. Biological attributes and alteration of the habitat to manipulate populations of Labops hesperius Uhler (Heteroptera: Miridae). M.S. the- sis, Oregon State University, Corvallis. 118 pp. Fuxa, J. R. and J. A. Kamm. 1976a. Effects of tem- VOLUME 97, NUMBER 1 perature and photoperiod on the egg diapause of Labops hesperius. Environmental Entomology 5: 505-507. [Labops hesperius diapauses as a well- developed embryo, normally overwintering in that stage. At least 60 days exposure to 3° to 9° C plus approximately 14 days incubation at 15° C are required for hatching to occur. No hatching oc- curred if the chilling occurred in August. Approx- imately 40% hatching occurred if the chilling was delayed until September or later. The combination of increasing temperatures and day length in early spring appears to regulate egg hatch, allowing the eggs to hatch at relatively cold temperatures, an ideal adaptation to early herbage growth of host grasses. ] Fuxa, J. R. and J. A. Kamm. 1976b. Dispersal of Labops hesperius on rangeland. Annals of the En- tomological Society of America 69: 891-893. [The adult population of Labops hesperius was analyzed at Vale, Oregon, in 1974; 43% were macropterous males, 53% brachypterous females, and only 4% macropterous females. Apparently the ovaries of macropterous females do not mature until the adults have completed their migratory phase of about three weeks. Dispersal capabilities were con- sidered to be limited because of the high percent- age of brachypterous females in the population. There was a positive correlation between old growth wheatgrass and bug density.] Gates, D. H. 1969a. ABC (another bug on crested)- problem. Oregon State University. United States Department of Agriculture, and Oregon Counties Cooperating. The Grazier. No. 127: 3. [Labops hesperius and Irbisia spp. are reported damaging wheatgrass in several localities in eastern Oregon.] Gates, D. H. 1969b. Black grass bug observations in Utah. Oregon State University, United States De- partment of Agriculture, and Oregon Counties Co- operating. The Grazier. No. 127: 5-6. [This report, taken from the Cooperative Economic Insect Re- port 16(25): 596, 1966, Plant Pest Control Divi- sion, Agricultural Research Service, United States Department of Agriculture, reviews several spe- cies of black grass bugs found in Utah: /rbisia brachycera, Irbisia spp., Labops hesperius, L. hir- tus, and L. utahensis. Damage was reported on planted crested wheatgrass, intermediate wheat- grass, other wheatgrasses and miscellaneous grass species. Labops spp. caused more damage at higher elevations and /rbisia spp. at lower elevations. Barley was attacked in Sanpete County by /rbisia spp., aS was rye in many areas of Utah. Seed dam- age to crested and intermediate wheatgrass was also common. It was felt that summer rains would result in additional grass growth, reducing the im- pact of the bugs’ feeding.] Gates, D. H. 1969c. Wheatgrass bugs (Labops spp.). 99 Oregon State University, United States Depart- ment of Agriculture, and Oregon Counties Co- operating. The Grazier. No. 127: 4-5. [This report was abstracted from the Cooperative Economic Insect Report, Plant Pest Control Division, Ag- ricultural Research Service, United States De- partment of Agriculture. General economic im- portance of these bugs is reported from western United States where damage to crested and inter- mediate wheatgrass occurred. Distribution infor- mation by state and province (Canada) is given. Life history information is given and the bug brief- ly described. ] Gibson, A. 1910. The entomological record, 1909. Entomological Society of Ontario, Fortieth An- nual Report: 110-128. [Labops burmeisteri is re- ported from Ontario, Canada; this is the first pub- lished record of this native species in North America.] Gibson, A. 1913. The entomological record, 1912. Entomological Society of Ontario, Forty-third An- nual Report: 113-140. [Irbisia brachycera and I. sericans are reported from the Queen Charlotte Islands, Canada.] Gillette, C. P. and C. F. Baker. 1895. A preliminary list of the Hemiptera of Colorado. (Colorado) State Agricultural College, Agricultural Experiment Sta- tion, Bulletin No. 31, Technical Series No. 1. 137 pp. [Labops hesperius is reported from several lo- calities in Colorado. /rbisia brachycera and I. pa- cifica are also reported (as Thyrillus brachycerus and 7. pacificus).] Gray, A. M. 1975. Nutritional quality and herbage production of intermediate wheatgrass (Agropyron intermedium (Host) Brauv.) when infested with black grass bugs (Labops hesperius Uhler). M.S. thesis, Utah State University, Logan. 42 pp. Hagen, A. F. 1976. Crested wheatgrass, L. hesperius control, 1975. Insecticide and Acaricide Tests 1: 102. [Several different chemicals, together with a control, were applied to plots in Scotts Bluff Coun- ty, Nebraska. All treatments gave highly effective results. } Hagen, A. F. 1982. Labops hesperius (Hemiptera: Miridae) management in crested wheatgrass by haying: An eight-year study. Journal of Economic Entomology 75: 706-707. [Annual harvest of a crested wheatgrass field significantly reduced the populations of Labops hesperius compared to a similar field that was not harvested.] Hall, I. M. 1959. The fungus Entomophthora erupta (Dustan) attacking the black grass bug, /rbisia so- lani (Heidemann) (Hemiptera, Miridae), in Cali- fornia. Journal of Insect Pathology 1: 48-51. [This fungus caused an epizootic at Riverside, Califor- nia. The fungus was previously known only from the mirids Lygus communis var. novascotiensis 100 Knight and Plagiognathus sp. Illustrations of the infected /rbisia are included together with a gen- eral description of the appearance of infected bugs.] Hansen, J. D. 1986. Differential feeding on range grass seedlings by Irbisia pacifica (Hemiptera: Mir- idae). Journal of the Kansas Entomological Society 59: 199-203. [Eighteen range grasses were tested for feeding preference by J. pacifica. Leymus ci- nereus (Scrib. & Merr.) Love was the species most preferred and Psathyrostachys juncea (Fisch.) Nevski and Agropyron fragile (Roth) Candargy, the least preferred.] Hansen, J. D. 1987. Feeding site selection by /rbisia pacifica (Hemiptera: Miridae) on four cool-season western range grasses. Journal of the Kansas En- tomological Society 60: 316-323. [Feeding sites by J. pacifica were examined on Great Basin wild- rye, Leymus cinereus (Scrib. & Merr.) Love, a crested wheatgrass hybrid and a hybrid between quack grass and blue bunch wheatgrass. Feeding density was greatest at the leaf apex and least at the base.] Hansen, J. D. 1988. Field observations of /rbisia pacifica (Hemiptera: Miridae): Feeding behavior and effects on host plant growth. Great Basin Nat- uralist 48: 68-74. [The interaction of J. pacifica with intermediate wheatgrass, Thinopyrum inter- medium (Host) Barkw. and D. R. Dewey, was ex- amined in northern Utah. Egg hatch began in April; ovarian development was completed by mid-June. The second and third leaves were the preferred feeding sites.] Hansen, J. D., K. H. Asay, and D. C. Nielson. 1985a. Screening range grasses for resistance to black grass bugs Labops hesperius and Irbisia pacifica (He- miptera: Miridae). Journal of Range Management 38: 254-257. [Three crested wheatgrasses and two hybrids between quack grass and blue bunch wheatgrass were tested for resistance to feeding by these mirids. No difference in feeding preferences was noted. Tolerance is suggested as a possible mechanism of resistance. ] Hansen, J. D., K. H. Asay, and D. C. Nielson. 1985b. Feeding preference of a black grass bug, Labops hesperius (Hemiptera: Miridae), for 16 range grass- es. Journal of the Kansas Entomological Society 58: 356-359. [Sixteen grass species and hybrids were tested for feeding preference by L. hesperius. Dactylis glomerata L., Phalaris arundinacea L. and Pascopyrum smithii (Rydb.) Love were among the least preferred and crested and intermediate wheatgrasses the most preferred.]} Hansen, J. D. and R. S. Nowak. 1988. Feeding dam- age by Irbisia pacifica (Hemiptera: Miridae): Ef- fects of feeding and drought on host plant growth. Annals of the Entomological Society of America 81: 599-604. [Green leaf area per tiller was re- duced by two-thirds in intermediate wheatgrass, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON for Great Basin wildrye (GBWR), the reduction was one-half. Drought conditions further reduced green leaf area on bug-infested GBWR but did not damage the previously affected intermediate wheatgrass. ] Hardee, D. D., H. Y. Forsythe, Jr., and G. G. Gyrisco. 1963. Asurvey of the Hemiptera and Homoptera infesting grasses (Gramineae) in New York. Jour- nal of Economic Entomology 56: 555-559. [La- bops hirtus is recorded from grass fields in New York.] Harling, J., J. M. Snyder, and D. M. Coletti. 1977. Insects collected from an alpine-sub alpine region in southeast British Columbia. Journal of the En- tomological Society of British Columbia 74: 34— 36. [Irbisia nigripes is reported from the Selkirk Mountains in southeastern British Columbia, Canada. ] Harrington, W. H. 1892. Fauna Ottawaensis, He- miptera. Ottawa Naturalist 6: 25-32. [Labops hes- perius is reported from the vicinity of Ottawa, Canada, and the long-winged form noted as rare.] Haws, B. A. 1972. Preliminary report on black grass bug Labops hesperius Uhler. United States De- partment of Agriculture, Forest Service, Inter Mountain Region, Range Improvement Notes 17: 1-3. [The Black Grass Bug project at Utah State University was established in 1971. Full obser- vations on the bug showed early seasonal growth of range grasses and concomitant development of early instar bugs, which were active at low air temperatures (22° F). There was evidence that con- siderable damage to the grass by bugs occurred early in the season.] Haws, B. A. (Compiler). 1978. Economic impacts of Labops hesperius on the production of high quality range grasses. Utah State University, Agricultural Experiment Station. Final Report to Four Corners Regional Commission. 267 pp. [The environmen- talimpact of L. hesperius, upon range grasses, chief- ly crested and intermediate wheatgrass, is dis- cussed. Other arthropods, including some natural enemies of L. hesperius, are reported.] Haws, B. A. 1979. Something is eating more grass that our livestock. Rangelands 1: 135-138. [This popular account of range insects includes several references to Labops hesperius and its damage to grasses. ] Haws, B. A. (Compiler). 1982a. An introduction to beneficial and injurious rangeland insects of the western United States. Utah State University, Utah Agriculture Experiment Station. Special Report 23. 64 pp. [This general introduction to the insects found on western rangeland includes many pho- tographs of Labops hesperius and Irbisia pacifica and their damage. The role of L. hesperius as a rangeland pest is discussed. ] VOLUME 97, NUMBER 1 Haws, B. A. (Compiler). 1982b. Rangeland improve- ment; Demonstration Project. FCRC No. 602-466- 080-4. Final Report of Five States to the Four Corners Regional Commission (Arizona, Colora- do, Nevada, New Mexico, Utah). Utah State Uni- versity, Logan. 282 pp. [A summary of coopera- tive work involving Nevada, Utah, Colorado, New Mexico and Arizona on Labops hesperius, L. hir- tus, L. utahensis, Irbisia brachycera and I. pacifica is included. Plantings of crested wheatgrass along highways were suggested to provide “bug free- ways”’ for dispersal of the insects.] Haws, B. A. 1986. The status of IPM strategies for controlling grass bugs infesting introduced grass- land monocultures, pp. 67—72. Jn Onsager, J. A., ed., Integrated Pest Management on Rangeland. United States Department of Agriculture, Agri- cultural Research Service. ARS-5O. [Strategies for managing grass bugs include chemical control, plant resistance, burning, grazing, planting heterocul- tures, and biological control. Future work on re- ducing the grass monocultures is suggested.] Haws, B. A. and G. E. Bohart. 1986. Black grass bugs (Labops hesperius Uhler) (Hemiptera: Miridae) and other insects in relation to crested wheatgrass, pp. 123-145. In Johnson, K. L., ed., Crested Wheat- grass: Its Values, Problems and Myths: Sympo- sium Proceedings. Utah State University, Logan. [The insects, native and introduced, range grasses, and the problems of management strategies to minimize the impact of insect damage are re- viewed. Basic biology of the bugs is compared to phenology of the wheatgrass. Many possible con- trol strategies are discussed. ] Haws, B. A.,D. D. Dwyer, and M.G. Anderson. 1973. Problems with range grasses? Look for black grass bugs! Utah Science 34: 3-9. [Information on the recognition, distribution, life history of /rbisia pa- cifica, Labops hesperius, L. hirtus, and L. utah- ensis, their damage, and management and control is included.] Haws, B. A., C. M. McKell, and J. Malechek. 1976. Do insects affect the validity of basic assumptions of grazing management? Society for Range Man- agement, 29th Annual Meeting, Abstract, p. 28. [Some reproductive structures of crested wheat- grass developed despite a heavy concentration La- bops hesperius (900 per sweep). Undergrazed rang- es had heavier bug infestations because of increased dead grass and plant debris, which provided sites for egg deposition. ] Hayward, C. L. 1948. Biotic communities of the Wa- satch chaparral, Utah. Ecological Monographs 18: 473-506. [Irbisia brachycerus is reported from the herb layer of the lower chaparral of the Wasatch Mountains, Utah.] Heidemann, O. 1900. Papers from the Harriman 101 Alaska Expedition. XIII. Entomological Results (7): Heteroptera. Proceedings of the Washington Academy of Sciences 2: 503-506. [/rbisia sericans, originally described from Sitka, Alaska, is record- ed from eight locations in Alaska during June and July in both macropterous and brachypterous forms. A collection record from near Hood River, Oregon, is included.] Heidemann, O. 1910. Description of a new capsid. Proceedings of the Entomological Society of Washington 12: 200-201. [I/rbisia solani is de- scribed (as Capsus solani) from Walnut Creek, California, with other specimens from California, Washington, and Utah. It was found on Lupinus sp. in California and injured potato plants.] Henry, T. J. and A. G. Wheeler, Jr. 1988. Miridae, pp. 25-507. In Henry, T. J., and R. C. Foreschner, eds., Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden. 958 pp. [Twenty-three species of /r- bisia are included. Only a single species, J. seri- cans, extends from the Pacific Northwest into east- ern Asia. Eight species of Labops are cataloged, including L. burmeisteri, a species known to occur in northern North America and west into the east- ern Palearctic Region.] Herms, W.B. 1926. An analysis of California’s major entomological problems. Journal of Economic En- tomology 19: 262-270. [Irbisia solani migrated from grass as it dried to feed on the California buckeye, Aesculus californica Nuttall. Droplets of plant sap exuding from the punctures made by the bugs were fed upon by bees.] Hewitt, G. B. 1975. Grass bugs may be present on your rangeland. Society for Range Management, International Mountain Section, Newsletter No. 1: 3. [This brief review of the presence of Labops and /rbisia on range grasses and grain includes a history of their occurrence and effect upon the vegetation. A chemical control method is cited from the publication by C. C. Burkhardt (1974).] Hewitt, G. B. 1980. Tolerance of ten species of Agro- pyron to feeding by Labops hesperius. Journal of Economic Entomology 73: 779-782. [Cultivars of ten different species and varieties of the wheat- grasses, Agropyron spp., were screened for feeding tolerance to L. hesperius and Capsus simulans Stal at Bozeman, Montana. Infested and non infested plants were compared for number of culms per plant, percentage of abnormal seed heads, seed production, seed weight, seed germination, forage production, and percentage leaf damage. Feeding reduced the number and weight of seeds, and the percentage of germination of most grass species. Silver top was recorded only in infested cages. Tall, slender, intermediate, and pubescent wheatgrasses appeared somewhat tolerant to grass bug feeding; 102 however, grass bugs preferred introduced wheat- grasses over native vegetation.] Hewitt, G. B. and W. H. Burleson. 1975. Arthropods associated with two crested wheatgrass pastures in central Montana. Journal of Range Management 28: 301-304. [ Two crested wheatgrass (Agropyron desertorum) (Fisch.) Schult.) pastures were sur- veyed for arthropods in 1972 and 1973. Although it was expected both /rbisia spp. and Labops spp. would be abundant, only 21 specimens of /rbisia and no Labops were collected during the two-year study.] Hewett, G. B. and W. H. Burleson. 1976. An inven- tory of arthropods from three rangeland sites in central Montana. Journal of Range Management 29: 232-237. [Three sites in central Montana were sampled in 1972 and 1973: a mountain, foothill, and plains rangeland. Seven orders of arthropods formed the bulk of the material collected. Labops hesperius was the second most abundant insect at the mountain site but was not collected at the other two sites; L. brooksi was taken in small numbers at the foothill site.] Hewitt, G. B., E. W. Huddleston, R. J. Lavigne, D. N. Veckert, and J. G. Watts. 1974. Rangeland En- tomology. Society for Range Management, Range Science Series No. 2. 127 pp. [This review of suck- ing insects of grasses and forbs in rangeland in- cludes /rbisia spp. and Labops spp. and brief dis- cussion of their damage, biology, and control.] Higgins, K. M. 1975. The effects of the black grass bug Labops hesperius Uhler, on several native and introduced grasses. M.S. thesis, Utah State Uni- versity, Logan. 82 pp. Higgins, K. M., J. E. Bowns, and B. A. Haws. 1977. The black grass bug (Labops hesperius Uhler): Its effect on several native and introduced grasses. Journal of Range Management 30: 380-384. [Six introduced grass species were more susceptible to damage by L. hesperius than native range grasses. Slender and intermediate wheatgrasses and Ken- tucky bluegrass were the most susceptible.] Homan, H. W. 1977. Labops grass bug, Labops utah- ensis. University of Idaho, Cooperative Extension Service. Idaho’s Insect Reporter, p. 33. [Labops utahensis caused minor damage to a grain field near Driggs, Idaho.] Horning, D. S., Jr. and W. F. Barr. 1970. Insects of Craters of the Moon National Monument, Idaho. University of Idaho, College of Agriculture, Mis- cellaneous Series No. 8. 118 pp. [/rbisia pacifica is reported from this area.] Jensen, F. 1971. Reseeding and Labops. United States Department of Agriculture, Forest Service, Inter Mountain Region. Range Improvement Notes 16: 6-9. [Severe damage to range grasses by L. hes- perius is reported in Dixie National Forest, Utah, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON where the areas have been plowed or otherwise disturbed and then seeded with crested wheat- grass, intermediate wheatgrass, and smooth brome. Large numbers of Labops were found on giant wild ryegrass. Several treatments were used on the range prior to reseeding, plowing, cabling, and chaining. Pesticide control of Labops was ineffective; crea- tion of a balanced plant community rather than a monoculture was recommended.] Jensen, F. 1973. Reseeding and Labops. Society for Range Management, 26th Annual Meeting Ab- stract, p. 23. [Labops hesperius inhabits much of the reseeded rangelands on the Dixie National Forest, Utah. Population densities and resulting damage are related to plant community diversity. Densities were greatest in plowed rangelands that had been seeded to virtual monocultures of crested wheatgrass, intermediate wheatgrass, or smooth brome. Planting for a more diversified plant com- munity is suggested, including the conservation of native grasses. ] Kamm, J. A. and J. R. Fuxa. 1977. Management practices to manipulate populations of the plant bug Labops hesperius Uhler. Journal of Range Management 30: 385-387. [Populations on wheat- grass increased with nitrogen fertilization; no re- sponse was shown to applications of phosphorus and potassium. Early curing of the herbage with paraquat, mechanical removal of the herbage, and heavy spring grazing all reduced bug populations in spring and summer. Grassbug survival is en- hanced by pastures with adequate oviposition sites, winter protection, and habitat.] Kamm, J. A. and R. R. Robinson. 1974. Labops, a plant bug, on Oregon rangeland. Oregon State University, Extension Service, Fact Sheet 211. 2 pp. [Labops hesperius is reported on native range- land seeded with introduced wheatgrasses in east- ern Oregon. Many different grasses, introduced and native, are recorded as host plants. The general life history is described. One hundred and twenty bugs per ft? reduced the nutritive value of inter- mediate wheatgrass by 18% midway in the growing season. This loss decreased to 23% if the rangeland was reserved for fall pasture. Greatest losses occur during severe summer drought.] Kamm, J. A., F. A. Sneva, and L. M. Rittenhouse. 1978. Insect grazers on the cold desert biome. Proceedings of the First International Rangeland Congress, D. N. Hyder (ed.), Society for Range Management, Denver, Colorado, pp. 479-483. [Population levels of Labops hesperius increased in response to widespread planting of introduced wheatgrasses. Increased use of resistant grass va- rieties could have occurred if the pest potential of this bug had been recognized.] Kelton, L.A. 1959. Male genitalia as taxonomic char- VOLUME 97, NUMBER 1 acters in the Miridae (Hemiptera). Canadian En- tomologist (Supplement | 1) 91: 1-72. [This review of male genitalia structure in mirids includes /r- bisia pacifica, I. sericans, Labops hesperius, L. hir- tus and L. sahlbergi.] Kelton, L. A. 1980. The insects and arachnids of Canada. Part 8. The plant bugs of the Prairie Prov- inces of Canada. Heteroptera: Miridae. Agricul- ture Canada. Publication 1703. 408 pp. [This re- view of the mirids of west-central Canada includes a brief description of each species with distribution maps and host information. Keys are given to al- low identification of each species. Some species (Labops hirtus) are illustrated with habitus draw- ings. The genus /rbisia (pp. 83-86) includes the four species found in the Prairie Provinces (J. bra- chycera, I. elongata, I. fuscipubescens, and I. ni- gripes). The genus Labops (pp. 189-194) includes L. brooksi, L. hesperius, L. hirtus, L. tumidifrons, and L. verae.] Kirkaldy, G. W. 1906. List of the genera of the pa- giopodus Hemiptera-Heteroptera, with their type species, from 1758 to 1904 (and also of the aquatic and semi-aquatic Trochalopoda). Transactions of the American Entomological Society 32: 117-156. [This important paper lists the type species of the genera of several families, including /rbisia and Labops.] Knight, H. H. 1918. Synoptic key to the subfamilies of Miridae (Hemiptera-Heteroptera). Journal of the New York Entomological Society 26: 40-44. [Details of the claw of Labops hesperius are illus- trated.] Knight, H. H. 1921. Scientific results of the Katmai expeditions of the National Geographic Society. XIV. Hemiptera of the family Miridae. Ohio Jour- nal of Science 21: 107-112. [/rbisia sericans is reported from Katmai, Alaska, where it was abun- dant and feeding on rye grass. Described originally from Sitka, Alaska, it is the only species of /rbisia to occur in the Old World (Siberia).] Knight, H. H. 1922. The North American species of Labops (Heteroptera-Miridae). Canadian Ento- mologist 54: 258-261. [Three species are consid- ered: L. hesperius, L. hirtus, and L. tumidifrons, the latter two described as new. Labops hirtus is described from New York with other specimens from across northern United States and southern Canada. Labops tumidifrons is described from British Columbia, Canada. Some notes are in- cluded on the taxonomic value of the shape of the male clasper.] Knight, H. H. 1923. Family Miridae (Capsidae), pp. 422-658. In Britton, W. E., ed., Guide to the In- sects of Connecticut. Part IV. The Hemiptera or Sucking Insects of Connecticut. State of Connect- icut, State Geological and Natural History Survey, 103 Bulletin No. 34. 807 pp. [A brief description of Labops hirtus and state records from Maine, Mas- sachusetts, and New York are given.] Knight, H. H. 1925a. Descriptions of a new genus and eleven new species of North American Mir- idae (Hemiptera). Canadian Entomologist 57: 89- 97. [Irbisia nigripes is described as new from Troy, Idaho, with additional specimens from Alberta, British Columbia, Canada, and Montana.] Knight, H. H. 1925b. A list of Miridae and Antho- coridae from Alberta, Canada (Hemiptera). Ca- nadian Entomologist 57: 181-182. [Labops hes- perius is reported from Nordegg, Alberta, Canada, and /rbisia nigripes from Waterton Lakes in the same province.] Knight, H. H. 1926. Capsus simulans (Stal) and La- bops burmeisteri Stal recognized from the Nearctic Region. (Hemiptera, Miridae). Canadian Ento- mologist 58: 59-60. [Labops burmeisteri is re- ported from the Abitibi Region of northern On- tario, Canada. This species was described originally from Siberia. Although there had been earlier re- cords of this species from North America, Knight was unable to locate any specimens.] Knight, H. H. 1927. On the Miridae in Blatchley’s “Heteroptera of Eastern North America.” Bulletin of the Brooklyn Entomological Society 22: 98- 105. [Blatchley’s concept of Labops hesperius, which includes species variation associated with geographic distribution, is discussed.] Knight, H. H. 1928. Family Miridae, pp. 110-134. In Leonard, M. D., ed., A List of the Insects of New York. Cornell University, Agricultural Ex- periment Station, Memoir 101. 1121 pp. [Local- ities of Labops hirtus for the state of New York are listed.] Knight, H.H. 1929a. Labops verae, new species, with Labopella, Nicholia and Pronotocrepis, new genera of North American Miridae (Hemiptera). Cana- dian Entomologist 61: 214-218. [A new high-al- titude, high-latitude species of Labops, L. verae, is described from Mt. Rainier, Washington, and Banff, Alberta.] Knight, H. H. 1929b. Rectifications for Blatchley’s “Heteroptera” with the description of a new spe- cies (Hemiptera). Bulletin of the Brooklyn Ento- mological Society 24: 143-154. [The identity of the Labops species treated by Blatchley (1926) is clarified. ] Knight, H. H. 1941la. New species of /rbisia Reuter (Hemiptera, Miridae). Bulletin of the Brooklyn Entomological Society 36: 75-79. [Three new spe- cies of /rbisia are described from western North America (/. elongata, I. fuscipubescens and I. shul- /i). Distributional information is included for each new species and for J. brachycera.] Knight, H. H. 1941b. The plant bugs, or Miridae of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Illinois. Illinois Natural History Survey, Bulletin. Volume 22: 1-234. [No Labops species were re- ported from Illinois; L. hirtus is reported from Colorado, Maine, Massachusetts, Montana, New York, and Ontario, Canada.] Knight, H. H. 1968. Taxonomic review: Miridae of the Nevada Test Site and the western United States. Brigham Young University, Science Bulletin, Bi- ological Services 9: 1-282. [This review of mirids found in western United States includes descrip- tions of many new species and keys for the iden- tification of many of the included species. Five species of /rbisia are briefly discussed and keyed (I. brachycera, I. elongata, I. nigripes, I. pacifica, and /. shulli). Information is largely restricted to collection localities. Labops hesperius, L. hirtus, and L. utahensis are listed with state distribution records only.] Knight, J. B. 1982. An initial survey of the insects associated with five grassland sites in central Utah. M.S. thesis, Utah State University, Logan. 81 pp. Knight, J. B. 1986. Range insects—pests and bene- ficials, pp. 73-75. Jn Onsager, J. A., ed., Integrated Pest Management on Rangeland. United States Department of Agriculture, Agricultural Research Service. ARS-50. [A brief review of the pest insects found on rangeland, including Homoptera, He- miptera, Lepidoptera, Diptera and Hymenoptera, is followed by general remarks on biological con- trol agents, chiefly of weeds.] Knowlton, G. F. 1931. Notes on Utah Heteroptera and Homoptera. Entomological News 42: 68-72. [[rbisia brachycerus var. solani Heidemann and J. pacifica (as Thyrillus pacificus) are reported from several localities in Utah.] Knowlton, G. F. 1932. Notes on Utah Hemiptera. Canadian Entomologist 64: 166-167. [Labops hir- tus 1s reported from Providence, Utah.] Knowlton, G. F. 1945. Labops damage to range grass- es. Journal of Economic Entomology 38: 707-708. [Labops tumidifrons fed in large numbers on giant wild rye, Elymus condensatus Presl., in Utah. Oth- er specimens were collected on bunch grass and large bunch grass at several sites in Utah and Ida- ho. Labops hesperius was collected on meadow grass and at several other localities in Utah on smooth broom. Labops hirtus was taken on range grasses in Utah and Idaho.] Knowlton, G. F. 1951. Bugs damage grass in Utah. Bulletin of the Brooklyn Entomological Society 46: 74-75. [Irbisia pacifica (as Thyrillus pacificus) damaged giant rye grass Timothy, blue grass brome, and several other grasses in Morgan County, Utah. Other Hemiptera found on the grasses were La- bops hirtus and Slaterocoris atritibialis (Knight) (as Strongylocoris atritibialis).] Knowlton, G. F. 1955a. Hemiptera of Utah—re- cords. Utah State Agricultural College, Extension Service, Mimeograph Series No. 140. 15 pp. [/r- bisia arcuata, I. brachycera, I. solani and Labops hesperius, L. hirtus, and L. utahensis are reported from Utah.] Knowlton, G. F. 1955b. Some Hemiptera and Ho- moptera of Utah—1955. Utah State Agricultural College, Extension Service, Mimeograph Service No. 145. 9 pp. [/rbisia elongata is reported from Utah.] Knowlton, G. F. 1966a. Insect conditions in Utah— 1966. Utah State University, Cooperative Exten- sion, Entomology Mimeo Series No. 114. 4 pp. [Approximately 200,000 acres were damaged by Labops hesperius, L. utahensis, Irbisia brachycera, I. shulli, and I. pacifica in Utah during 1966.] Knowlton, G. F. 1966b. Grass bugs, range and crop pests in Utah. Utah State University, Cooperative Extension, Entomology Mimeo Series No. 119. 5 pp. [This brief general discussion of the extent and range of damage to grasses and crops in Utah by mirids gives detailed distributional information for 15 species including Labops hesperius, L. hir- tus, L. utahensis, Irbisia brachycera, I. pacifica, and J. shulli.] Knowlton, G. F. 1967. Grass bugs: A serious range problem in 1966. Utah Academy of Sciences, Arts and Letters 43: 20-21. [Grass bugs extensively damaged crested wheatgrass, intermediate wheat- grass, giant rye grass, and other planted and native grasses in Utah. The species included /rbisia bra- chycera, I. pacifica, I. shulli, Labops hesperius, L. utahensis, Leptopterna ferrugata (Fallén), Steno- dema pilosipes Kelton, S. vicinum (Provancher), and Trigonotylus dohertyi (Distant). At least 200,000 acres were damaged, especially at higher elevations. Wheat, barley, and rye were also dam- aged.] Knowlton, G. F. 1973. Some Hemiptera of Curlew Valley. Utah State University, Terrestrial Arthro- pod Series No. 5. 8 pp. [A partial list of the He- miptera of the Utah portion of Curlew Valley in- cludes records of Jrbisia brachycera on crested wheat and giant rye grasses and /. brachycerus so- lani (probably J. serrata) (no hosts given) and J. pacifica on crested wheatgrass and giant ryegrass. Labops hesperius was reported from crested wheatgrass at Cedar Creek, Utah.] Knowlton, G. F. and F. C. Harmston. 1940. Utah insects. Hemiptera. Utah State College, Utah Ag- ricultural Experiment Station, Mimeograph Series 200 (Technical). Part 5. 10 pp. [Labops hesperius and L. hirtus are reported from several localities in Utah as are Irbisia brachycera, I. brachycera solani, I. mollipes, I. arcuata, and I. nigripes.] Kumar, R., R. J. Lavigne, J. E. Lloyd, and R. E. Pfadt. 1976. Insects of the Central Plains Experiment VOLUME 97, NUMBER 1 Range, Pawnee National Grassland. University of Wyoming, Agricultural Experiment Station, Sci- ence Monograph 32. 74 pp. [Labops hesperius 1s reported from the Central Plains Experimental Range (Pawnee National Grasslands) in Colora- do.] La Follette, R. A. 1915. Preliminary list of common Heteroptera from the Claremont-Laguna region. Journal of Entomology and Zoology 7: 123-129. [Irbisia politus Uhler is reported from Claremont, California and nearby localities. (This must be a mistake; there is no combination, /rbisia politus Uhler, and one species is described as “‘very dark olive green, legs yellowish brown.”’)] Lange, W. H., Jr. 1941. The artichoke plume moth and other pests injurious to the globe artichoke. University of California, Agricultural Experiment Station, Bulletin 653. 71 pp. [/rbisia solani is re- ported as damaging artichokes; the report is at- tributed to Tavernetti (1933).] Larochelle, A. 1984. Les punaises terrestres (Hém- iptéres: Geocorises) du Québec. Fabreries, Sup- plement 3. 1-513. [Irbisia sericans, Labops hes- perius, L. hirtus, and L. burmeisteri are reported from Quebec, Canada. An extensive bibliography on the hemipteran fauna of Quebec is included with keys for identification. (The record for J. ser- icans has not been duplicated (Schwartz 1984, p. 263).)] Larochelle, A. and M. C. Lariviere. 1979. Le genre Labops Burmeister du Québec, Canada (Heter- optera: Miridae): Repartition geographique, hab- itat et biologie. Bulletin d’inventaire des insectes du Québec 1(4): 61-67. [This review of Labops spp. found in the province of Quebec, Canada, includes Labops burmeisteri and L. hirtus. Labops hesperius, reported from Quebec by early authors, is removed from the faunal list because no spec- imens could be located to document its occur- rence.] Leonard, M. D. (ed.). 1928. A list of the insects of New York. Cornell University, Agricultural Ex- periment Station. Memoir 101. 1121 pp. [Labops hirtus is reported from several localities in New York State.] Lindsay, H. G. 1970. A serious threat: Black grass bugs. Utah Farmer. August 6, 1970, p. 12. [This article discusses black grass bugs in Utah, es- pecially Labops spp. Details on life history, dam- age, and possible chemical control methods are given and the need for research on this problem is stated.] Ling, Y. H. 1982. Scanning electron microscopic (SEM) studies on range grasses and their resistance to black grass bugs. M.S. thesis, Utah State Uni- versity, Logan. 94 pp. Ling, Y. H., W. F. Campbell, B. A. Haws, and K. H. 105 Asay. 1985. Scanning electron microscope (SEM) studies of morphology of range grasses in relation to feeding by Labops hesperius. Crop Science 25: 327-332. [Selected grass varieties were subjected to similar bug densities in rearing chambers. Re- sults suggested that selecting clones of wheatgrass- es with large trichomes should confer some resis- tance to this insect.] Loan, C. C. 1965. Life cycle and development of Leiophron pallipes Curtis (Hymenoptera: Bracon- idae: Euphorinae) in five mirid hosts in the Belle- ville district. Proceedings of the Entomological So- ciety of Ontario 95: 115-121. [The braconid parasite Leiophron pallipes was reared from La- bops hirtus near Belleville, Ontario, Canada. The parasite larva emerged from the adult bug and then pupated, spinning a silken cocoon. The adult wasp emerged the following year.] Loan, C.C. 1980. Plant bug hosts (Heteroptera: Mir- idae) of some euphorine parasites (Hymenoptera: Braconidae) near Belleville, Ontario, Canada. Na- turaliste Canadien 107: 87-93. [The braconid par- asite Peristenus pallipes (Curtis) was reared from Labops hirtus collected near Belleville, Ontario, Canada, as well as from four other mirid species.] Lockwood, S. 1933. Insect and mite scars of Cali- fornia fruits. California Department of Agricul- ture. Monthly Bulletin 22: 319-345. [/rbisia solani was suspected of causing scarring of peaches, es- pecially in orchards close to grassy or weedy slopes. The bugs became very abundant in the orchards after the surrounding vegetation dried up.] Lockwood, S. 1937. Farm sanitation aids pest con- trol. California Cultivator 84: 537, 557. [Black grass bugs (no scientific name given) are reported as a pest of cultivated crops, especially peaches, moving to these plants as the grasses dried up.] Lockwood, S. and E. T. Gammon. 1949. Incidence of insect pests. California Department of Agricul- ture. Bulletin 38: 190-203. [/rbisia sp. is reported cat-facing peaches, cherries, and plums in parts of Riverside County, California. DDT was used for control.] MacGillivray, A. D. and C. O. Houghton. 1903. A list of insects taken in the Adirondack Mountains, N.Y.—III. Entomological News 14: 262-265. [La- bops hesperius is reported from the Adirondack Mountains of New York.] Malechek, J. C., A. M. Gray, and B. A. Haws. 1977. Yield and nutritional quality of intermediate wheatgrass infested by black grass bugs at low pop- ulation densities. Journal of Range Management 30: 128-131. [Populations of Labops hesperius at 156 bugs per m? did not affect herbage yields of intermediate wheatgrass in Utah. Seed head pro- duction was reduced 56%, resulting in a slight in- 106 crease in crude protein and a small decrease in cell content.] Markgraf, P. M. 1974. Effects of wheatgrass bug in- festation on range grasses. Society for Range Man- agement, 27th Annual Meeting. Abstract, p. 31. [Labops hesperius was studied on soil bank lands seeded to wheatgrasses in northeast Oregon. Feed- ing by mature nymphs and adults was heavy in May, reducing current annual growth of air-dry herbage by 13%. The quality of the resulting fo- liage was 5% less digestible. Although chemical control of the bugs is possible, resource manage- ment to reduce litter and straw accumulation was considered more economical.] McAtee, W. L. 1923. Heteroptera. Jn A biological survey of the Pribilof Islands, Alaska, Part II. In- sects, arachnids, and chilopods. North American Fauna No. 46. p. 145. [/rbisia sericans is reported from the St. Paul Island and St. George Island, Alaska, from June to September.] McKendrick, J. D. and D. P. Bleicher. 1980. Obser- vations of a grass bug on bluejoint ranges. Agro- borealis 12: 15-18. [A native bluejoint reedgrass (Calamagrostis canadensis (Michx.) Beauv.) was damaged by /rbisia sericans near Homer, Alaska. The range of this bug includes parts of eastern Siberia and Alaska south to San Francisco, Cali- fornia. Evidence showed a negative correlation be- tween insect damage and total nonstructural car- bohydrates in the grass. The possibility of some resistance to insect damage because of silica in the grass was suggested. The bug was also reported from fireweed (Epilobium angustifolium L.).] Mills, H. B. 1939. Montana insect pests for 1937 and 1938. Montana State College, Agricultural Exper- iment Station, Bulletin 366. 32 pp. [Labops hes- perius and Conostethus n. sp. were found on range grasses and winter wheat at sites in Montana. The number of bugs declined in late May and the wheat outstripped the injury. The range grasses attacked were Koeleria cristata (L.) Pers., Poa secunda Presl., Stipa comata Trin. and Rupt., Stipa williamsi Scrbn., and an unidentified species.] Mills, H. B. 1941. Montana insect pests 1939 and 1940. Montana State College, Agricultural Exper- iment Station, Bulletin 384. 28 pp. [Labops hes- perius moved into wheat near Bozeman, Montana, causing mottling of the leaves.] Moore, G. A. 1944. A list of Hemiptera taken at Hudson Heights, Quebec. Canadian Entomologist 76: 40-44. [Labops hesperius is reported from Hudson Heights, Quebec. (Note: This identifica- tion is doubtful, see Larochelle and Lariviere (1979).)] Moore, G. A. 1950. Checklist of Hemiptera of the Province of Quebec. Naturaliste Canadien 77: 233- 271. [Labops hesperius and L. hirtus are reported PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from Quebec; the record of L. hesperius is based on Van Duzee (1916c).] Osborn, H. 1893. Notes on the distribution of He- miptera. Proceedings of the lowa Academy of Sci- ences 1: 120-123. [Labops hesperius is reported from New Hampshire. This record almost cer- tainly refers to L. hirtus.] Oshanin, B. 1912. Katalog der palaéarktischen Hem- ipteren (Heteroptera, Homoptera-Auchenorhyn- cha-und Psylloideae). R. Friedlander and Sohn, Berlin. 187 pp. [This catalog of the Old World Hemiptera includes the Heteroptera and part of the Homoptera. Irbisia sericans is reported from northern Siberia and the Neartic Region. Four spe- cies of Labops are included: L. setosus from Si- beria; L. sahlbergi from Scandinavia, northern and middle Russia and Siberia; L. burmeisteri from Siberia; and L. nigripes from Siberia and Mon- golia. (Labops burmeisteri is also found in North America.)] Osman, D. H. 1979. The toxicity, metabolism and distribution of carbaryl in three species of Labops with and without piperonyl butoxide treatment (Hemiptera: Miridae). M.S. thesis, Utah State University, Logan. 85 pp. Osman, D. H. and W. A. Brindley. 1981. Estimating monooxygenase detoxification in field popula- tions: Toxicity and distribution of carbaryl in three species of Labops grass bugs. Environmental En- tomology 10: 676-680. [In tests of monooxygen- ase detoxification of carbaryl by field populations of Labops hesperius, L. hirtus, and L. utahensis, males were more susceptible than females. The three species differed in susceptibility, with L. hes- perius the most tolerant and L. utahensis the most susceptible. ] Ostlie, K. R. 1979. Labops hesperius Uhler, abun- dance and dispersal in relation to vegetation. M.S. thesis, Utah State University, Logan. 198 pp. Paraqueima, O. L. 1977. Some effects of different temperatures on the development of the black grass bug Labops hesperius Uhler, from the egg through the adult stage. M.S. thesis, Utah State University, Logan. 84 pp. Parshley, H. M. 1917. Fauna of New England. 14. List of the Hemiptera-Heteroptera. Occasional Papers of the Boston Society of Natural History 7: 1-125. [Labops hesperius is recorded from Maine, New Hampshire, Vermont, and Massa- chusetts. (Note: Henry and Wheeler (1988) suggest these records apply to L. hirtus.)] Parshley, H. M. 1919. On some Hemiptera from western Canada. University of Michigan. Occa- sional Papers of the Museum of Zoology. No. 71. 35 pp. [Irbisia brachycerus solani and I. pacificus (as Thyrillus pacificus) are reported from Vernon, British Columbia, Canada.] VOLUME 97, NUMBER 1 Parshley, H. M. 1921. A report on some Hemiptera from British Columbia. Proceedings of the Ento- mological Society of British Columbia. Systematic Series No. 18: 13-24. [Labops burmeisteri and L. hesperius are reported from Chilcotin, British Co- lumbia.] Parshley, H. M. 1922. Report on a collection of He- miptera-Heteroptera from South Dakota. South Dakota State College, Technical Bulletin No. 2. 22 pp. [/rbisia brachycera and Labops hesperius are reported from several localities in South Da- kota.] Parsons, G. L., G. Cassis, A. R. Moldenke, J. D. Lattin, N. H. Anderson, J. C. Miller, P. Hammond, and T. D. Schowalter. 1991. Invertebrates of the H. J. Andrews Experimental Forest, Western Cascade Range, Oregon. V: An annotated list of insects and other arthropods. United States Department of Agriculture, Forest Service, Pacific Northwest Re- search Station, General Technical Report. PNW- GTR-290. 168 pp. [This report documents 3454 species of insects and other arthropods from an old-growth Douglas-fir forest in western Oregon, including /rbisia cascadia (as I. inurbana Bliven) and /. serrata, from grasses in open areas in the forest. Irbisia cascadia is distributed in the Coast and Cascade of Oregon and northern California, and J. serrata occurs throughout much of western North America.] Pepper, J. H. 1962. Montana insect pests, 1961 and 1962. 39th Report of the State Entomologist. Montana State College, Agriculture Experiment Station. Miscellaneous Publication. No. 4. 8 pp. [Irbisia sp. damaged intermediate wheatgrass in Montana. |] Pepper, J. H., N. L. Anderson, G. R. Roemhild, and L. N. Graham. 1956. Montana insect pests 1955 and 1956. Montana State College, Agricultural Ex- periment Station, Bulletin 526. 27 pp. [The crested wheat plant bug, Labops hesperius, infested crested wheatgrass fields as well as wheat and barley. Sten- odema sp. damaged tall wheatgrass and crested wheatgrass in Montana.] Pepper, J. H., J. P. Corkins, L. N. Graham, D. R. Merkley, and N. L. Anderson. 1953. Montana insect pests. 1951 and 1952. Montana State Col- lege, Agricultural Experiment Station. Bulletin 484. 34 pp. [The margins of winter and spring wheat fields were damaged by Labops hesperius moving from adjacent crested wheatgrass fields. Crested wheatgrass plantings were severely damaged at several locations in Montana and were often killed when the bug infestation was accompanied by grazing. | Pepper, J. H., J. P. Corkins, R. Schmiedeskamp, C. R. Hunt, N. L. Anderson, and J. C. Wright. 1951. Montana insect pests, 1949 and 1950. Montana 107 State College, Agricultural Experiment Station. Bulletin 474. 35 pp. [Labops hesperius occurred throughout southern Montana where it severely damaged planted crested wheatgrass. ] Pepper, J. H., G. R. Roemhild, and L. N. Graham. 1954. Montana insect pests 1953-1954. Montana State College, Agricultural Experiment Station, Bulletin 504: 1-27. [Labops hesperius damaged crested wheatgrass fields in Cascade and Chouteau Counties. ] Pepper, J. H., G. R. Roemhild, and L. N. Graham. 1958. Montana insect pests, 1957 and 1958. 37th Report of the State Entomologist. Montana State College, Agricultural Experiment Station. Miscel- laneous Publication No. 2. 19 pp. [The crested wheat plant bug, Labops hesperius, damaged grass in two Montana counties. Leptopterna dolabrata (as Miris dolobratus) damaged crested wheatgrass. ] Pepper, J. H., G. R. Roemhild, and L. N. Graham. 1960. Montana insect pests, 1959-60. Montana State College, Montana Agricultural Experiment Station, Miscellaneous Publication No. 3. 11 pp. [Irbisia sp. damaged intermediate and crested wheatgrass grown for seed in Montana.] Perry, E. 1954. New ranges for old. Western Live- stock Journal, 32 (July, 1954): 26-27, 39. [This article is a nontechnical review of the use of crested wheatgrasses in western United States.] Provancher, L. 1885-1890. Petite Faune Entomo- logique du Canada. Vol. III. Cinquieme Ordre les Hémiptéres. Naturalist Canadien. 354 pp. [Labops hesperius is reported from Ontario, Canada.] Rees, N. E. and G. B. Hewitt. 1977. Effects of specific cultural practices on immediate rangeland arthro- pod populations. Montana State University, Ag- ricultural Experiment Station. Bulletin 695. 38 pp. [Seven sites in northern Montana that had re- ceived different rangeland renovation treatments were sampled for insects for three years. Treat- ments included sagebrush removal, scalping, and interseeding. Irbisia sp. (probably J. serrata) was reported from four of the seven sites. Additional grass-feeding mirids reported included Litomiris debilis (Uhler), Stenodema vicinum (Provancher), and Trigonotylus tarsalis (Reuter).] Reuter,O.M. 1879. De Hemipteris e Siberia orientali nonnullis adnotationes criticae. Ofrersigt Finska Vetenskaps-Societatens Foérhandlingar 21: 42-63. {Reuter described the genus /rbisia, type Lepto- merocoris sericans Stal. and stated that the genus was allied with Orthocephaus Fieber. Reuter also reported Labops burmeisteri from Siberia.] Reuter,O.M. 1890. Adnotationes Hemipterologicae. Revue d’Entomologique 9: 248-254. [The system- atic position of /rbisia is noted.] Reuter, O. M. 1896. Dispositio generum palaearc- ticorum divisionis Capsaria familiae Capsidae. 108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ofrersigt af Finska Vetenskaps-Societatens For- handlingar 38: 156-171. [/rbisia is included in a key to the genera of Capsidae (Miridae) of the Palearctic Region. The genus 7hyrillus is a syn- onym of /rbisia.] Schuh, R. T. 1975. The structure, distribution, and taxonomic importance of trichobothria in the Mir- idae (Hemiptera). American Museum of Natural History. Novitates No. 2585. 26 pp. [Labops bur- meisteri and L. hirtus are included in a study of the systematic value of the trichobothria (sensory setae).] Schwartz, M. D. 1981. A revision of the black grass bug genus /rbisia Reuter (Heteroptera: Miridae). M.S. thesis, Orgeon State University, Corvallis. 222 pp. Schwartz, M. D. 1984. A revision of the black grass bug genus /rbisia Reuter (Heteroptera: Miridae). Journal of the New York Entomological Society 92: 193-306. [This modern revision includes keys to 23 recognized species from throughout the range of the genus and synonyms of some names used previously. The genus is characterized and sepa- rated from related genera of Miridae. Each species is described and discussed in detail. Habitus draw- ings of J. pacifica, I. knighti and I. cascadia are included with morphological drawings of portions of each species. /rbisia bliveni and I. cascadia are described as new. Detailed information on host plant associations is included for each species as available. ]} Schwartz, M.D. and J. D. Lattin. 1983. Irbisia knigh- ti, a new mirine plant bug (Heteroptera: Miridae) from the Pacific Northwest. Journal of the New York Entomological Society 91: 413-417. [/rbisia knighti is described from the Pacific Coast of Brit- ish Columbia, Washington, Oregon and northern California. This species, which occupies a very narrow portion of the Vancouveran Zone, was found on Agropyron repens (L.) Beauv., Festuca rubra L., Holcus lanatus L., Poa pratensis L., Poa sp. and Carex sp.] Slater, J. A. 1950. An investigation of the female genitalia as taxonomic characters in the Miridae (Hemiptera). Iowa State College Journal of Science 25: 1-81. [Details of the sclerotized portions of the female genitalia of /rbisia sericans, I. shulli, 1. pacifica (as Thyrillus pacificus), Labops hesperius, and L. hirtus are described and illustrated.] Slater, J. A. 1954. Notes on the genus Labops, Bur- meister in North America, with the descriptions of three new species (Hemiptera: Miridae). Bul- letin of the Brooklyn Entomological Society 49: 57-65, 89-94. [This revision includes descriptions of each species and a key to the species.] Slater, J. A. 1974. A preliminary analysis of the der- ivation of the Heteroptera fauna of the northeast- ern United States with special reference to the fau- na of Connecticut. Connecticut Entomological Society, Memoirs, 25th Anniversary, New Haven, pp. 145-213. [A single species (L. Airtus) is re- ported in this biogeographical analysis of north- eastern Heteroptera. The genus Labops is sug- gested to have reached North America in post- Pleistocene times.] Slater, J. A. and R. M. Baranowski. 1978. How to Know the True Bugs (Hemiptera-Heteroptera). W. C. Brown Co., Dubuque, Iowa. 256 pp. [This book contains keys to the mirid genera of the United States, including Labops and Irbisia. An illustra- tion of L. hesperius and a brief discussion of its appearance and distribution are included. Some western species of Labops are mentioned as range- land pests.] Slosson, A. T. 1895. Additional list of insects taken in alpine region of Mt. Washington. Entomolog- ical News 6: 316-321. [Labops hesperius is re- ported from the alpine zone of Mt. Washington, New Hampshire. ] Spangler, S. M. 1984. Arthropod faunas of reseeded rangelands: Effects of vegetation structure. M.S. thesis, Utah State University, Logan. 77 pp. Spangler, S. M. and J. A. MacMahon. 1991. Arthro- pod faunas of monocultures and polycultures in reseeded rangelands. Environmental Entomology 19: 244-250. [Irbisia brachycera and Conostethus americanus (Knight) were the dominant mirid species occurring in grass monocultures. They were present during the early stages of growth when the root reserves of carbohydrates were the lowest. Species richness was lowest in monocultures. Di- versifying grass species mixes for reseeding is sug- gested as a means of reducing plant bug numbers and increasing their natural enemies.] Stal, C. 1858. Beitrag zur Hemipteren-Fauna Sibi- riens und des Russischen Nord-Amerika. Ento- mologische Zeitung Herausgegeben von dem En- tomologische Vereine zu Stettin 19: 175-198. [Irbisia sericans is described (in Leptomerocoris) from Sitka, Alaska. Labops burmeisteri is de- scribed from Kamchatka, Siberia.] Stephens, G. M., III. 1982. The plant bug fauna (Het- eroptera: Miridae) of grasses (Poaceae) of the Med- icine Bow Mountains and Pole Mountain Ranger District, Wyoming. University of Wyoming, Ag- ricultural Experiment Station, Science Monograph 43. 175 pp. [This analysis of the plant bug fauna found on grasses in southeastern Wyoming in- cludes Labops hesperius, L. hirtus and L. utah- ensis. A brief description of the first two species is combined with information on host grasses, grass communities, and general ecological data.] Stonedahl, G. M. and W. R. Dolling. 1991. Heter- VOLUME 97, NUMBER 1 optera identification: A reference guide, with spe- cial emphasis on economic groups. Journal of Nat- ural History 25: 1027-1066. [More than 350 references useful for identifying Heteroptera from around the world are provided.] Strickland, E. H. 1953. An annotated list of the He- miptera (S.L.) of Alberta. Canadian Entomologist 85: 193-214. [Five species of /rbisia are reported from Alberta: J. arcuata, I. brachycera, I. solani, I. nigripes and I. fuscipubescens. Two species of Labops are reported, L. hesperius and L. verae. Strickland listed Labops hirtus as a synonym of L. hesperius.} Sweet, H. E. 1930. An ecological study of the animal life associated with Artemesia californica Less, at Claremont, California. Journal of Entomology and Zoology 22: 57-70, 75-103, 121-151. [/rbisia sita is recorded from Claremont, California, from sev- eral different plant associations that included Ar- temesia. This early-spring species (March to May) was uncommon. ] Tavernetti, A. A. 1933. Production of the globe ar- tichoke in California. University of California, Berkeley. California Agricultural Extension Ser- vice, Circular 76. 24 pp. [/rbisia solani is reported as a pest of the globe artichoke in California, the bugs moving onto the crop when the host grasses dried up.]} Thomas, D. B.and F.G. Werner. 1981. Grass feeding insects of the western ranges: An annotated check- list. The University of Arizona, Agricultural Ex- periment Station. Technical Bulletin No. 243. 50 pp. [Some host and distribution information is provided for Irbisia brachycera, I. oreas, I. paci- fica, I. serrata, I. shulli, I. solani, Labops hesperius, L. hirtus, and L. utahensis. Leptopterna ferrugata fed on seed heads of Agropyron sp.] Todd, J. G. 1973. Labops hesperius Uhler: Biology and impact in Oregon rangelands (Hemiptera: Miridae). M.S. thesis, Oregon State University, Corvallis. 91 pp. Todd, J. G. 1974. Biology of the wheatgrass bug in Oregon rangelands. Society for Range Manage- ment. 27th Annual Meeting, Abstract, p. 31. [The life history of Labops hesperius was studied on rangeland seeded to intermediate wheatgrass in central Oregon (see Todd and Kamm 1974).] Todd, J. G. and J. A. Kamm. 1974. Biology and impact of a grass bug Labops hesperius Uhler in Oregon rangeland. Journal of Range Management 27: 453-458. [The biology of a univoltine grass bug Labops hesperius is outlined for a site in Baker County, Oregon. Winter was passed as an egg in grass straw, eggs hatched in late March, and the nymphs reached the adult stage by late April. About two weeks later, new eggs were deposited in dry grass from the previous year. The nutrient value 109 of the new season’s growth of intermediate wheat- grass was reduced about 18% half way through the season; the losses were reduced to 2% by the time the grass had matured. The effect of insect feeding on range productivity varied with rainfall, grazing period, and drought.]} Uhler, P. R. 1871. A list of Hemiptera collected in eastern Colorado and northeastern New Mexico, by C. Thomas, during the expedition of 1869, pp. 471-472. In Hayden, F. V., Preliminary Report of the United States Geological Survey of Wyo- ming and Portions of Contiguous Territories. [La- bops hesperius and Irbisia pacifica are reported without specific localities. ] Uhler, P. R. 1872. Notices of the Hemiptera of the western territories of the United States, chiefly from the Surveys of Dr. F. V. Hayden, pp. 392-493. In Hayden, F. V., Preliminary Report of the United States Geological Survey of Montana and Portions of the Adjacent Territories 5: 392-493, United States Government Printing Office, Washington, D.C. [Irbisia pacifica (as Rhopalotomus pacificus) is described from Montana, and J. brachycera (as Rhopalotomus brachycerus) and Labops hesperius are described from Colorado and Canada (Lake Winnipeg and Great Bear Lake).] Uhler, P. R. 1876. List of Hemiptera of the region west of the Mississippi River, including those col- lected during the Hayden explorations of 1873. Bulletin of the United States Geological and Geo- graphical Survey of the Territories 1: 267-361, plates 19-21. [Irbisia pacifica (as Rhopalotomus pacificus) is reported from Montana, Idaho, and California and J. brachycera (as Rhopalotomus brachycerus) is reported from California and Col- orado. Labops hesperius is reported from Colo- rado, Montana, and Canada.] Uhler, P. R. 1877. 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., M.D. Bulletin of the United States Geological and Geographic Survey of the Terri- tories 3: 355-475, 765-801, plates 27-28. [Labops hesperius is reported from near Gray’s Peak, Col- orado.] Uhler, P. R. 1886. Check-list of the Hemiptera Het- eroptera of North America. Brooklyn Entomolog- ical Society. 32 pp. [/rbisia pacifica (as Capsus pacificus), I. brachycera (as Capsus brachycorus), and Labops hesperius are listed all from the “west- ern states.”’] Uhler, P. R. 1894. Observations upon the heterop- terous Hemiptera of Lower California, with de- scriptions of new species. Proceedings of the Cal- ifornia Academy of Sciences. Second Series 4: 223- 295. [The genus Thyrillus is described to include 110 Rhopalotomus pacificus and R. brachycerus. (Note: Thyrillus Uhler is a synonym of /rbisia Reuter.) Irbisia pacifica is reported from Lower (Baja) Cal- ifornia, southern California, San Francisco, south- ern Nevada, and Yakima, Washington. /rbisia brachycera, reported from Lower California, is considered common in California (state).] United States Department of Agriculture, Animal and Plant Health Inspection Service, Plant Protection and Quarantine Program. 1951-1975. Cooper- ative Economic Insect Report. Vol. 1-25. [This weekly series, listing site-specific occurrences of various economic and non-economic insects from throughout the United States, includes occasional references to Labops and Irbisia.]} United States Department of Agriculture, Animal and Plant Health Inspection Service. Plant Protection and Quarantine Programs. 1976-1980. Coop- erative Plant Pest Report. Vol. 1-5. [This series superseded the Cooperative Economic Insect Re- port cited above.] Usinger, R. L. 1934. Blood sucking among phytoph- agous Hemiptera. Canadian Entomologist 66: 97— 100. [/rbisia solani bit a person in California and produced a small, red spot that persisted for a short time.] Van Duzee, E. P. 1889. Hemiptera from Muskoka Lake District. Canadian Entomologist 21: 1-11. [Labops hesperius is reported from an oat field along the Muskoka River, Canada.] Van Duzee, E. P. 1905. List of Hemiptera taken in the Adirondack Mountains, pp. 546-556. Jn Felt, E. P., 20th Report of the [New York] State En- tomologist for 1904. New York State Museum Bulletin 97. [Labops hesperius is reported from Axton, New York, in the Adirondack Mountains. ] Van Duzee, E. P. 1912a. A few days’ work and play in Canada. Ottawa Naturalist 26: 68-70. [Labops hesperius is reported from Hull, Ontario, Canada.] Van Duzee, E. P. 1912b. Synonymy of the Provanch- er collection of Hemiptera. Canadian Entomolo- gist 44: 317-329. [The identity of Labops hesperius in the Provancher collection is confirmed. (Note: the Provancher collection is located in Laval Uni- versity, Quebec City, Canada.)] Van Duzee, E. P. 1914. A preliminary list of the Hemiptera of San Diego County, California. Transactions of the San Diego Society of Natural History 2(1): 1-57. lrbisia pacifica and I. brachy- cera are reported from San Diego County; the lat- ter species is considered the most common mirid in cultivated regions of southern California.] Van Duzee, E.P. 1916a. Synoptical keys to the genera of the North American Miridae. University of Cal- ifornia Publications, Technical Bulletins, Ento- mology 1: 199-216. [The keys include /rbisia and Labops.]| PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Van Duzee, E. P. 1916b. Notes on some Hemiptera taken near Lake Tahoe, California. University of California Publications, Technical Bulletins, En- tomology 1: 229-249. [Labops hesperius is re- ported from Lake Tahoe.] Van Duzee, E. P. 1916c. Check list of the Hemiptera (excepting the Aphididae, Aleurodidae and Coc- cidae) of America, north of Mexico. New York Entomological Society, New York. 111 pp. [Dis- tribution information for /rbisia sericans, I. bra- chycera, I. solani, I. pacifica (as Thyrillus pacifi- cus), Labops hesperius, and L. burmeisteri is in- cluded. ] Van Duzee, E. P. 1917a. Catalogue of the Hemiptera of America north of Mexico. University of Cali- fornia Publications, Technical Bulletins, Ento- mology 2: 1-902. [/rbisia sericans, I. brachycera, I. pacifica (as Thyrillus pacificus), Labops bur- meisteri, and L. hesperius are included.] Van Duzee, E. P. 1917b. Report upon a collection of Hemiptera made by Walter M. Giffard in 1916 and 1917, chiefly in California. Proceedings of the California Academy of Sciences. Fourth Series 7: 249-318. [Irbisia californica (as I. sericans, see Schwartz 1984, p. 2-34) is reported from several localities in California. [rbisia mollipes, described as a variety of sericans from specimans near San Francisco, is now considered a valid species. /r- bisia pacifica (as Thyrillus pacificus) is reported from California.] Van Duzee, E. P. 1921a. A study of North American grass-bugs of the genus /rbisia. Proceedings of the California Academy of Sciences. Fourth Series 11: 145-152. [Six new species of /rbisia are described from western North America: J. arcuata (a syn- onym of brachycera), I. californica, I. castanipes, I. parta (a synonym of brachycera), I setosa, and I. sita. A key is included to the ten known species in the genus (J. brachycera, I. mollipes, I. sericans, and /. solani, plus the six new species). Distribu- tion information is included with the original de- scriptions of the six new species and for /rbisia mollipes.] Van Duzee, E. P. 1921b. Insects of the Pribilof Is- lands, Alaska. Orthoptera, Neuroptera, Hemip- tera and Lepidoptera. Proceedings of the Califor- nia Academy of Sciences. Fourth Series 11: 193- 195. [Irbisia sericans is reported from St. Paul Island and St. George Island.] Van Duzee, E. P. 1926. Labops burmeisteri Stal. Pan- Pacific Entomologist 2: 163. [Labops burmeisteri is recorded from the Adirondack Mountains of New York and comments are made on L. hirtus.] Vosler, E. J. 1913. A new fruit and truck crop pest CUrbisia brachycerus Uhler). California (State) Commission of Horticulture. Monthly Bulletin 2: 551-553. [Irbisia solani (as Irbisia brachycera, see VOLUME 97, NUMBER 1 Schwartz 1984, p. 281) injured garden crops and fruit in California. Damage to lettuce, radishes, onions, peaches and rhubarb is recorded. Weedy plants in uncultivated areas near the crops dried up, resulting in the bugs moving into crops.] Watts, J. G., E. W. Huddleston, and J.C. Owens. 1982. Rangeland entomology. Annual Review of Ento- mology 27: 283-311. [This broad overview in- cludes a concise treatment of black grass bugs and other mirids.] Wheeler, A. G., Jr.and T.J. Henry. 1992. ASynthesis of the Holarctic Miridae (Heteroptera): Distribu- tion, Biology, and Origin, with Emphasis on North America. Entomological Society of America, Tho- mas Say Foundation, Volume 15, Lanham, Mary- land. 282 pp. [/rbisia sericans and Labops bur- meisteri are discussed with information on distribution, host plants, habits, and zoogeogra- 111 phy. Distribution maps in North America are in- cluded.]} Windig, W., H. L. C. Meuzelaar, B. A. Haws, W. F. Campbell, and K. H. Asay. 1983. Biochemical differences observed in pyrolysis mass spectra of range grasses with different resistance to Labops hesperius Uhler attack. Journal of Analytical and Applied Pyrolysis 5: 183-198. [Range grass species and hybrids differing in susceptibilities towards feeding damage by Labops hesperius were exam- ined by means of pyrolysis mass spectrometry. Clear correlations between pyrolysis patterns and susceptibility to insect damage were found. A pos- sible attractive role of chloroplast components and a possible repellant role of phenolic moieties that might influence the feeding behavior of the grass bug are suggested.] PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 112-116 NOTES ABOUT THE OLD WORLD GENUS HEXAMEROCERUS REUTER (HETEROPTERA: REDUVIIDAE: ECTRICHODIINAE) J. MALDONADO CAPRILES Department of Crop Protection, University of Puerto Rico, Mayagiiez, Puerto Rico 00681; mailing address: Urb. Aponte 6 I 1, Cayey, Puerto Rico 00736. Abstract. —Hexamerocerus and a syntype of H. punctatus (Stal) are redescribed, H. quadrimaculatus, sp. nov. is described from Mashonaland, South Africa, and a key to the four species of the genus is given. Key Words: latus, n. sp., key Reuter (1881) described the South Afri- can genus Hexamerocerus for the type spe- cies H. nobilis from Zanzibar. Thanks to Mrs. J. Margerison-Knight, from the British Natural History Museum, I was able to ex- amine a female syntype of H. punctatus, describe a new species, and prepare the notes discussed below. Measurements are given in mm. Hexamerocerus Reuter Reuter 1881 (1883). 12: 306. Type Hex- amerocerus nobilis Reuter 1881, 12: 307. Body parallel-sided, narrow, not flat- tened; head finely and sparsely granulose, both lobes thinly corrugate, broader behind than in front of eyes, without ventrolateral pouches behind eyes; antennae six seg- mented, the first about half as long as head or second segment, second the longest, about twice as long as each of the last segments, all segments short, with fine decumbent se- tae, thickened; rostrum moderately thick, first segment almost as long as last two com- bined, slightly dilated apically; eyes rela- tively small, not reaching upper or lower margins of head. Thorax wider than long, without collar, anterior margin below dorsal Reduviidae, Hexamerocerus, H. punctatus redescription, H. quadrimacu- surface of anterior lobe; anterior lobe short- er than posterior lobe, coarsely punctate above and laterally, lateral margins cari- nate; posterior lobe transversely corrugate (only the corrugations in the mediolongi- tudinal impression illustrated in Fig. 4); me- dian longitudinal sulcus of pronotum ex- tending across transverse constriction just before base of posterior lobe; mesopleurae horizontally and metapleura vertically cor- rugate; mesosternum radially and metaster- num transversely corrugate, the latter with a broad median depression; scutellum qua- drangular, wider than long, two widely sep- arated, short, converging prongs, on apex. Anterior femur incrassate, second slightly incrassate, third slightly thickening toward apex; pro- and mesotibia slightly expanded apically, both with spongy fossa on slanted apical surface; each femur ventrally with a postmedian toothlike spine following one or two small spines. Tarsi 3, 3, 3; third segment as long as first two combined; claws slightly expanded basally. Hemelytral surface smooth. Abdomen coarsely punctate dor- sally and ventrally, slightly compressed lat- erally, parallel sided, connexivum exposed, apical connexival angles slightly produced as elevated, rounded angulations, sternal LULL. HK UM MMMM Ma bh. WLLL — 208 SHO NOM 2 Figs. 1-14. 1-9. Hexamerocerus punctatus (Stal), female syntype. 1, head, lateral view. 2, head, dorsal view. 3, hemelytron and connexivum, dorsal and lateral view, respectively; colors of hemelytron inverted. 4, pronotum, dorsal view. 5, anterior leg, external view. 6, evaporation area of mesopleural scent gland, lateral view, arrow: opening of gland, P: pronotum, M: mesopleuron. 7, external genitalia, lateral view. 8, scutellum, lateral view. 9, external genitalia, caudal view. 10-14. Hexamerocerus quadrimaculatus, n. sp., female holotype. 10, hind femur, external view. 11, evaporation area of mesopleural scent gland, lateral view. 12, external genitalia, lateral view. 13, external genitalia, caudal view. 14, hemelytron and connexivum, dorsal and lateral view respectively; colors of hemelytron inverted. All scale lines equivalent to 1.0 mm. 114 sutures crenulate; spiracles circular, each on a globose area. Species with blackish-blue or black me- tallic bodies, apical half of some connexival segments yellow or pale orange, some spe- cies with small yellowish areas in different parts of body; hemelytra dull, usually with a grayish semicircular area and with or with- out a longitudinal fascia on corium; legs un1- formly dark, polished or with base and apex of segments fulvous; anterior lobe of pro- notum darker than posterior lobe. Basal and apical connexival segments with caudal halves yellow above and below, the middle segments with connexivum colored as cor- responding sterna (Fig. 14). Reuter considered Hexamerocerus close to Labidocoris Mayr, Mendis Stal, and Clep- tria Stal. It can be separated from all African ectrichodiine genera by the narrow, metal- lic-blue body, a six-segmented antenna, the relatively thick rostrum with the first seg- ment almost as long as last two combined, the coarsely punctured body, and the trans- versely corrugate posterior lobe of prono- tum. Not mentioned before in the literature is a narrow, vertical evaporation area, prob- ably associated with a scent gland, anteri- orly on the mesopleura (Fig. 6). It differs from other pleural sclerites by its dull sur- face; in Fig. 6 the double lines in the sur- rounding sclerites represent carinae or cor- rugations. The opening of the gland is above, just below the beginning of the caudal ex- tension of the posterior lobe of the prono- tum (Fig. 6, arrow). A similar area occurs in at least some other ectrichodiine genera. Hexamerocerus punctatus (Stal) Figs. 1-9 Pirena punctata Stal, 1863, 3: 47, Caffraria. Ectrichodia punctata: Walker, 1873, 8: 46. Hexamerocerus punctatus: Bergroth, 1894, 382547. Female.— Coloration metallic black: head, anterior lobe of pronotum laterally and dor- sally, meso- and metapleurae, scutellum, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON basal half of connexival segments II, III, IV, VI, and VII, connexival segment V, basal half of ventral connexival segments II, III, VI, and VII, thoracic sterna, and genital seg- ments. Antenna brown; legs dark brown, tarsi pale brown. Caudal half of connexival segments mentioned above yellow (Fig. 3). Abdominal sterna metallic dark blue. Pos- terior lobe of pronotum metallic dark brown, a shade paler than legs. Hemelytra: clavus and corium, very dark red, membrane black; a gray, suboval transverse spot across on apex of corium; hemelytra reaching middle of last tergum. Head (Figs. 1, 2).—Anterior lobe of pro- notum dorsally and laterally grossly punc- tate (Fig. 4), lateral margin carinate; median longitudinal sulcus shallow and narrow along anterior half, on caudal half gradually broader and deeper, crossing over onto pos- terior lobe; median longitudinal sulcus rel- atively broad anteriorly, tapering to and ending before basal margin of lobe. Evap- oration area of mesopleuron as in Fig. 6. Scutellum width 0.6, length 0.5, with discal quadrangular depression, slightly raised to- wards apex (Fig. 8). Legs: pro- and meso- femora incrassate (Figs. 5, 10), metafemur slightly thickening toward apex, slightly constricted apically; large spine on femora about 4% to '4 diameter of segment. Head—length 1.40, width across eyes 1.02, width behind eyes 0.87, width in front of eyes 0.62, distance from anterior margin of eye to apex of antennophore 0.31, from anterior margin of eye to apex of head 0.62, from posterior margin of eye to base of pos- terior lobe 0.43, width of eye 0.19, interoc- ular space 0.64, from interocular suture to apex of head 1.00, from interocular suture to base of posterior lobe 0.4, collum 0.18. Antennal segments: I, 0.62; II, 1.19; III, 1.0; IV, 0.5; V, 1.0, VI missing. Rostral seg- ments: 1.0; 0.62; 0.4. Thorax length 1.93, width 2.12; anterior lobe—length 0.81, greatest width 1.50; posterior lobe—length 1.12, greatest width 2.12. Scutellum width 0.69, length 0.50. Legs: length and depth of VOLUME 97, NUMBER 1 femora: 1.56°x+0:5168: * 0:5; 2:3.1-x..0.5; tibiae length: 1236, 1.50, 2:01 (Figs. 5; 10). Length of abdomen 5.51, margins of con- nexival segments straight, greatest abdom- inal width at segment V, 2.62. External gen- italia as in Figs. 7 and 9. Total length of body 10.30. Syntype female, from Pt. Natal, SOUTH AFRICA; pin with: red-margined card- board circle labeled ‘“‘type’’; blue-margined circle labeled “‘syntype’’; cardboard label typed “Ectrichodia punctata”’; handwritten label ‘‘punctata Stal’; in Natural History Museum, London. Hexamerocerus quadrimaculatus, n. sp. Maldonado Figs. 10-14 Female.—Coloration dark metallic blue: head, anterior lobe of pronotum dorsally and laterally, meso- and metapleura, meso- and metasternum, abdominal sterna, and scutellum; abdominal tergites black; pos- terior lobe of pronotum brown, with a me- tallic bluish tinge. Antenna: segments I and II dark brown, III and IV black, others miss- ing; rostrum dark brown. Legs: coxae brown, polished; trochanters pale brown; profemur polished dark brown, apex shortly ringed with stramineous; mesofemur: middle % polished dark brown, apex and base ful- vous; metafemur with same colors as me- sofemur, as in Fig. 10; protibia: external surface with basal half and apex brown, preapical stramineous ring, internal surface with basal 34 whitish, apical 4 brown; me- sotibia with both surfaces as protibia exter- nally; metatibia brown, dark stramineous preapically. Tarsi pale brown. Hemelytra black, with grayish corial longitudinal stripe parallel to clavocorial suture, a gray, sub- hemispherical area at end of corium and over base of membrane (Fig. 14). Colora- tion of connexival segments: dorsal caudal halves of II narrowly, III, [V, VI, and VII entirely yellow; ventral caudal halves of IT, III, 1V narrowly, VI and VII entirely yellow, the last two fused above and below: spirac- 1S ular areas VI and VII yellow, apparently fused. Head—length 1.62, width across eyes 1.16, width behind eyes 1.00, width in front of eyes 0.75, from anterior margin of eye to apex of antennophore 0.40, from anterior margin of eye to apex of head 0.89, from posterior margin of eye to base of posterior lobe of head 0.37, width of eye 0.19, inter- ocular space 0.68, from interocular suture to apex of head 1.02, from same to base of posterior lobe of head 0.6, collum 0.19. An- tennal segments: I, 0.75; II, 1.43; III, 0.62; IV, 0.4; V and VI missing. Rostral seg- ments: 1.0; 0.75; 0.37. Thorax: length 2.29, greatest width 2.62, posterior margin slight- ly angularly produced; anterior lobe length 0.93, width 1.81; posterior lobe length 1.36, width 2.62; evaporation area of mesopleura as in Fig. 11. Scutellum width 0.87, length 0.68. Legs—femora length and depth: 1.81 x 0.5: 1.94 x 0.5; 3.06 x 0.5. Tibiae lengths: 1.62, 1.87, 2.75. Hemelytra smooth; slightly surpassing apex of abdomen. Abdomen: margins of connexival segments slightly concave, length 6.77, greatest width at V segment 3.00 (Fig. 14). External genitalia as in Figs. 12 and 13. Total body length 11.06. Holotype female, Mashonaland, SOUTH AFRICA; 1897, collector unknown; in Nat- ural History Museum, London. KEY TO THE SPECIES IN HEXAMEROCERUS 1. Corium of hemelytra with longitudinal fascia close and parallel to the clavacorial commis- Sures(Eigeb4 ip eee rns ment ee en nee. 2 Corium without such longitudinal fascia (Fig. 3) ee ones es ee: cre ee heme ee tO 3 2. Hemelytra and antennae black; dorsal connex- ival segments II, III, 1'V, VI, VII with caudal halves yellow to orange; ventral segments II, III, IV, VI, and VII colored similarly; small yellowish area near each mucron of scutellum, acetabulum, coxae, and first abdominal ster- num; length 10.3; Zanzibar .... H. nobilis Reuter Antenna brown basally, gradually turning to black toward apex; hemelytra dull dark brown; yellow apical half of connexival segments II, III, IV, and yellow of VI and VII fused; body without small yellow areas; length 1 1.1; Mash- 116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bi 3 hs WA eee H. quadrimaculatus, n. sp. 3. Legs black, base and apex of pro- and meso- femora luteus; hemelytra black; Mozambique Re PR Ty oy ee one H. junodi Distant Legs uniformly black; clavus and corium dark red, membrane black; 8.84 mm; South Africa Ee A RA A tert a ake H. punctatus (Stal) LITERATURE CITED Bergroth, E. 1894. Rhynchota Aethiopica II. Annales Societe Entomologique Belgique 38: 539-547. Distant, W.L. 1908. Rhynchotal miscellanea (second series). Part I. Rhynchota from the Transvaal, Mashonaland, and British Nyasaland. Annals of the South African Museum 3: 43-62. Reuter, O. M. 1881 (1883). Ad cognitionem Redu- viidarum mundi antiqui. Acta Societatis Scientia- rum Fennicae 12: 269-339. Stal, C. 1863. Formae speciesque novae Reduvi- idum. Annales Societe Entomologique France 4: 47-68. Walker, F. 1873. Catalogue of the specimens of Het- eroptera-Hemiptera in the collection of the British Museum. Part VII. British Museum, London 7: 1-213. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 117-122 PODALIA BOLIVARI (LEPIDOPTERA: MEGALOPYGIDAE): A HIGHLY SEXUALLY DIMORPHIC NEOTROPICAL PEST Scott E. MILLER, VITOR O. BECKER, AND RAUL VELEZ-ANGEL (SEM) Bishop Museum, Box 19000-A, Honolulu, Hawaii 96817, U.S.A.; (VOB) Centro de Pesquisa Agropecuaria dos Cerrados, Caixa postal 08223, 73301-Planaltina, Brasil; (RVA) Facultad de Ciencias, Universidad Nacional de Colombia, Apartado Aereo 3840, Medellin, Colombia. Abstract. —Podalia bolivari (Heylaerts) is unusual among the Megalopygidae in its ex- treme sexual dimorphism, as well as the male habitus. Association of the sexes is confirmed and the species is redescribed to allow its identification. Observations on the natural history of P. bolivari are given. Problems of generic concepts in Megalopyginae are dis- cussed. Key Words: Podalia bolivari is unusual among the Megalopygidae in its extreme sexual di- morphism, as well as the male habitus. This has caused confusion in its classification and identification. Because this species can be a pest of cultivated palms and ferns in Co- lombia, we redescribe it here. The taxonom- ic portion of this paper was prepared by SEM and VOB, the natural history obser- vations are by RVA. The male was described by Heylaerts (1884) as Pentophora bolivari, but the fe- male has been named three times (Megal- opyge pellucens Dognin, 1912; Unduzia gis- tinda Dyar, 1914; and U. phaule Dyar, 1914). Dognin (1916) recognized the syn- onymy of pellucens and phaule, and Joicey and Talbot (1922, based on unpublished 1n- formation from Dyar) synonymized pellu- cens and gistinda. Hopp (1926, 1935) rec- ognized the association of the male and female, but other workers (unpublished notes in USNM) doubted the association because of the extreme dimorphism. We can now confirm the association, based on males and females reared together. Megalopygidae, Zygaenoidea, Megalopyge, palms, ferns The sexual dimorphism of Podalia boli- vari is the most extreme that we are aware of in Megalopygidae, both in size and wing shape (similar to, but more extreme than, dimorphism in Phobetron in the Limacod- idae). Forewing lengths in male bolivari are 7-9 mm; female lengths are 17-22 mm. The usual dimorphism in megalopygids is that the largest males are slightly smaller than the smallest conspecific females. In addition to size, bolivari is strongly dimorphic in habitus. The females are fairly typical Po- dalia in appearance (although the wings are semitranslucent), but the males are very un- usual for Megalopygidae. The males have narrow, hyaline wings, yielding the appear- ance of Zygaenidae, Psychidae, or Arctiidae such as small species of Paracles (Becker and Miller 1991); Podalia bolivari can be distinguished from most of these by the stalking of forewing veins R4 and R5. This species can be a pest of palms and ferns in the vicinity of Medellin, Colombia. Gallego (1946) recorded Podalia bolivari as a pest of Washingtonia palms, and it is quite common on the introduced palm Chrysal- 118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1, 2. wing (USNM) (forewing length 19 mm); same specimen illustrated against both black and white backgrounds. Both sexes figured at same scale to show dimorphism. idocarpus lutescens H. Wendl. (= areca palm or bamboo palm). It is also found on an ornamental fern, Preridium sp., and Cype- rus diffusus Vahl (Cyperaceae) on the Univ- ersidad Nacional de Medellin campus. Lar- vae develop much more slowly on Cyperus than Preridium. Genty et al. (1978: 382- Podalia bolivari wings: | (upper), male wing (USNM) (forewing length 9 mm); 2 (lower), female 383) noted P. bolivari as a secondary pest of oil palms. All the species of Megalopyginae are cur- rently placed in either Megalopyge Hiibner or Podalia Walker (Hopp 1935), except for the enigmatic genus Psychagrapha Walker (transferred to Megalopyginae by Epstein VOLUME 97, NUMBER 1 niger: and Becker 1994: 313, Becker 1994). The generic classification of the subfamily needs revision (Miller 1994). Such a revisionary study may result in the lumping of all spe- cies into Megalopyge or the splitting of the subfamily into additional genera. The ge- neric name Unduzia was proposed by Dyar (1914: 252), with U. gistinda as type species. We follow Hopp (1935) and Forbes (1942) in placing bolivari in Podalia until the ge- neric classification of the subfamily can be revised. REDESCRIPTION Podalia bolivari (Heylaerts) Figs. 1-4 Pentophora bolivari Heylaerts, 1884: xli. Hypogymna bolivari: Kirby, 1892: 490. Unduzia bolivari: Hopp, 1926: 193.—Gal- lego, 1946: 455.—Morales, 1982: 168. 119 Podalia bolivari, male genitalia (USNM 28024). Podalia bolivari: Hopp, 1935: 1098, pl. 163g.—Forbes, 1942: 404.—Genty et al., 1978: 382-383. Megalopyge pellucens Dognin, 1912: 171. Unduzia pellucens: Dognin, 1916: 22. Unduzia gistinda Dyar, 1914: 252. Unduzia phaule Dyar, 1914: 252. Diagnosis.—Male dark grey, with nar- row, hyaline wings, yielding the appearance of Zygaenidae or Psychidae (Fig. 1). Female fairly typical of Podalia in appearance, but with semitranslucent wings; forewing with postmedial white spots (Fig. 2). Adult male (Fig. 1).— Forewing length 7- 9 mm. Head: Pale fuscous, densely hairy. Antennae dark grey, broadly bipectinate. Thorax: Dark grey dorsally, slightly paler ventrally. Densely covered with long hairs. Forewings sparsely covered with long nar- row scales, dense only along wing margins; 120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Podalia bolivari, female genitalia (USNM 28162, paralectotype of pellucens). scaling especially sparse in middle of wing, creating translucent appearance. Hindwings as in forewings, but densely scaled from CuP to posterior margin. Wings ventrally as dor- sally, slightly paler. Legs dark grey. Abdo- men: Dark grey, densely covered with long hairs. Genitalia as in Fig. 3. Adult female (Fig. 2).—Forewing length 17-22 mm. Entire moth pale fuscous, cov- ered with long hairs. Head: Densely hairy. Antennae narrowly bipectinate. Thorax: Densely hairy. Forewings pale fuscous, sparsely covered with long, curled hairs, re- sulting in appearance of undulating bands, translucent; postmedian line consisting of patches of elongate white hairs, one patch in each cell, midway between each vein after M2. Hindwings pale fuscous, more sparsely clothed than forewing. Abdomen: Pale fus- cous, densely hairy. Genitalia as in Fig. 4. Types.—Holotype male, ZMHB (boli- vari); Lectotype female, here designated [lectotype bears Dognin’s handwritten label “‘type,”’ three paralectotypes bear “‘co- type’’], USNM 29855 (pellucens); Lectotype female, here designated, USNM 16097 (gis- tinda), Holotype female, USNM_ 16098 (phaule). [all examined] Type localities. — Venezuela, Mérida (bo- livari); Venezuela, Mérida (pellucens); Pan- ama, Rio Trinidad (gistinda); Venezuela, Mérida (phaule). Hosts. — Washingtonia filifera (L. Linden) H. Wendl. (Gallego 1946: 455); Elaeis gui- neénsis Jacq. (Genty et al. 1978); Cyperus diffusus Vahl; Chrysalidocarpus lutescens H. Wendl. (USNM); Pteridium sp. (USNM). Gallego (1946: 455) also recorded Wash- ingtonia comunis and Washingtonia erecta, but we have been unable to identify these taxa; they are not valid species of Wash- ingtonia. Immature stages. — Larvae densely hairy, reddish, up to 30 mm long (see below). Flight period.— March, May, August, and December (in Panama). Distribution.— Panama, Colombia, and Venezuela. VOLUME 97, NUMBER 1 Material examined [by SEM and VOB].— 10 males and 18 females. COLOMBIA: An- tioquia: Medellin, [no date], F. L. Gallego M. (USNM), VIII-1985, ‘“‘ex helecho”’ [reared from Pteridium sp.], J. A. Quiroz & F. Serna (USNM), X-1985, “‘en palma are- ca” [reared from Chrysalidocarpus lutes- cens], R. Velez (USNM); Boyacad: Muzo, 400-800 m, [no date], A. H. Fass] (USNM); PANAMA: Barro Colorado Island, 5-VIII- 1940, N.S. Scrimshaw (MCZ), 10-XII-1934, M. Bates (MCZ); Cabima, 16-30-V-1911, A. Busck (USNM); Cano Saddle, V-[no year], R. C. Shannon (USNM); Rio Trini- dad, 15-31-III-1912, Busck (USNM, lec- totype of gistinda), same but III-1912 (USNM, paralectotype of gistinda); VEN- EZUELA: Mérida: Mérida, [no date] (USNM, lectotype and 3 paralectotypes of pellucens), 1890 (USNM), [no date], S. Bri- ceno (USNM, holotype of phaule). Discussion. — Hopp (1926, 1935; repeat- ed by Forbes 1942) mentions México, but we have not seen any specimens to confirm this distribution record. We have seen a sin- gle female from Peru ([Valle de] Canchcha- mayo, I-VII-1901, W. Hoffmanns, BMNH), evidently the specimen noted by Hopp (1935), which might be P. bolivari but it differs slightly in coloration and venation. We have examined the “type” male of Podalia bolivari in ZMHB, as did Hopp (1926). The original description implies that Heylaerts had only one specimen, so we consider this specimen the holotype. It is in good condition, except for lacking the ab- domen (as noted in the original description) and right antenna. It bears a handwritten locality label ‘““Merida/Hahnel.” Paul Hah- nel collected in Venezuela from 1875-1879 (Horn and Kahle 1935: 102). Podalia bolivariis most similar to P. dyari (Joicey and Talbot 1922: 302), known only from the female holotype (BMNH) from Ec- uador (specific locality not known). The ho- lotype of dyari is much larger than bolivari, with a forewing length of 27 mm. The wing shape 1s more pointed in dyari, with the 121 Table 1. Life cycle of Podalia bolivari on leaves of areca palm (Chrysalidocarpus lutescens) at 24°C and 67% R.H. Mean No. life span individuals Life stage (days) observed Egg 9 200+ Larvae Males 49-50 34 Females 65-70 33 Pupae Males 25-28 24 Females 21-26 23 Adults (males & females) 1 47 outer margin more oblique than in bolivari. The wings of dyari are also more transparent and the forewings lack the postmedial white spots, although this is hard to evaluate with only one specimen. We have not dissected the genitalia of dyari, as there is no mor- phological knowledge of megalopygid fe- male genitalia for comparative evaluation (Fig. 4 in this paper is the first published illustration of a megalopygid female geni- talia). NATURAL HISTORY Adult and eggs: Adults eclose at night under laboratory conditions. After emerg- ing, both sexes show slow movements dur- ing the day but fly actively after dark. About 3 days after mating, the female lays a mass of eggs (close to 200 in each group), fre- quently deposited on its old cocoon. The eggs are very small, spherical and surround- ed by masses of web. In the field, the egg masses are covered completely with pili- form scales from the female abdomen. Adults live about 7 days when fed on ab- sorbent cotton soaked in a solution of honey (30%) and water (see Table 1). Larvae and pupae: The larvae, soon after hatching from the eggs, start feeding on fo- liage of the host plants reported. The larvae are covered with a dense coat of fine, long, reddish hairs (figured by Genty et al. 1978: 122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 382), show slow movements and prefer the underside of the leaves they feed on. The larvae have urticating setae beneath the long hairs that cause skin irritation. Last instar larvae reach about 3.0 cm long. Mature lar- vae seek hidden pupation sites on walls or similar surfaces close to their feeding site. The pupae can be easily sexed by the clearly larger size of the females. Natural enemies: Egg parasites, Teleno- mus sp. (Hymenoptera: Scelionidae), were observed emerging from an egg mass col- lected on C. diffusus. A fly (possibly Ta- chinidae) was seen after emerging from a dead larva. Signs and symptoms similar to those of affliction with a viral disease have also been noticed in larvae. ACKNOWLEDGMENTS Research facilities for SEM and VOB were provided by the Smithsonian Institution (USNM), and the photographs were taken by Victor Kranz of the Smithsonian Insti- tution. H. J. Hannemann loaned the type of Podalia bolivari. Additional specimens were borrowed from the Museum of Com- parative Zoology, Harvard University (MCZ) and the Natural History Museum, London (BMNH). Isabella Forster translat- ed Hopp’s paper for us. D. R. Davis, M. E. Epstein, and N. L. Evenhuis reviewed the manuscript. L. Masner, Agriculture Cana- da, identified the parasitic wasp. LITERATURE CITED Becker, V. O. 1994. Megalopygidae. Jn Heppner, J. B., ed., Atlas of Neotropical Lepidoptera: Check- list. Association for Tropical Lepidoptera, Gaines- ville. (In press.) Becker, V. O. and S. E. Miller. 1991. Three unusual species of Paracles from South America (Lepi- doptera: Arctiidae). Journal of Research on the Lepidoptera 28: 283-288 (1989). Dognin, P. 1912. Hétérocéres nouveaux de l’Amé- rique du Sud. Mémoires de la Société Entomo- logique de Belgique 19: 121-177. 1916. Hétérocéres nouveaux de l’Amérique du Sud. Fascicule X. Imprimerie Oberthiir, Rennes. 25 pp. Dyar, H.G. 1914. Report on the Lepidoptera of the Smithsonian Biological Survey of the Panama Ca- nal Zone. Proceedings of the United States Na- tional Museum 47: 139-350. Epstein, M. E. and V. O. Becker. 1994. Combinations and synonymies in New World Limacodidae, Me- galopygidae, Lasiocampidae and Arctiidae (Lep- idoptera). Revista Brasileira de Zoologia 10: 289- 319 (1993). Forbes, W. T. M. 1942. The Lepidoptera of Barro Colorado Island, Panama. No. 2. Bulletin of the Museum of Comparative Zoology 90: 265-406. Gallego M., F. L. 1946. Catalogo de insectos deter- minados correspondientes a la orden Lepidoptera existentes en la Seccion de entomologia de la Fa- cultad Nacional de Agronomia.— Medellin. Parte I. Diurnas, Rhopalocera o Achalinoptera. Univ- ersidad Nacional de Colombia, Facultad de Agronomia, Revista 6: 294-314, 415-473. Genty, P., R. Desmier, J. P. Marin, and C. A. Koryt- kowski. 1978. Les ravaguers du palmier a huile en Amérique Latine. Oléagineux 33: 325-419. Heylaerts, F. J. M. 1884. Description de deux bom- bycides exotiques nouvelles. Bulletin de la Société Entomologique de Belgique 28: xli—xliv. Hopp, W. 1926. Megalopygiden-Studien II. Deutsche entomologische Zeitschrift 1926: 193-198. 1935. Megalopygidae, pp. 1071-1101. Jn Seitz, A., ed., The Macrolepidoptera of the World, Volume 6. Alfred Kernan, Stuttgart. Horn, W. and I. Kahle. 1935-1937. Uber entomo- logische Sammlungen, Entomologen und Entomo- Museologie. Entomologische Beihefte 2-4: vi, 536 pp., 38 pl. Joicey, J. J.andG. Talbot. 1922. New forms of moths from New Guinea and South America. Bulletin of the Hill Museum 1: 300-302. Kirby, W. F. 1892. A synoptic catalogue of Lepi- doptera Heterocera. (Moths.) Vol. I. Sphinges and bombyces. Gurney and Jackson, London. xiii + 951 pp. Miller, S. E. 1994. Systematics of the Neotropical moth family Dalceridae (Lepidoptera). Bulletin of the Museum of Comparative Zoology 153: 301- 495. Morales Soto, G. 1982. Notas sobre taxonomia y clasification de los insectos. Parte segunda del cur- so entomologia general y systematica. Universi- dad Nacional de Colombia, Medellin. 257 pp. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 123-127 THE IDENTITY AND SYNONYMY OF NEPA FUSCA LINNAEUS, 1758 (HETEROPTERA: NEPIDAE) JOHN T. POLHEMUS University of Colorado Museum, 3115 S. York St., Englewood, Colorado 80110. Abstract. —The nomenclatural history of Nepa fusca Linnaeus, 1758 is reviewed. Usages by other authors of the name Nepa fusca are also reviewed, some of which refer to different species, and occasionally to Belostomatidae. The synonymy of Nepa fusca Linnaeus, 1758 and Laccotrephes brachialis Gerstaecker, 1873 is established (New Synonymy). Key Words: Nepa fusca has remained an enigma ever since it was described by Linnaeus (1758). It has had a confusing nomenclatural his- tory because of uncertainty as to both its identity and provenance, the latter various- ly attributed to America, Asia and Africa. Linnaeus (1758, 1767) gave “Habitat in Calidis regionibus”’ but in 1764 gave simply ‘“‘Habitat—.’’ Gmelin (1790) gave ‘‘Habitat in Indiae aquis ... ,” Turton (1802) states that it ‘““Inhabits East India,” and Fabricius variously gave “Habitat in America” (1775), ‘Habitat in Americae meridionalis aquis”’ (1781), and ‘Habitat in Indiae orientali aquis” (1787, 1794, 1803). Fabricius (1775) described Nepa fusca as “N. ecaudata, scutello rugoso, alis niveis” indicating that the specimen was belosto- matid. He cites the third description of Lin- naeus (1767), but gave “Habitat in Amer- ica.” He further gave “Duplo major N. cinerea: tota fusca, solis alis albis,”’ leading Esaki (1926) to suggest that the species might be Lethocerus (Benacus) griseus (Say). Fabricius (1787; same in 1794) described Nepa fusca differently, as ““N. cauda biseta, scutello rugoso, alis niveis” clearly indicat- ing that the species was a nepid. In both (Fabricius 1787, 1794) he listed Stoll 1780, pl. I, fig. I as a citation for fusca, but this Heteroptera, Nepidae, Belostomatidae, Laccotrephes, synonymy must have been a lapsus (cf. Esaki 1926) because this is not a figure of a nepid, but an American belostomatid (Zaitha stollii Amyot and Serville, 1843: 430); pl. I, fig. II may have been intended, which 1s a Lac- cotrephes. As further evidence that this was a lapsus on the part of Fabricius, Stoll, in his citations for pl. I, fig. I, gave ““FABRI- CIUS Syst. Entom. pag. 691. Nepa 2. Rus- tica” (= Fabricius 1775) which is clearly an American belostomatid (but not the Nepa rustica of Fabricius 1787, 1794, 1803, which is synonymous with Nepa plana Sulzer, 1776, a belostomatid from India). On page 11, Stoll cited pl. I fig. II for Nepa cinerea Linnaeus, but on pages 35-36 he also cited it as illustrating his “Scorpion-aquatique gris” from Tranquebar and Coromandel (India). Amyot and Serville (1843) gave bi- nominal names to most of Stoll’s aquatic Heteroptera, but attributed his pl. I, fig. I solely to Nepa cinerea, and did not mention his ““Scorpion-aquatique gris.” Ferrari (1888), on the other hand, attrib- uted Stoll’s ““Scorpion-aquatique noir de Tranquebar’’ to fusca, with the citation as p. 29, tab. VIII, fig. 5, 1792, a German lan- guage edition of Stoll’s original work (1780- 88) which was printed in parallel Dutch and French; this citation refers to p. 35, tab. VII, 124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fig. V in the original. Amyot and Serville (1843) give this species in error as “Nepa rubra Linn.,” later shown by Lundblad (1933) to pertain instead to grossus Fabri- cius (= L. kohlii Ferrari sensu Esaki, 1926). Esaki (1926) studied the types of fusca in Uppsala, reviewed the works of Fabricius, and concluded that at least two of the Fa- bricius descriptions refer to other species. For instance the description of Nepa fusca by Fabricius in 1775 apparently refers to a belostomatid, and his 1803 descripton is of a species smaller than Nepa cinerea, or only about half the length of Linnaeus’ types of fusca. Fabricius’ 1794 description matches the Linnaean species more closely; at least it is approximately the same size. Esaki thought the latter might be Ferrari’s (1888) Laccotrephes kohlii, but this species has since been synonymized with Laccotrephes gros- sus Fabricius, 1787, described next to L. fusca Fabricius, 1787 in the same work so this cannot be. Lundblad, however (1933: 23) has tentatively cited L. fusca Fabricius, 1787 as belonging to his Laccotrephes oc- cultus, new name for “Laccotrephes fuscus auct. in coll. partim, non Laccotrephes fus- cus Linné, 1758.” Esaki did not mention Fabricius’ ‘‘Species Insectorum”’ (1781) where he repeated the description from his “Systema Entomologiae” (1775) but gave the habitat as “Habitat in Americae meri- dionalis aquis,”’ nor his ““Mantissa Insec- torum” (1787) wherein he gave the same description and provenance used in his “Entomologia Systematica”’ (1794). Stal (1868: 135) studied the Linnaean types of fusca in Uppsala, and provided a redescription in Latin. Esaki (1926) also studied the Linnaean types of fusca in Upp- sala, stated that the two specimens were both females with lengths of 31 and 33 mm, with caudal filaments of lengths 22.5 + X and 21 + X mm respectively, with apices of the caudal filaments broken (with ““+ X” Esaki indicated that an unknown additional length of caudal filament once existed). No further description was given. Fortunately Lundblad (1933) carefully restudied the Linnaean types of Nepa fusca and found that Esaki’s (1926) examination had been cursory indeed. The types are both males, not females as Esaki had stated, and do not belong to any Asian species known to Lundblad. Lundblad dissected and fig- ured the paramere of one male, and pro- vided a dorsal habitus photograph of the unspread specimen, presumably also the one he dissected. This specimen is here desig- nated as lectotype. The key characters given by Lundblad are: interocular space about twice the width of an eye, low prosternal carina, shape of male paramere, size of body (See also Poisson 1949: 30-31). Lundblad gave the body length of the lectotype as 30 mm, the siphon as 22 mm. The photograph clearly shows the characteristic widened abdomen. Poisson (1949) noted that the parameres of Laccotrephes brachialis closely match those of Laccotrephes fuscus, but he failed to formally equate the two species because he thought fuscus was an Asian species, from “Inde,” citing Lundblad (1933) as the source for this provenance. Gerstaecker described Laccotrephes bra- chialis from East Africa (from a locality giv- en as “See Jipe’’). He described the proster- nal carina of L. brachialis as low and of even height all along its length except for the an- terior tubercle, which is quite removed from the anterior margin; he gave the length of the body as 40 mm, the siphon 35 mm. Poisson (1949: 30-31; 1965: 240) re- stricted L. brachialis brachialis to those specimens with a body length of 33-40 mm and the sides of the abdomen enlarged, and gave the distribution as West Africa, Ugan- da, Zaire, etc. He assigned those of 37-45 mm with the sides of the abdomen parallel to L. brachialis oculatus Montandon, known from Sierra Leone and Zaire. Poisson (1954) has also described a considerably smaller subspecies, L. brachialis kazibae from Zaire, with body length given as 23-30 mm. Be- cause they all are said to occur in Zaire, the VOLUME 97, NUMBER 1 status of these subspecies should be care- fully scrutinized, for if any two are found together they must either be considered syn- onyms or separate species. SYNONYMY The evidence presented here leaves little doubt concerning the synonymy of Nepa fusca Linnaeus, 1758 and Laccotrephes bra- chialis Gerstaecker, 1873 (new synonymy). This species has three salient characteristics that in combination set it apart from any other Laccotrephes species: 1) The unique shape of the distal part of the male para- mere, which is somewhat like a crochet hook. All three subspecies share this char- acter to a degree but have other differences (see above). 2) The shape of the abdomen, slightly expanded just behind the middle. Laccotrephes ampliatus (Montandon) is similar in this regard, but is definitely broader than L. brachialis. 3) The unique shape of the prosternal carina. No other spe- cies I have examined (including most of the known species) has the odd characteristics of a low carina with an isolated anterior tumescence removed from the anterior margin. The usage of the names fusca and bra- chialis are about equal in the literature, so it is difficult to make a case for suppression of the name fusca, particularly in view of the previous evidence for the synonymy given by both Lundblad (1933) and Poisson (1949). The only plausible explanation of why one of these authors did not formalize the synonymy must have been their appar- ent belief and mind-set that fusca was Ori- ental. Laccotrephes fuscus (Linnaeus, 1758) Nepa fusca Linnaeus, 1758: 440. Lectotype here designated. Nepa fusca: Linnaeus, 1764: 166. Nepa fusca: Linnaeus, 1767: 713. Nepa fusca: Miller, 1774: 472. Nepa fusca: Goeze, 1778: 173. 125 Nepa fusca: Olivier, 1811: 189. [in part] Laccotrephes fuscus: Stal, 1868: 135. [re- description of Linnaean type] Laccotrephes brachialis Gerstaecker, 1873: 422. New Synonymy Laccotrephes brachialis: Montandon, 1914: 1273 Nepa fusca: Esaki, 1926: 179. [in part] Laccotrephes fuscus: Lundblad, 1933: 22- 23. [redescription of Linnaean type] Laccotrephes brachialis: Poisson, 1949, 30, fig. 32A, B, C, fig. 33A, B. Laccotrephes fuscus: Poisson, 1949: 31. [comparison with L. brachialis] Laccotrephes brachialis: Poisson, 1954: 17, fig. SA. Laccotrephes brachialis: Poisson, 1965: 240, fig. 3C. Laccotrephes brachialis: Linnavuori, 1971: B57. References to L. fusca referring to other species: Nepa fusca: Fabricius, 1775: 692. [belos- tomatid sp.; Benacus griseus? Scorpion-aquatique gris Stoll, 1780; 35-36. pl. I, fig. H. [Laccotrephes griseus (Gue- rin)?] Nepa fusca: Fabricius, 1781: 333. [belos- tomatid sp.; = fusca Fabricius, 1775] Nepa fusca: Fabricius, 1787: 277. [Lacco- trephes occultus Lundblad?; not L. grossa, which was described in the same work.] Nepa fusca: Gmelin, 1790: 2121. [Lacco- trephes occultus Lundblad?; = L. fusca F., 1787.] Nepa fusca: Fabricius, 1794: 62. [Lacco- trephes occultus Lundblad?; = L. fusca F., 1787.] Nepa fusca: Turton, 1802: 607. [Lacco- trephes occultus Lundblad?; = L. fusca F., 1787.] Nepa fusca: Fabricius, 1803: 107. [Lacco- trephes griseus (Guerin)?] Nepa fusca: Olivier, 1811: 189. [in part] Nepa fusca: Ferrari, 1888: 184. [Lacco- trephes sp.?; synonymy; redescription; nomen inquerendum] 126 Nepa fusca: Esaki, 1926: 179. [Laccotrephes spp.; in part] ACKNOWLEDGMENTS I am indebted to the following for helpful reviews of the manuscript: I. M. Kerzhner, Zoological Institute, St. Petersburg; S. L. Keffer, James Madison University, Harri- sonburg, VA; R. L. Sites, University of Mis- souri, Columbia. F. C. Thompson, National Museum of Natural History, Washington, D.C. provided a rare book from his private library. LITERATURE CITED Amyot, C. J. B. and J. G. A. Serville. 1843. Histoire naturelle des insectes. Hémiptéres. Libraire En- cyclopedique de Roret, Paris, Ixxvi + 675 + 6 pp., 12 pls. Esaki, T. 1926. Remarks on the Linnéan species of Nepa and Laccotrephes (Heteroptera: Nepidae). Bulletin of the Brooklyn Entomological Society, N. S. 21: 177-181. Fabricius, J.C. 1775. Systema entomologiae, sistens insectorum classes, ordines, genera, species adiec- tis synonymis, locis, descriptionibus, observation- ibus. Libraria Kortii, Flensbergi et Lipsiae, xxx + 832 pp. 1781. Species insectorum exhibentes eorum differentias specificas, synonyma auctorum, loca natalia, metamorphosin adjectis observationibus, descriptionibus. Carol. Ernest. Bohnii, Hamburgi et Kiloni, Vol. 2, 494 pp. + Appendix pp. 495- 514 + Index pp. 515-517. 1787. Mantissa insectorum sistens species nuper detectas adiectis synonymis, observationi- bus, descriptionibus, emendationibus. Christ. Gottl. Proft, Hafniae, Vol. 2, 382 pp. . 1794. Entomologia systematica emendata et aucta, secundum classes, ordines, genera, species adjectis synonymis, locis, observationibus, des- criptionibus. Christ. Gottl. Proft, Hafniae, Vol. 4, vi + 434 pp. + Appendix pp. 435-462 + Expo- litiones specierum pp. 463-472 + Index, 5 pp. (unn.). 1803. Systema Rhyngotorum secundum or- dines, genera, species adjectis synonymis, locis, observationibus, descriptionibus. C. Reichard, Brunsvigae, x + 314 pp. Farrari, E. 1888. Die Hemipteren-Gattung Nepa Latr. (sens. natur.). Annalen des Naturhistorischen Mu- seums, Wien 3: 161-194, 2 pls. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Gerstaecker, A. 1873. Baron Carl Claus von der Decken’s Reisen in Ost.-Afrika. C. F. Winter, Leipzig, x + pp. 363-460, 3 col. pls. Gmelin, J. F. 1790. Systema naturae per regna tria naturae, secundum classes, ordines, genera, spe- cies cum characteribus, differentiis, synonymis, lo- cis. Georg Emanuel Beer, Lipsiae, Tomus I, Pars IV, pp. 1517-2224, Classis V, Insecta (Coleoptera, Hemiptera). Goeze, J. A. E. 1778. Entomologische Beytrage zu des Ritter Linné zw6lften Ausgabe des Natursys- tems. Weidmanns Erben und Reich, Leipzig, Vol. 2; xxii + 352 pp. Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, spe- cies, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. L. Salvii, Hol- miae, v + 824 pp. 1764. Museum Ludovicae Ulricae Reginae. L. Salvius, Holmiae, 8 + 720 pp. . 1767. Systema naturae per regna tria naturae secundum classes, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Edi- tio duodecima, Reformata. L. Salvii, Holmiae, vol. I, pars 2, pp. 533-1327 + 3 (Nomina generica) + 21 (Nomina specierum, Synonyma) + 2 (Termini artis) + 10 (Appendix synonymorum, Addenda). Linnavuori, R. 1971. Hemiptera of the Sudan, with remarks on some species of the adjacent countries. Annales Zoologici Fennici 8: 340-366. Lundblad, O. 1933. Zur Kenntnis der aquatilen und semiaquatilen Hemipteren von Sumatra, Java und Bali. Archiv fiir Hydrobiologie, 1933, Supplement Band 12, Tropische Binnengewasser 4: 1-195, 263- 498. Montandon, A.L. 1914. Naucoridae, Nerthridae, Be- lostomatidae et Nepidae. Jn Voyage de Ch. Al- luaud et R. Jeannel en Afrique Orientale (191 1- 1912). Résultats Scientifiques. Hemiptera, II. A. Schultz, Paris, pp. 115-130. Miiller, P.L.S. 1774. Vollstaéndiges Natursystem des C. v. Linné mit einer Eklaérung. Gabriel Nicolaus Kaspe, Niirnberg, 4 + 8 + 758 pp., 22 folding pls. Olivier, A.G. 1811. Encyclopédie Méthodique. Dic- tionnaire de Insectes. Pankouke, Paris, Vol. VIII, 2 Teil, Mou-Pan, 722 pp. Poisson, R. A. 1949. Hémiptéres Aquatiques. Ex- ploratin de la Parc Nationale Albert, Mission G. F. Witte, Fascicle 58, 94 pp. 1954. Hémiptéres Aquatiques. Exploration du Parc National de Upemba. Institut des Parcs Nationaux du Congo Belge, Fascicle 31, 53 pp. 1965. Catalogue de Hétéroptéres Hydrocor- ises Africano-Malgaches de la Famille des Nepi- dae (Latreille) 1802. Bulletin de la Institut Fran- cais d’Afrique Noire (A, 1)27: 229-269. Stal C. 1868. Hemiptera Fabriciana. 1. Konglika VOLUME 97, NUMBER 1 127 Svenska Vetenskaps-Akademien Handlingar 7(11): Turton, W. 1802. A general system of nature through 1-148. the three grand kingdoms of animals, vegetables, Stoll, C. 1780-88. Représentation exactement colo- and minerals; ... Translated from Gmelin’s last rée d’aprés nature des punaises, qui se trouvent edition of the celebrated Systema Naturae, by Sir dans les quatre parties du monde, l’Europe, l’Asie, Charles Linné, ammended and enlarged . . . Print- Afrique et l’Amerique, reassemblée et decrites. ed by Voss & Allen, Swansea, for Lackington, Al- Jan Christian Sepp, Amsterdam, 172 pp., 41 col. len & Co., London, Vol. II, 717 + 2 pp. (errata). pls. (Lief 1, pp. 1-20, Taf. 1-4, 1780; Lief 2, pp. 21-36, Taf. 5-8, 1780). PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 128-152 A REVIEW OF THE GENUS STRIDULIVELIA HUNGERFORD AND TWO NEW SPECIES (HETEROPTERA: VELIIDAE) FROM SOUTH AMERICA JOHN T. POLHEMUS AND PAUL J. SPANGLER (JTP) University of Colorado Museum, 3115 S. York, Englewood, Colorado 80110; (PJS) Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Abstract.—Two new species are described, Stridulivelia ayacucho from the Territorio Federal Amazonas of Venezuela and S. anta from Amazonas, Brazil. Stridulivelia nama (Drake 1957) is placed as a junior synonym of S. tersa (Drake & Harris 1941) (new synonymy). Habitat and distributional data and maps are given for the South American species along with a key to species of the nominate subgenus Stridulivelia. Key Words: Heteroptera, Veliidae, Stridulivelia, new species, water strider, synonymy, key to subgenera, key to species, South America Hungerford (1929) established the sub- genus Stridulivelia for those members of the genus Vel/ia Latreille 1804 that share a gen- eral facies that he characterized as follows: “The sides of the thorax are devoid of hair and sculptured with depressed figures. The middle legs are the longest. Two or more of the anterior ventral abdominal segments bear transverse grooves and all of the spe- cies, except Velia cinctipes Champion, 1898, are provided with a stridular patch on the hind femur and with a row of stridular pegs on the submargin of the abdomen.” Pol- hemus (1976) elevated Stridulivelia to ge- neric rank and later Polhemus (1979) es- tablished the subgenus Aenictovelia for the primarily Mesoamerican species that lack the stridulatory mechanism. The Meso- american species were revised by Drake and Menke (1962) and two additional species were added by Polhemus (1979) and Pol- hemus and Polhemus (1985); thus, the fau- na of that region is adequately known. This contribution, therefore, deals only with the South American species assigned to Stri- dulivelia—S. (A.) cinctipes (Champion); S. (S.) alia (Drake); S. (S.) anta Polhemus & Spangler, new species; S. (.S.) astralis (Drake & Harris); S. (S.) ayvacucho Polhemus & Spangler, new species; S. (S.) quadrispinosa (Hungerford); S. (S.) raspa (Hungerford); S. (S.) stridulata (Hungerford); S. (S.) strigosa (Hungerford); S. (S.) tersa (Drake & Harris); S. (S.) transversa (Hungerford). Of the five species in the subgenus Aenictovelia, only Stridulivelia (A.) cinctipes occurs in north- ern South America and Mesoamerica and is included in this study. Both authors have made substantial col- lections of these insects in South America and we believe the taxonomy, habitat, and distribution of the South American mem- bers of Stridulivelia (Stridulivelia) are now reasonably well understood. This informa- tion and the synonymy are given for each species and a key to the species of Stridu- livelia (Stridulivelia) is provided. Paratypes of the new species are depos- VOLUME 97, NUMBER 1 Fig. 1. Stridulivelia (Aenictovelia) epeixis (Drake & Menke): a, metasternum and scent gland channels; b, body, lateral view (from Drake and Menke 1962) ited in the U.S. National Museum of Nat- Genus Stridulivelia Hungerford ural History, Smithsonian Institution, Stridulivelia Hungerford, 1929: 55 (as sub- Washington (NMNH) and, as material per- genus of Velia Latreille, 1804). mits, paratypes will be deposited in the $trjdulivelia; Polhemus, 1976: 509 (raised American Museum of Natural History, New to generic rank). York (AMNH) and the J. T. Polhemus Col- Aenictovelia Polhemus, 1979: 46 (as sub- lection (JTPC). genus of Stridulivelia). 130 as — ‘ VE LOG | io are) oF es 5 —t G — / a hugh Baad pee a Se , + i PS / Yy ae oe f- ae X mo Ao 0 ° ° © had oS oO a S——— 0° fe} ate = ie C2 \ oO ee 4 a 2 aS } — wv = ( —- = —— \ Pat @\_* Fig. 2. Stridulivelia species, left parameres: a, S. (Stridulivelia) tersa (Drake & Harris); b, S. (Aenicto- velia) cinctipes (Champion). (From Drake and Menke 1962) Discussion: The genus Stridulivelia is re- stricted to the New World. Its Old World sister group is 7etraripis, a genus Andersen (1982) placed in the subfamily Rhagoveli- inae on account of a swimming plume re- sembling that of Rhagovelia; however, Te- traripis has been transferred by D. A. Polhemus (in press) to the Veliinae by rea- son of the morphology of the scent gland channels (i.e. angled anteriorly in Striduli- velia; Fig. 1a), fore tibial grasping comb in females, and swimming plumes of a differ- ent nature than in Rhagovelia, representing parallelism in these pretarsal structures. Stridulivelia Hungerford KEY TO SUBGENERA Stridulatory apparatus present in both sexes, con- sisting of rastrate patch on hind femur and pegs or ridges on connexival margin. Claws slender, downcurving arolia slender. ...........5 Stridulivelia Stridulatory apparatus absent in both sexes. Claws blade-like, downcurving arolia blade-like Discussion: The subgenus Aenictovelia is primarily Mesoamerican; however, S. (4.) cinctipes (Champion) occurs widely in PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON northern South America. This species and all others of the subgenus are easily distin- guished from those of the subgenus Stri- dulivelia by the lack of stridulatory struc- tures (Fig. 1b) and the complex (Fig. 2b) rather than simple (Fig. 2a) parameres. In the taxonomic treatment below, a key is giv- en only for species of the subgenus Stridu- livelia, which is restricted to South America including Trinidad. Stridulivelia (Aenictovelia) cinctipes (Champion) Figs. 2b, 3 Velia cinctipes Champion, 1898: 143, pl. 9, fig. 9. (Type from Santarem, Brazil; in BMNH.) Velia (Stridulivelia) cinctipes; Hungerford, 1929-55. Stridulivelia cinctipes; Polhemus, 1976: 509. Stridulivelia (Aenictovelia) cinctipes; Pol- hemus, 1979: 46. Material examined. —COLOMBIA: An- tioquia: 4 6 micr., 4 2 micr., trib to Rio Claro, W of Doradal, CL 2406, 21.VII.1989, J. T..& D: A. Polhemus (}FP@); 1 6 micr.. 1 6 macr., 2 2 macr., Quebrada Cristalina, 18 km W of Doradal on Hwy. 60, 350 m, water temp. 22.5°C, CL 2407, 22. VII.1989, J.T. & D. A. Polhemus (JTPC). GUYANA: Mazaruni-Potaro Dist.: 1 @ micr., Takatu Mtns., 6°15’N, 59°5’W, 3-10.XII.1983, P. Spangler, R. Faitoute (NMNH). VENE- ZUELA: Amazonas: 3 ? micr., 42 km S. Puerto Ayacucho, Tobogan, CL 2371, 19.1.1989, J. T. Polhemus (JTPC); 3 4 micr., 5 2 micr., 42 km S. Puerto Ayacucho, small trib to Cano Coromoto, at Tobogan, CL 2388, 26.1.1989, J. T. Polhemus (JTPC); 1 é micr., tiny stream, trib to Rio Gavilan, nr. bridge, CL 2379, 23.1.1989, J. T. Pol- hemus (JTPC). Dist. Federal: 2 6 micr., 1 9° micr., Los Caracas, 19.1.1985, P. Spangler, R.; Faitoute,. W... Steiner; A;. Conover (NMNH). Known distribution (Fig. 3).— Brazil, Co- lombia, Costa Rica, Guatemala, Guyana, VOLUME 97, NUMBER 1 131 Fig. 3. Stridulivelia (Aenictovelia) cinctipes (Champion), known South American distribution. Mexico, Panama, Venezuela. Only South American localities are plotted. Discussion.—This predominantly mi- cropterous species is easily distinguished from all other South American species of the genus by the lack of an evident stridu- latory mechanism and the usual presence in males of a long stout spine on the hind tro- chanter. The length of this spine on the hind trochanter is extremely variable, essentially lacking in some specimens and extremely long in others. Stridulivelia cinctipes 1s com- mon throughout Panama and Costa Rica, where we have collected many specimens; it was reported from Guatemala and Mex- ico by Drake and Menke (1962), but we have not seen specimens from Mexico. I. Stridulivelia (Stridulivelia) Hungerford KEY TO SPECIES First five (visible) ventral abdominal segments with transverse glabrous grooves. (Male also has small groove on segment VI) .......... transversa (Hungerford) First four or fewer ventral abdominal segments with transverse glabrous grooves ........... 2 . First four ventral abdominal segments with transverse glabrous grooves (female may have just a spot on segment IV; male has small groove on segment V, Fig. 18b) .. tersa (Drake & Harris) First three or fewer ventral abdominal seg- ments with transverse glabrous grooves Pronotal humeral angles spinose (Fig. 4a) ... A BAe Ets Le dele ate i Rei eens alia (Drake) Pronotal humeral angles not spinose (Figs. | Sa, 18a) 132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4. First two ventral abdominal segments with transverse glabrous grooves ............... 5) First three ventral abdominal segments with transverse glabrous grooves ............... 6 5. Length of second antennal segment subequal to width of head across eyes (male) or at most 1.13 times greater (female). Male without a dig- itate process ventrally on first genital segment. Female without spine-like processes on pos- terior margin of abdominal tergite VII ...... I Ae AR ery eas ce ae ayacucho, n. sp Length of second antennal segment clearly greater than width of head across eyes; 1.22 (female) to 1.38 (male) times greater. Male with a digitate process ventrally on first genital seg- ment. Female with two posteriorly directed spine-like processes on posterior margin of ab- dominal tergite VII ...... stridulata (Hungerford) 6. Length of first antennal segment at least 1.5 times width of head across eyes. Body and legs long SlEndere meet ioe strigosa (Hungerford) Length of first antennal segment no more than 1.2 times width of head width across eyes. Body stout, legs stout, hind femur incrassate 7. Length of first antennal segment about 1.2 times width of head across eyes. Males with pair of prominent projections ventrally on last ab- dominal segment. Stridulatory patch on hind femur with about 26-32 sharp pegs not orga- MIZEGHNETOW SH er ae etc: anta, n. sp. Length of first antennal segment equal to or less than width of head across eyes. Males with or without pair of prominent projections ven- trally on last abdominal segment. Stridulatory patch on hind femur may or may not be or- PAaniZedineTOWSEer ye Tea eee oe 8 8. Length of first antennal segment about 0.8 width of head across eyes. Males without pair of prominent projections ventrally on last ab- dominal segment. Female genital segment, in dorsal view, sharply triangular with length equal to basal width. Stridulatory patch on hind fe- mur organized in about 17 rows of tiny spinulae Th TAA cette tet ee astralis (Drake & Harris) Length of first antennal segment about equal to width of head across eyes. Males with pair of prominent projections ventrally on last ab- dominal segment. Female genital segment, in dorsal view, rounded or broadly triangular with length clearly less than basal width. Stridula- tory patch on hind femur may or may not be organized in rows 9. Stridulatory patch on hind femur with about 40 sharp pegs not organized in rows; connex- ival margin with a row of fine vertical ridges. Male (ventrally) and female (dorsally) with prominent projections posteriorly on seventh abdominal segment .. qguadrispinosa (Hungerford) Stridulatory patch on hind femur organized in about 17 rows of tiny spinulae; connexival margin with a thin row of tiny pegs. Male (ven- trally) with prominent projections posteriorly on seventh abdominal segment; female (dor- sally) without prominent projections posteri- orly on seventh abdominal segment ........ Jaan asG ee raspa (Hungerford) Stridulivelia (Stridulivelia) alia (Drake) Figs. 4, 5 Velia alia Drake, 1957: 115 (Type from Bar- tica District, Guyana; in the National Museum of Natural History). Stridulivelia alia; Polhemus, 1976: 509. Material examined (all JTPC, unless not- ed).—BRAZIL: Amazonas: 6 6 micr., 2 6 macr., 4 2 micr., | 6 macr., Reserva Ducke, 25 km NE of Manaus, Igarape de Anta, 60 m, water temp. 24.5°C, 25.VIII.1989, CL 2472, J. T. and D. A. Polhemus; 1 4 micr., 8 6 macr., 5 2@macr., Reserva Ducke, Igarape Barro Branco, nr. headquarters, 50 m, 27.VIII.1989, CL 2475, J. T. and D. A. Pol- hemus; 6 6 micr., 33 6 macr., 9 ? micr., 39 ? macr., forest stream at INPA forest man- agement station, 98 km NW Manaus, 90 m, water temp. 25°C, 29 Aug. 1989, CL 2477, J. T. & D. A. Polhemus; 7 4 macr., 15 ? macr., stream near viewing tower, 90 km NW Manaus, 90 m, 29 Aug. 1989, CL 2478, J. T. & D. A. Polhemus; 29 6 micr., 26 2 micr., — stream mr, @Eglen Reserve: 30. VIII.1989, CL 2479, J. T. and D. A. Pol- hemus. Para: 5 6 macr., 3 2 macr., Rio Xin- gu, camp, 52°22’W, 3°39’S, ca. 60 km S Al- tamira, 3.X.1986, P. Spangler, O. Flint (NMNH). GUYANA: 6 4 micr., | 2 micro, 40 km S Georgetown, 4°29.9'N, 58°13.1'W, 13.1V.1994, P. J. Spangler, colln #27 (NMNH); | 6 micro, | 2 micro, 1 2 macr., Dubulay Ranch, 5°39.8'N 57°53.4'W, War- niabo River, 10.1V.1994, P. J. Spangler & R. Parris, colln #24 (NMNH). SURINA- ME: 1 6 macr., 1 @ macr., Zanderijsavanne, 4.1X.1969, SN 117B, N. Nieser; 1 2 micr., Coesewijn Project, 24.111.1970, SN 371, N. Nieser; | 6 micr., 1 2 micr., Coesewijn Pro- VOLUME 97, NUMBER 1 133 : Wael) ; mnt}, iv Ky {\ sh Mt ii. DN i | "il Fig. 4. Stridulivelia (Stridulivelia) alia (Drake): a, dorsal habitus, micropterous female; b, connexiva, lateral view, showing stridular file of fine vertical ridges; c, hind femur, showing stridular denticles. 134 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 5. ject, 3.111.1970, SN 368, N. Nieser; 1 2 macr., Ist streamlet, Hanover Rd., 4. VIII.1969, SN 061, N. Nieser; 1 6 micr., Troelikreek, 18.X1.1969, SN 237, N. Nieser. VENE- ZUELA: Amazonas: 9 ¢ micr., 9 2 micr., small clear stream with sandy bottom, 0.5 km N of Alto Mavaca Base Camp, 2°1'30’N, 65°7'0"W, 228 m, water temp. 22°C, CL 8006, 4.11.1989, D. A. Polhemus; 2 6 micr., 1 2 micr., small shallow forest stream, trib to upper Rio Siapa, 1°43’N, 64°30’W, 635 m, water temp. 21.5°C, CL 8011, 8.11.1989, D. A. Polhemus. Known distribution (Fig. 5).— Brazil, Guyana, Suriname, Venezuela. Discussion. — Stridulivelia alia (Drake) is the most easily distinguished Stridulivelia Stridulivelia (Stridulivelia) alia (Drake), known distribution. species because of the unique spines on the humeral angles of both sexes. Stridulivelia (Stridulivelia) anta Polhemus and Spangler, NEw SPEcIES Figs. 6-8 Diagnosis.—Stridulivelia anta, new spe- cies, is most closely related to S. quadris- pinosa (Hungerford 1929) and S. alia Drake. It differs from S. guadrispinosa by the small- er size, shorter antennae, fewer stridulatory pegs on the hind femur, and differently formed abdominal terminalia in both males and females. Stridulivelia anta is very sim- ilar to S. alia (Fig. 4) in body size, shape, coloration and stridulatory mechanism; however, S. anta is distinguished by the ab- VOLUME 97, NUMBER 1 sence of large spines on the humeral angles, longer glabrous groove on male abdominal sternite IV, less tumid male genital seg- ments, and male proctiger with much short- er and blunt lateral protuberances (Fig. 7). The female of S. anta is without spines pro- truding posteriorly from the first genital seg- ment. Description. — Micropterous male: Length, mean = 4.09 mm (N = 10, min. 3.77, max. 4.50). Width, mean = 1.07 mm (N = 10, min: 1.00; max. 1.17): Ground color orange brown, venter somewhat lighter. Pronotum anteriorly lighter, anterolaterally with prominent tri- angular silvery spots. Head light orange brown; tips of antennal tubercles, bucculae, anteclypeus yellowish; rostrum yellowish brown, distal segment black. Legs and an- tennae yellow to light brown, lighter ven- trally; antennal segment 4 broadly lighter medially. Head short, almost vertical anteriorly; bucculae prominent, short; rostral cavity closed behind; collar not prominent, set off by a few weak pits; length 0.39; width of eye/interocular space, 0.22/0.28. Pronotum long, humeri not prominent; weakly cari- nate on midline, carina evanescent anteri- orly and posteriorly; with shallow pits, de- pressed part of each pit with several glabrous light reflecting facets; disc raised; posterior margin rounded at apex; length : width, 1.00: 1.05. Micropterous wing pads with silvery setae, reaching onto base of abdominal ter- gite I. Dorsum clothed with short semi-erect pubescence and scattered longer setae. Ab- dominal tergites I, II-VI subequal in length (0.22-0.28), III, VII longer (0.33); connex- ival margins of II, III, and basal part of IV with a stridular file of fine vertical ridges (similar to those of S. alia, Fig. 4b); distal connexival spines acuminate, extending to distal part of first genital segment. Abdom- inal sternite II (first visible) medially cari- nate; VII modified, with stout tubercles pos- teroventrally; sternites H-IV with long, 135 t+—— 0.2 mm ——__4 Fig. 6. Stridulivelia (Stridulivelia) anta Polhemus & Spangler, n. sp., male paramere: a, lateral view; b, anterior view; c, posterolateral view. Fig. 7. Stridulivelia (Stridulivelia) anta Polhemus & Spangler, n. sp., male genital capsule, lateral view. glabrous, transverse striae laterally, striae shorter on IV. Legs and antennae thickly clothed with short to moderate length setae; with scattered longer setae ventrally on all femora and tibiae. Posterior trochanters un- armed. Posterior femur tumid, denticulate ventrally, basally with a few scattered den- ticles increasing in length and frequency dis- 136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Stridulivelia (Stridulivelia) anta Polhemus & Spangler, n. sp., known distribution. tally, also with a stout tooth at distal 7 fol- lowed by a distal row of small denticles; stridulatory patch small, dorsal, at basal 1%, consisting of 26-32 small but stout, black denticles not organized into rows (similar to those of S. alia, Fig. 4c). Posterior tibia ventrally with numerous denticles over en- tire length, not organized into rows; with a very large stout apical spur. Antennal formula I:II:HI:1V; 0.80:0.56: 0.56:0.56. Claws long, slender, preapical; dorsal and ventral arolia slender, promi- nent. Measurements of legs as follows: Fe- mur, tibia, tarsal-1, tarsal-2, tarsal-3 of male fore-leg, 0.97, 0.97, 0.06, 0.05, 0.22; mid- dle-leg, 1.55, 1.66, 0.11, 0.61, 0.42; hind- leg 12565 1689, '0106,°0:23, 039% Male genital capsule as shown in Fig. 7. Proctiger sculptured, with blunt lateral pro- jections (Fig. 7). Paramere long, slender (Fig. 6). Macropterous male: Length, mean = 4.42 mm (N = 3, min. 4.33, max. 4.50). Width, mean = 1.44 mm (N = 3, min. 1.44, max. 1.44). Similar in most respects to microp- terous male, except pronotum longer and wider, broadly V-shaped posteriorly. He- melytra chocolate brown, reaching to tip of abdomen; each with two (1 + 1) spots of silvery setae, one basally, the other along costal margin at basal '4; distally with single median, light colored, V-shaped mark opening posteriorly, sometimes followed by light streak; basal half with scattered patch- es of short golden setae, absent medially and distally. VOLUME 97, NUMBER 1 Micropterous female: Length, mean = 4.12 mm (N = 10, min. 4.00, max. 4.27). Width, mean = 1.14 mm (N = 10, min. 1.11, max. 1.17). Color and most structures as in male, except slightly broader. First genital segment truncate, with slight lateral posteriorly directed projections. Macropterous female: Length, mean = 4.38 mm (N = 4, min. 4.22, max. 4.61). Width, mean = 1.44 mm (N = 4, min. 1.39, max. 1.50). Except for sex, similar to ma- cropterous male. Ecological notes. — At the type locality, S. anta was taken along with 5 other species of Stridulivelia—S. alia (Drake), S. tersa (Drake and Harris), S. stridulata (Hunger- ford), S. strigosa (Hungerford), and S. trans- versa (Hungerford); thus 6 of the 10 known species of this genus were collected at a sin- gle site. The type locality is a moderately sized, clear, headwater stream flanked with small springs and pools in the original rain forest of Reserva Ducke, a biotope located in low sandy hills. Etymology.—The name anta, a noun in apposition, refers to the type locality, Igar- ape da Anta, in Reserva Ducke, Amazonas, Brazil. Material examined.— Holotype, microp- terous male: BRAZIL: Amazonas, Reserva Ducke, 25 km NE Manaus, Igarape da Anta, 60 m, water temp. 24.5°C., 25 Aug. 1989, CL 2472, J.T. & D.A. Polhemus (INPA). Paratypes: BRAZIL: Amazonas: 9 4 micr., 11 2 micr., same data as holotype (JTPC, AMNH, NMNH); 8 6 micr., 9 2 micr., Reserva Ducke, 25 km NE Manaus, Igarape Barro Branco, 50 m, water temp. 2BES2©-po Aes 1989 C2475... 7 s&:D: A. Polhemus (JTPC); 3 6 macr., 4 2 macr., stream near viewing tower, 90 km NW Ma- naus, 90 m, 29 Aug. 1989, CL 2478, J. T. & D. A. Polhemus (JTPC); 2 6 macr., rain- forest stream at INPA Forest Mgnmt. Sta., 98 km NW Manaus, 90 m, 29 Aug. 1989, C241 72); Pe Di As Polhemus (J TPC): VENEZUELA: Amazonas: 2 6 micr., Cerro de la Neblina, white water str., | km S. Base- 137 camp, O°SON, 66°10W, 140 m, 8.1I.1985, W. E. Steiner, R. Holling (NMNH). Known distribution (Fig. 8).— Brazil, Venezuela. Stridulivelia (Stridulivelia) astralis (Drake & Harris) Fig. 9 Velia astralis Drake & Harris, 1938: 200 (Type from Matto Grosso, Brazil; in Mu- seo de La Plata). Stridulivelia astralis; Polhemus, 1976: 509. Material examined (all JTPC, unless not- ed).— BRAZIL: Goias: 1 ¢ micr., 1 2 micr., 2 6 macr., 1 2 macr., 48 km S. Peixe, 1. VI1956, EF: Truxal; Matto ‘Grosso: 16 micr., 1 2 micr., Capitao Vasconselos, on Rio Tuatuari, Up. Xingu _ Basin, 31.VII.1957, B. Malkin. PARAGUAY: Cordillera: 2 2 macr., Dist. Caacupe, Ca- banas, 9.XII.1980, R. D. Cave (NMNH),; 1 2 micr., Depto. Cordillera, Piribebuy, Rio Piribebuy, 3.V.1985, T. Bonace (NMNH). Central: 1 2 macr., Asuncion, 9.XI.1974, J. Sedlacek; 1 2 macr., Horqueta, 17.XII.1974, Alperto Schulze. Known distribution (Fig. 9).— Brazil, Paraguay. Discussion.—S. astralis and S. raspa are small, stout-bodied, closely related species with a small, rastrate stridulatory patch on the hind femur. They may be distinguished by the characters given in the key. Stridulivelia (Stridulivelia) ayacucho Polhemus and Spangler, NEw SPECIES Figs. 10-12 Diagnosis.—Stridulivelia ayacucho, new species, most closely resembles S. guadris- pinosa (Hungerford) in size, general habitus, female abdominal terminalia, and most other features. However, the male of S. ay- acucho lacks the paired protuberances found on abdominal sternite VII of S. guadrispi- nosa and the stridulatory patch on the hind femur is a rastrate area of parallel sclero- 138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 9. Stridulivelia (Stridulivelia) astralis (Drake & Harris), known distribution. tized ridges unlike the patch of black den- ticles found on S. quadrispinosa. Stridulivelia ayacucho is perhaps most closely related to S. stridulata (Hungerford) (Fig. 15b, c), as the stridulatory mechanisms are the same, as well as most other features. In S. stridulata, however, the abdomen is more strongly narrowed in both sexes (Fig. 15a); the males have longer upturned con- nexival spines, spatulate parameres, and a strong digitate process on the first genital segment (Hungerford, 1929, Figs. 3, 8), and the females have two long processes on ab- dominal tergite VIII (Fig. 15a). Description. — Micropterous male: Length, mean = 4.73 mm (N = 10, min. 4.38, max. 4.99). Width (across base of abdomen), mean = 1.32 mm (N = 10, min. 1.28, max. 1.39). Ground color brown. Abdominal venter yellowish on each side of midline and along connexival margins. Pronotum anteriorly slightly lighter, anterolaterally without sil- very spots; connexival margins yellowish. Head light brown; antennal tubercles light- er; rostrum yellowish brown, distal segment black. Legs and antennae yellow to light brown, lighter ventrally. Posterior femur with broad dark annulus just beyond middle and another distally. Antennal segment 4 light on distal %. Head short, almost vertical anteriorly; bucculae prominent, short, rostral cavity closed behind; collar not prominent, set off by a few weak pits plus elongate transverse pit behind each eye; length 0.47; width of eye/interocular space, 0.28/0.28. Pronotum VOLUME 97, NUMBER 1 long, humeri not raised; weakly carinate on midline, carina evanescent posteriorly; with shallow pits, depressed part of each with several glabrous light reflecting facets; disc raised; posterior margin rounded at apex, set off by row of pits; length: width, 1.22: 1.17. Micropterous wing pads with silvery setae, reaching onto base of abdominal ter- gite I. Dorsum clothed with short, semi-erect pubescence and scattered, dark, erect, long setae. Abdominal tergites I-VI subequal in length (0.28-0.33), VII longer (0.39); con- nexival margins I-IV with stridular file of about 30 black, knob-like denticles (similar to that of S. stridulata, Fig. 15b); distal con- nexival spines long, acuminate, extending to middle of genital segments. Abdominal sternite II (first visible) medially carinate; VII modified, tumescent, tumescence abruptly terminating posteriorly; sternites II-III with long, glabrous, transverse striae laterally, located about midway between in- tersegmental sutures. Legs and antennae thickly clothed with short to moderately long setae; with scattered longer setae ventrally on all femora and tibiae. Posterior trochan- ters with 8-10 tiny black denticles. Posterior femur tumid, denticulate ventrally; with 2 ragged parallel rows of small denticles plus a few more distally, all subequal in size, except stout tooth at distal 74 on anterior row; dorsally near base with long, broad stridulatory patch consisting of about 50 parallel sclerotized ridges; each ridge inter- rupted 7 to 20 times, ridges very thin and closely packed anteriorly, progressively coarser posteriorly, extending onto poste- rior face of femur (similar to that of S. stri- dulata, Fig. 15c). Posterior tibia ventrally with numerous denticles over entire length, mostly organized into 2 rows; with a stout apical spur. Antennal formula [:II:HI:IV; 1.10:0.89: 0.69:0.61. Claws long, slender, preapical; dorsal and ventral arolia slender, prominent. Measure- ments of legs as follows: Femur, tibia, tar- 139 ————O2mm a Fig. 10. Stridulivelia (Stridulivelia) ayacucho Pol- hemus & Spangler, n. sp., male paramere: a, lateral view; b, anterior view; c, posterolateral view. Fig. 11. Stridulivelia (Stridulivelia) ayacucho Pol- hemus & Spangler, n. sp., male genital capsule, lateral view. sal-1, tarsal-2, tarsal-3 of male fore-leg, 1.22 1.22, 0.06, 0.05, 0.22; middle-leg, 1.89, 2.05, 0.06, 0.61, 0.50; hind-leg, 1.72, 1.83, 0.06, 0:39.40533: Male genital capsule as shown in Fig. 11. Proctiger sculptured, with acuminate lateral 140 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 12. Stridulivelia (Stridulivelia) ayacucho Polhemus & Spangler, n. sp., known distribution. projections (Fig. 11). Paramere long, broad, blade-like (Fig. 10). Macropterous male: Unknown. Micropterous female: Length, mean = 4.93 mm (N = 10, min. 4.77, max. 5.16). Width (across base of abdomen), mean = 1.45 mm (N = 10, min. 1.39, max. 1.50). Color and most structures as in male, except slightly broader. First genital segment trun- cate; with prominent, acuminate, lateral, posteriorly directed projections. Macropterous female: Length, mean = >/30mme(N = 2eemin: (5227maxs' 5233), Width (across humeral angles), mean = 1.78 mm (N = 2, min. 1.78, max. 1.78). Similar in most respects to microperous female, ex- cept pronotum longer and wider, broadly V-shaped posteriorly, caudally with short (0.14) posteriorly projecting digitate pro- tuberance. Hemelytra chocolate brown, reaching to tip of abdomen; each with two (1 + 1) spots of silvery setae, one large tri- angular spot basally, another small spot along costal margin at basal '; distally with single median white almost circular mark, with small V-shaped notch posteriorly; with scattered patches of short golden setae ba- sally, patches absent medially and distally. Habitat data.— The type locality is a flow- ing stream in original forest, with alternat- ing long pools and gravel bottomed riffles. The insects were taken from a low, partially VOLUME 97, NUMBER 1 flooded cavity deeply hollowed out under a steep bank laced with tree roots. Etymology.—The name ayacucho, a noun in apposition, refers to the city of Puerto Ayacucho, which 1s near the type locality. Holotype, apterous male: VENEZUELA: Amazonas: 21 km S. Tobogan Jct. (53 km S of Pto. Ayacucho), “Missionary Stream,” small river with alternating riffles and pools, Cra23 75, 221.1989. J 1; Polhemus (NMNH). Paratypes: BRAZIL: Para: 1 2 micr., Rio Xingu, camp, 52°22'W, 3°39’S, ca. 60 km S Altamira, 3.X.1986, P. Spangler, O. Flint (NMNH). GUYANA: 1 6 macr., Ishezr.- Tun., Lat 2°N, 16 km E of Rupununi Riv., 1937, Terry-Hulden Exp. (JTPC). PARA- GUAY: Paraguari: 3 6 micr., 4 2 micr., Cer- ro Acahay, Arroyo, 31.V.1985, T. Bonace (NMNH). PERU: Madre Dios: | ¢ micr., 1 @ micr., Parque Manu, Pakitza, 12°07’S, 70°58'W, 250 m, side pools of stream, colln. 39, 13.1X.1989, R. A. Faitoute (NMNH); | @ micr., Parque Manu, same, but stream, colln. 44, 17-18.1X.1989, R. A. Faitoute (NMNH). VENEZUELA: Amazonas: 5 4 micr., 10 2 micr., same data as holotype (JTPC); 1 2 micr., 6 km S. Tobogan Jct. (38 km S of Pto. Ayacucho), brook, CL 2374, 22.1.1989, J. T. Polhemus (JTPC); 4 6 micr., 10 2 micr., tiny stream, trib to Rio Gavilan, nr bridges ‘CLE 2379, 23.71.1989, J. T. Pol- hemus (JT PC); 2 2 micr., 16 km S Pto. Aya- cucho, Puente Pulda, CL 2387, 26.11.1989, J. T. Polhemus (JTPC); 2 6 micr., 4 2 micr., small shallow forest stream, trib to upper Rio Siapa, 1°43’N, 64°30’W, 635 m, water temp lesc@. Cl SOll Ss IL1989 Dp. A. Polhemus (JTPC, NMNH); 23 6 micr., 23 2 micr., small clear stream with sandy bot- tom, 0.5 km N of Alto Mavaca Base Camp, 2°1'30"N, 65°7'0”W, 228 m, water temp. 22°C, CL 8006, 4.11.1989, D. A. Polhemus (JTPC, NMNH); | é micr., 1 2 micr., 39 km S Pto. Ayacucho, brook, 15.XI.1987, P. Spangler, R. A. Faitoute (NMNH). Known distribution (Fig. 12).—Brazil, Guyana, Paraguay, Peru, Venezuela. 14] Stridulivelia (Stridulivelia) quadrispinosa (Hungerford) Fig. 13 Velia (Stridulivelia) quadrispinosa Hunger- ford, 1929: 52. (Type from Santarem, Brazil; in BMNH.) Stridulivelia quadrispinosa; Polhemus, 1976: 509. Material examined.—BOLIVIA: Beni: 2 $6 macr., 2 2 macr., rainforest stream 40 km S of Rurrenabaque, nr. Monte Redondo SawmillCE 2514, 12.1X61989. Jo ide. D: A. Polhemus (JTPC). BRAZIL: Amazonas: 1 6 macr., Rio Crynyn, Missao, A 88-1, 1211961. E. J. Fittkau @@EO)s Para: 279 macr., Rio Xingu, camp, 52°22’W, 3°39’S, ca. 60 km S Altamira, 3.X.1986, P. Span- gler, O. Flint (NMNH). GUYANA: Lethem (30 km SE), 3°18’N, 59°39’W, Moco Moco River, 3.1V.1994, Paul J. Spangler, 1 2 macr. (NMNH). PERU: Loreto: | 6 macr., 2 2 macr., stream near Napo Camp, CL 2459, 15.VIII.1989, J. T. & D. A. Polhemus (JTPC); 1 émacr., 1 2 macr., small rainforest stream near Explorama Inn, CL 2465, 18.VIII.1989, J. T. & D. A. Polhemus (JTPC). VENEZUELA: Amazonas: 2 4 macr., 3 2 macr., 21 km S. Tobogan Jct. (53 km S of Pto. Ayacucho), ‘“‘Missionary Stream,” small river with alternating riffles and: pools. CL, 237 5.22 11989) J. 1. bol hemus (JTPC); 3 6 macr., 3 2 macr., small clear stream with sandy bottom, 0.5 km N of Alto Mavaca Base Camp, 2°1'30’N, 65°70" W228) 1m: water temp: 22°C. CL 8006, 4.11.1989, D. A. Polhemus (JTPC). Known distribution (Fig. 13).— Bolivia, Brazil, Guyana, Peru, Venezuela. Discussion.— This species was described from Brazil. We have specimens from sev- eral localities matching the original descrip- tion and figures given by Hungerford (1929), but the species seems to be uncommon. In three small series from Loreto Dept., Peru, and Beni Dept., Bolivia, the males have the first genital segment more tumid ventrally and the females lack the strongly 142 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 13. Stridulivelia (Stridulivelia) quadrispinosa (Hungerford), known distribution. projecting spines on tergite VIII; otherwise these specimens match those from Vene- zuela and Brazil in every respect. For the present, these are considered as variant pop- ulations of S. guadrispinosa. Stridulivelia (Stridulivelia) raspa (Hungerford) Fig. 14 Velia (Stridulivelia) raspa Hungerford, 1929: 51. (Type from Manacaparu, Amazonas, Brazil; in SEMC.) Stridulivelia raspa; Polhemus, 1976: 509. Material examined (all JTPC).—BRA- ZIL: Amazonas: 2 2 micr., Rio Madeira, Ig. (Igarape) Tres Casas, 10.XI.1941, S67, H. Sioli (JTPC); 2 2 micr., Rio Madeira, Ig. Tres Casas, 12.XI.1941, S73, H. Sioli GLRPEC): Known distribution (Fig. 14).— Brazil. Discussion. — This species has been found only in the lowlands west and south of Ma- naus, Amazonas, Brazil, and is one of eight Stridulivelia species now known from the vicinity of Manaus. Stridulivelia (Stridulivelia) stridulata (Hungerford) Figs. 15, 16 Velia (Stridulivelia) stridulata Hungerford, 1929: 53. (Type from Guyane, Haut-Car- sevenne; in Paris Museum.) Stridulivelia_ stridulata; Polhemus, 1976: 509. VOLUME 97, NUMBER 1 Fig. 14. Stridulivelia (Stridulivelia) raspa (Hungerford), known distribution. Material examined (all JTPC unless not- ed).—BRAZIL: Amazonas: 4 ¢ micr., 4 2 micr., Reserva Ducke, 25 km NE of Ma- naus, Igarape da Anta, 60 m, water temp. 245°C. 25. VIL L989. (C2472. T. and D. A. Polhemus; 9 ¢ micr., 11 2 macr., Re- serva Ducke, Igarape Barro Branco, nr. headquarters, 50 m, 27.VIII.1989, CL 2475, J. T. and D. A. Polhemus; 4 2? micr., forest stream at INPA forest management station, 98 km NW Manaus, 90 m, water temp. 25°C, 29 Aug. 1989, CL 2477, J. T. & D. A. Pol- hemus; | 4 micr., | 2 micr., Igarape Acara, 4 km SE of Ducke Reserve Hdq., 60 m, water temp. 24.5°C, 26 Aug. 1989, CL 2474, J iu& DA. Polhemus: 1. 9: miucr., Rio Cuieiras, Branquinho, 22.VII.1969, A 214- 1,E. J. Fittkau; 1 6 micr., 2 2 micr., Manaus, Ig. (Igarape) Gigante, 3.VII.1961, A 198-6, E. J. Fittkau (JTPC, ZSMC); 3 2 micr., Lago Salgado, Ig. S. Benedito, 15.1V.1948, A 223, H. Sioli (JTPC, ZSMC). SURINAME: 1 ¢ micr., Zanderijsavanne, Carolinakreek, 22.VIII.1969, SN 095, N. Nieser; 3 6 micr., 1 @ micr., Brokopondo, Kakaterekreek, 3.X1.1969, SN 216, N. Nieser; 1 6 micr., Saramacca, road to Saramacca-brug, 12 km W of jct. with rd. to Matta, 26.1.1970, SN 299, N. Nieser. Known distribution (Fig. 16).—Brazil, French Guiana, Suriname. Discussion.— This species is known only from the micropterous form (Fig. 15a). It is the largest Stridulivelia species known. 144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 15. Stridulivelia (Stridulivelia) stridulata (Hungerford): a, dorsal habitus, micropterous female; b, con- nexiva, lateral view, showing row of stridular denticles; c, hind femur, showing rastrate stridular structure. VOLUME 97, NUMBER 1 Fig. 16. Stridulivelia (Stridulivelia) stridulata (Hungerford), known distribution. Stridulivelia (Stridulivelia) strigosa (Hungerford) igs 7, Velia (Stridulivelia) strigosa Hungerford, 1929: 50. (Type from Guyane, Haut-Car- sevenne; in Paris Museum.) Stridulivelia strigosa; Polhemus, 1976: 509. Material examined (all JTPC, unless not- ed).—BRAZIL: Amazonas: 1 ¢ micr., 1 ? micr., Reserva Ducke, 25 km NE of Ma- naus, Igarape da Anta, 60 m, water temp. DA SoC. 325. VII 1989. 2G 2472.2)--Pyand D. A. Polhemus; 10 4 micr., 4 2 micr., Re- serva Ducke, 25 km NE Manaus, Igarape Barro Branco, 50m, water temp: 23.5°C., 27 Aug. 1989, CL 2475, J. T. & D. A. Pol- hemus (JTPC); 1 6 macr., 1 2 micr., stream near viewing tower, 90 km NW Manaus, 90 m.29" Aug. 1989) CL2478. Jaiiark& DivA: Polhemus; 3 ¢ micr., 1 2 micr., rainforest stream at INPA Forest Mgmt. Sta., 98 km NW Manaus, 90 m, 29 Aug. 1989, CL 2477, J.T. & D. A. Polhemus; 3 6 micr., 2 2 micr., Reserva Ducke, 25 km NE of Manaus, Igar- ape Acara, 26. VIII.1989, CL 2474, J. T. and D. A. Polhemus; 2 ¢ micr., 2 2 micr., stream nr. Egler Reserve, 30.VIII.1989, CL 2479, J. T. and D. A. Polhemus; 5 ¢ micr., 5 2 micr., Rio Negro, Igarape Barro Branco, 30.VI1I.1962, A 400, E. J. Fittkau (JTPC, ZSMC). Matto Grosso: 9 ¢ macr., 11 2 macr., Serra Roncador, 16-17.VIII.1965, A 558, E. J. Fittkau (JTPC, ZSMC). Para: 1 6 macr., Quellgebeit des Rio Xingu, Fliessgewasser zwischen Goiana und Cuiaba, 5.1X.1965A 146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 17. 574-2, E. J. Fittkau (ZSMC); 2 6 macr., Rio Xingu, camp, 52°22'W, 3°39’S, ca. 60 km S Altamira, 3.X.1986, P. Spangler, O. Flint (NMNH). GUYANA: 15 6 micro, 10 2 mi- cro, Dubulay Ranch, 5°39.8’N, 57°53.4'W, Warniabo River, 10.1V.1994, P. J. Spangler & R. Parris, colln #24 (NMNH). PERU: Madre Dios: 1 2 micr., Parque Manu, Pak- itza, 12°07'S, 70°58’W, 250 m, side pools of stream, colln. 42, 16.1X.1989, R. A. Fai- toute (NMNH). SURINAME: 1 ¢ micr., 1 2 micr., Zanderijsavanne, Ist trib. Colak- reek crossing rd. to Matta, SN 116, N. Nies- er; 1 émicr., Zanderijsavanne, 28. VIII.1969, SN 099, N. Nieser; 1 6 micr., 3 2 micr., Zan- derijsavanne, Carolinakreek, 22. VII.1969, SN 092, N. Nieser; 7 6 micr., 6 2 micr., Zander- ysavanne, Ist trib. Colakreek, 8.1X.1969, SN Stridulivelia (Stridulivelia) strigosa (Hungerford), known distribution. 127, SN 128, N. Nieser; 21 4 micr., 23 2 micr., 2 6 macr., Saramaca, 2nd trib. Troelikreek, 18.X1.1969, N. Nieser; 1 ¢ micr., Sabakoek- reek, 29.VII.1965, SN 044, N. Nieser; 2 6 micr., 3 2 micr., Carolinakreek, 8.1V.1962, P. H. van Doesburg; | ¢ micr., Bosgivak, Creek, 29. XII.1950, Geiskes. VENEZUELA: Ama- zonas: | 2 micr., Cerro de la Neblina, Base- camp, margins Rio Baria, 0°50'N, 66°10'’W, 140 m, 20.11.1985, P. J. & P. M. Spangler, R. A. Faitoute, W. E. Steiner (NMNH); 2 2 micr., Cerro de la Neblina, same, but 1 km S of Basecamp (NMNH). Known distribution (Fig. 17).—Brazil, French Guiana, Guyana, Peru, Suriname, Venezuela. Discussion.—This long, slender species does not resemble any of the other species VOLUME 97, NUMBER 1 of the genus except S. fersa, with which it forms a species pair. These species are dis- tinguishable from each other by the com- plement of striae on the abdominal sternites as given in the key and the short dark region on the hind femur of S. tersa (Fig. 18a, c) as opposed to a longer, more diffuse, dark region in S. strigosa. They share a similar size and shape, large triangular silvery spots on the pronotum (Fig. 18a), an extensive covering of silvery setae on the base of the hemelytra of macropters, and large rastrate stridular patches on the slender hind fem- ora; the stridular patches are only modestly thickened distally (Fig. 18c). Stridulivelia strigosa 18 usually micropterous. Stridulivelia (Stridulivelia) tersa (Drake & Harris) Figs. 18, 19 Velia tersa Drake & Harris, 1941: 338. (Type from Trinidad, B.W.I.; in the National Museum of Natural History). Velia nama Drake, 1957: 114. (Type from Quarto Ojos, Rio Piray, Dept. Santa Cruz, Bolivia; in the National Museum of Nat- ural History) (New synonymy). Velia (Stridulivelia) tersa; Drake & Menke, 1962: 415. Stridulivelia tersa; Polhemus, 1976: 509. Material examined (all JTPC, unless not- ed).—BOLIVIA: Beni: 1 6 micr., rainforest stream 40 km S of Rurrenabaque, nr. Monte Redondo Sawmill, CL 2511, 12.1X.1989, J. dace: A; Polhemus: La Paz: 3yéanicr., 9 2imucr;, 3\ km) SEof Sapecho, “CE, 2515, 12.1X.1989, J. T. & D. A. Polhemus. Santa Cruz: 1 2° macr., Ichilo Prov., stream 10 km S Buena Vista, 380 m, 20.1X.1989, CL 2357, J. T. and D. A. Polhemus. BRAZIL: Ama- zonas: 1 2 macr., Reserva Ducke, 25 km NE of Manaus, Igarape da Anta, 60 m, water temp.24:5°C, 25. Vill. 1989, CL.2472. J. T. and D. A. Polhemus; 6 4 macr., 5 2 macr., Reserva Ducke, Igarape Acara, 26. VIII.1989, CL 2474, J. T. and D. A. Polhemus; 2 2 macr., Reserva Ducke, Igarape Barro Branco, nr. 147 headquarters, 50 m, 27.VIII.1989, CL 2475, J. T. and D. A. Polhemus; 3 6 macr., rain- forest stream at INPA Forest Mgmt. Sta., 98 km NW Manaus, 90 m, 29 Aug. 1989, CL 2477, J.T. and D. A. Polhemus. Matto Gros- so: 4 6 macr., 8 2 macr., Serra Roncador, 16- 17. VIII.1965, A 558, E. J. Fittkau; 9 ¢ macr., 9 2 macr., Serra Roncador, nr. Acampamento Sarape,skm 125, lg2Vilbig6s" A 559, E. J. Fittkau (JTPC, ZSMC). GUYANA: | 4 macr., 1 @ macr., Karanambo, 18.1X.1957, R. McConnell. PERU: Loreto: 7 6 macr., 5 9 macr., stream near Napo Camp, CL 2459, 15.VIII.1989, J. T. & D. A. Polhemus; 7 2 macr., 6 2 macr., stream near Explorama Lodge, CL 2457, 14. VIII.1989, J. T. & D. A. Polhemus; 3 6 macr., 4 2 macr., small rain- forest stream near Explorama Inn, CL 2465, 18.VIII.1989, J. T. & D. A. Polhemus. SU- RINAME: 1 ¢ micr., 5 6 macr., 4 2 macr., Zanderijsavanne, Carolinakreek, 22. VIII.1969, SN 092, N. Nieser; | 2 micr., same but 18. VIII. 1969, SN 080, N. Nieser; 1 2 macr., Coesewijn Project, Brokopondo, 9 km along Rd. to S. branch, 7.1V.1970, SN 402, N. Nieser. TRINIDAD: | 6 micr., 27.X.1938, C. J. Drake. VENEZUELA: Amazonas: 1 ¢ micr., 2 2 macr., 39 km S. Puerto Ayacucho, brook, coll. 4, 15.XI.1987, P. J. Spangler & R. A. Faitoute (NMNH); 30 ¢ micr., 6 2 macr., 42 2 micr., 4 2 macr., 21 km S. To- bogan Jct. (53 km S of Pto. Ayacucho), “Missionary Stream,” small river with al- ternating riffles and pools, CL 2375, 22.1.1989, J. T. Polhemus; 7 4 micr., 4 2 micr., 6 km S. Tobogan Jct., brook, CL 2374, 22.1.1989, J. T. Polhemus; 2 4 micr., 2 2 micr., Puente Pulda, 16 km S Pto. Ayacucho, CL 2387, 26.1.1989, J. T. Polhemus; 4 ¢ micr., 2 6 macr., 3 2 micr., 4 2 macr., 30 km S Pto. Ayacucho, Rio Paria Chico, CL 2372, 21.1.1989, J. T. Polhemus; 6 4 micr., 7 6 macr., 4 2 micr., 8 2 macr., stream, 0.1 km S of Tobogan-Coromoto Jct., CL 2383, 25.1.1989, J. T. Polhemus; 4 6 macr., 5 2° macr., 38 km N Pto. Ayacucho, CL 2386, 25.1.1989, J. T. Polhemus; 4 6 macr., 1 2 macr., small clear stream with sandy bottom, 0.5 km N of Alto 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON i fy \ \y us, micropterous male; b, body, Fig. 18. Stridulivelia (Stridulivelia) tersa (Drake & Harris): a, dorsal habit cles on the connexivum; c, hind lateral view, showing lateral abdominal glabrous grooves and stridulatory denti femur, showing basal rastrate stridular structure. (From Drake and Menke 1962) VOLUME 97, NUMBER 1 149 Fig. 19. Stridulivelia (Stridulivelia) tersa (Drake & Harris), known distribution. Mavaca Base Camp, 2°1'30’N, 65°7'0’”W, 228 m, water temp. 22°C, CL 8006, 4.1I.1989, D. A. Polhemus. Prov. Unknown: | é micr., 2 ° micr., Barinas, Rio Bocono, 13.1X.1957, B. Malkin. Known distribution (Fig. 19).— Bolivia, Brazil, Guyana, Peru, Suriname, Trinidad, Venezuela. Discussion.— The type of S. tersa is a mi- cropterous male. This species is quite com- mon in South America and we now have long series including both micropterous and macropterous forms; the macropterous morph is predominant. See comparative notes under S. strigosa. The macropterous type of S. nama was compared with a macropterous male of S. tersa by Dr. D. A. Polhemus who found it to be the same in every important respect. Drake (1957) stated in the description of nama: ““Abdomen beneath with first three abdominal segments deeply transversely grooved on each side,” which would be a key character distinguishing this species from tersa; however, the type of nama has these grooves on the first 5 (visible) abdom- inal segments, exactly as shown in the splen- did figure of tersa given by Drake and Menke (1962, plate 1, fig. b; our Fig. 18b), and the stridulatory mechanism also is exactly the same. We recently collected specimens of nama from several localities in Bolivia very near the type locality, but when we could not distinguish these from fersa, and they 150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 20. Stridulivelia (Stridulivelia) transversa (Hungerford), known distribution. matched the description of nama except for the abdominal grooves, we reverted to the types to establish the synonymy. Stridulivelia (Stridulivelia) transversa (Hungerford) Fig. 20 Velia (Stridulivelia) transversa Hungerford, 1929: 54. (Type from Guyane, Haut-Car- sevenne; in Paris Museum.) Stridulivelia transversa; Polhemus, 1976: 509. Material examined (all JITPC).—BRA- ZIL: Amazonas: 11 6 micr., 11 2 micr., Re- serva Ducke, 25 km NE of Manaus, Igarape da Anta, 60 m, water temp. 24.5°C, 25. VIII.1989, CL 2472, J. T. and D. A. Pol- hemus; 9 é micr., 9 2 micr., Reserva Ducke, Igarape Barro Branco, nr. headquarters, 50 m, 2/.VI11.1989. CL 2475, J. Land Ds A- Polhemus; 26 4 micr., 1 6 macr., 24 2 micr., 1 2 macr., forest stream at INPA forest man- agement station, 98 km NW Manaus, 90 m, water temp. 25°C, 29 Aug. 1989, CL 2477, J.T. & D.A. Polhemus; 2 6 micr., 4 6 macr., 4 2 micr., 2 2 macr., stream near viewing tower, 90 km NW Manaus, 90 m, 29 Aug. 1989, CL 2478, J. T. & D. A. Polhemus; 9 émicr., 7 2 micr., stream, nr. Egler Reserve, 30. VITI.1989, CL 2479, J. T. and D. A. Pol- hemus; | 6 macr., Mont Alegre, Mulaba, 23.1X.1954, $303-6, H. Sioli; 1 2 macr., Lago VOLUME 97, NUMBER 1 Salgado, Ig. (Igarape) Agua Dolce, 23.1V.1948, $225, H. Sioli; 1 ¢ micr., Rio Negro, Ig. (Igarape) Barro Branco, 30.VII.1962, A 400, E. J. Fittkau. SURI- NAME: | é micr., 1 6 macr., 1 2 micr., Zan- derijsavanne, Ist trib. of Colakreek crossing rd. to Matta, 19.IX.1969, SN 150, N. Nies- er: 2 2 macr., Ist trib of Colakreek, 8.IX.1969, SN 127, SN 128, N. Nieser; 1 6 mich... 1) 2 micr.;, Coesewijn.. Project, 3.111.1970, SN 368, N. Nieser; 3 6 micr., | 2 micr., Ist streamlet, Hanover Rd., 4.VIII.1969, SN 058, N. Nieser. Known distribution (Fig. 20).—Brazil, French Guiana, Suriname. Discussion.—This is the smallest and most delicate of the Stridulivelia species, easily distinguishable by its small size, small stridulatory patch on the hind femur, com- plement of transverse abdominal grooves, and strongly narrowed abdomen of the fe- males. ACKNOWLEDGMENTS Our special thanks go to Dr. Nico Nieser, Tiel, The Netherlands, for the generous loan and gift of specimens from his extensive collections of aquatic Heteroptera from Su- riname. We are also indebted to the late H. H. Weber, Kiel, for the gift and loan of spec- imens (collection now in Munich, ZSMC). We are indebted to the following individ- uals, without whose assistance the research could not have been completed: Dr. R. C. Froeschner, Smithsonian Institution, Washington, D.C., for access to the Drake Collection at the National Museum of Nat- ural History and arranging for the use of figures 1, 2, 4, 15, and 18 drawn by Patricia J. Hogue; Dr. Dan A. Polhemus, Bishop Museum, Honolulu, for assistance in col- lecting specimens and comparing type ma- terial at the Smithsonian Institution. For assistance in the field, J. T. Polhemus thanks the following individuals: Gabriel Roldan P., Luis Fernando Roldan, Luisa Fernanda Alvarez, Medellin, Colombia; 151 Raquel T. de M. Sampaio, Victor Py-Dan- lel, INPA, Manaus, Brazil; Peter Jensen, Angel Ocman, Iquitos, Peru; Gerardo La- mas, J. Ruben Tejada D., Lima, Peru; Ed- uardo Forno, Fernando Guerra, La Paz, Bo- livia; Paolo Betella, Santa Cruz de la Sierra, Bolivia; Robin Clarke, Guy Cox, Buena Vista, Bolivia. We thank colleagues Terry Bonace, Ronald Cave, and Adele Conover who collected and donated specimens to the Smithsonian Institution that are included in this review. The following personnel of the Smithson- ian Institution contributed to the study in various ways and we appreciate their assis- tance: Robin A. Faitoute, Oliver S. Flint, and Warren E. Steiner for assistance in col- lecting specimens; and Young T. Sohn for Figs. 6, 7, 10, and 11 and assistance with maps. The field work of J. T. Polhemus and D. A. Polhemus during which many specimens included in this study were collected was supported in part by grant #4092-89 from the National Geographic Society, Washing- ton, D.C. The field work of P. J. Spangler, during which some included specimens were collected, was supported in part by the fol- lowing: Brazil: The Consércio Nacional de Engenheiros Consultores S.A., through Dr. Paulo Vanzolini and the Smithsonian In- stitution (Neotropical Lowland Research Program, R. Heyer). Guyana: The Center for Field Research, Earthwatch Expedition: The Biological Diversity of the Guianas Program, V. Funk. Venezuela: The Foun- dation for the Development of Physics, Mathematics, and Natural Sciences of Ven- ezuela; and the Smithsonian Institution (Scholarly Research Fund; Research Op- portunities Fund). We thank Dr. John E. McPherson, Southern Illinois University; Dr. Dan A. Polhemus, Bishop Museum, Honolulu; and Dr. Robert W. Sites, University of Missou- ri, for useful, critical reviews of the manu- script. 152 LITERATURE CITED Andersen, N. M. 1982. The semiaquatic bugs (He- miptera, Gerromorpha), phylogeny, adaptations, biogeography, and classification. Entomonograph 3: 1-455. Champion, G. C. 1898. Insecta, Rhynchota, Hemip- tera-Heteroptera. Biologia Centrali-Americana 2: 1-416, pls. 1-22. Drake, C. J. 1957. New Neotropical water-striders (Hemiptera). Proceedings of the Biological Society of Washington 70: 111-118. Drake, C. J. and H. M. Harris. 1938. y Sudamericanos descriptos por Carlos Berg. Notas del Museo de La Plata. Zool- ogia. 3(13): 199-204. 1941. A new Velia from Trinidad (Hemip- tera). Revista de Entomologia 12: 338-339. Drake, C. J. and A. S. Menke. 1962. Water striders of the subgenus Stridulivelia from Mexico, Central America, and the West Indies (Hemiptera: Veli- idae). Proceedings of the United States National Museum 113(3460): 413-419. Hungerford, H. B. 1929. Some new semi-aquatic He- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON miptera from South America with a record of stridulatory devices. (Veliidae-Velia). Journal of the Kansas Entomological Society 2: 50-59. Latreille, P. A. 1804. Histoire naturelle, Genérale et Particuliere des Crustacés et des Insectes 12: 269. Polhemus, D. A. In press. Systematics, phylogeny and zoogeography of the genus Rhagovelia (Het- eroptera: Veliidae) in the western hemisphere (ex- clusive of the angustipes complex). Thomas Say Monographs, Entomological Society of America. Polhemus, J.T. 1976. A reconsideration of the status of the genus Paravelia Breddin, with notes and a check-list of species (Veliidae: Hemiptera). Jour- nal of the Kansas Entomological Society 49: 509- ails), 1979. A new species of Stridulivelia from Mexico, and a new subgenus from Middle Amer- ica (Hemiptera: Veliidae). Pan-Pacific Entomol- ogist 55: 46-50. Polhemus, J. T. and D. A. Polhemus. 1985. Studies on Neotropical Veliidae (Hemiptera) VIII: New species and notes. Pan-Pacific Entomologist 61: 163-169. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 153-160 A NEW SPECIES OF CENTISTES FROM BRAZIL (HYMENOPTERA: BRACONIDAE: EUPHORINAE) PARASITIZING ADULTS OF DIABROTICA (COLEOPTERA: CHRYSOMELIDAE), WITH A KEY TO NEW WORLD SPECIES Scott RICHARD SHAW Department of Plant, Soil, and Insect Sciences, University of Wyoming, P.O. Box 3354, Laramie, Wyoming 82071. Abstract. — A new species of euphorine braconid, Centistes gasseni Shaw from Brazil is described, diagnosed, and illustrated. This species is a koinobiont endoparasitoid of the adult stage of Diabrotica, and is currently being studied as a possible biocontrol agent for the Diabrotica pest-complex including the southern corn rootworm in North America. An identification key for the known Centistes species in the New World region is included. Three species, C. agilis (Cresson), C. claripennis (Ashmead), and C. /aevis (Cresson) are newly recorded as occurring in Wyoming, representing new far western distribution records for these species. Key Words: The purpose of this paper is to describe a new species of Centistes parasitizing Di- abrotica that was recently discovered in Passo Fundo, Rio Grande do Sul, Brazil. The new Centistes was reared from Dia- brotica speciosa (Germar) in 1992 by Dirceu N. Gassen, of EMBRAPA-CNPT, Passo Fundo, working in collaboration with Rob- ert F. W. Schroder, USDA-ARS, Insect Bio- control Laboratory, Beltsville, Maryland. It is currently being studied by Dr. Schroder as a possible biocontrol agent for the Dia- brotica pest-complex including the southern corn rootworm, Diabrotica undecimpunc- tata howardi Barber. D. undecimpunctata 1s a serious pest of corn and cucurbits in the southern United States, and Diabrotica spe- closa is increasingly becoming a problem in Brazil. The new Centistes was imported to the United States in 1992 and has been suc- cessfully raised from D. undecimpunctata at the Maryland Department of Agriculture quarantine facility in Annapolis. A permit Braconidae, Euphorinae, Centistini, neotropical, biological control was recently obtained for laboratory studies at the Beltsville Agriculture Research Cen- ter. Consequently, providing a scientific name for this species is critical at this point in time. The genus Centistes is the most diversi- fied lineage of the euphorine tribe Centistin1 (Shaw 1985). The Centistini can be most easily diagnosed by the broad first meta- somal tergum with spiracles very near the front of the segment (Fig. 13) and the very smooth fused tergum 2 + 3 without a lateral fold or crease (Fig. 12). Centistes are the most common members of the centistine lineage with an extremely short, broad, and densely setose ovipositor sheath (Figs. 15, 16). A less obvious, but phylogenetically significant, character is the extreme reduc- tion or complete absence of vein M beyond the Rs in Centistes. The genus 1s essentially worldwide in distribution, except for the Australian continent. Most of the described species are holarctic in distribution, but that 154 is probably only an artifact of poor sampling and lack of taxonomic revisions in the trop- ics. For example, while there is a total of six named Centistes species in North Amer- ica (Shaw 1985), there are at least eight un- described Centistes species occurring in Costa Rica alone (Shaw unpublished data). While no Centistes species have been de- scribed from South America, one closely related species has recently been named in the genus Centistoides (van Achterberg 1992). Centistes can be identified to genus using the keys of Shaw (1985), van Achter- berg (1985, 1992), or Marsh et al. (1987). The tribe Centistini is defined by Shaw (1985) and van Achterberg (1985, 1992). The subfamily Euphorinae is defined by Shaw (1985) and can be identified using the keys of Shaw and Huddleston (1991) or Goulet and Huber (1993). Descriptive terminology follows that of Shaw (1985, 1993) and Marsh et al. (1987). Terminology for microsculpture follows that of Harris (1979). Wing venation terminol- ogy is that of Goulet and Huber (1993). Scanning electron microscopy was done with a JEOL model 35CF SEM at an operating voltage of 25 kv. Specimens from this study are deposited at the University of Wyo- ming, Rocky Mountain Systematic Ento- mology Laboratory, Laramie (RMSEL), the United States National Museum of Natural History, Washington, D.C. (USNM), the Universidade Federal de Sao Carlos, Sao Paulo, Brazil (UFSC), and EMBRAPA- CNPT, Passo Fundo, Rio Grande do Sul, Brazil. Centistes gasseni Shaw, NEw SPECIES Figs. 1-18 Holotype female.— Body length 2.8 mm; forewing length 3.5 mm. Head (Figs. 1, 2).—Shortest distance be- tween eyes 1.28 x greater than clypeus width; face, clypeus, and gena evenly setose; frons, vertex, and temple smooth and devoid of setae; temple width 0.5 x eye width; eyes, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in anterior view, parallel and not converging ventrally; eye 1.61 x taller than wide, smooth, devoid of minute setae; scape cy- lindrical, 1.36 longer than wide; pedicel compact, 0.88 x longer than wide; antenna with 26 flagellomeres; F1 3.13 x longer than wide; F10 2.86 x longer than wide; F26 2 x longer than wide basally, terminating api- cally in a sharp point; malar space 0.22 x eye height, about equal to basal width of mandible; mandibles when closed overlap- ping for 0.6 mandible length; mandible width basally 0.25x mandible length, strongly narrowing apically; ocellar triangle equilateral, posterior margin of median ocellus slightly ahead of anterior margin of lateral ocellus; ocelli large, lateral ocellus separated from compound eye by distance 0.83 x ocellar width; occipital carina strong and complete, but situated very low on oc- ciput, apex of occipital carina about level with midpoint of eye. Mesosoma (Figs. 3—11).— With scattered large setae abouth same size as those on head posteriorly, except pronotum laterally, mesonotum, scutellum, central disc of mesopleuron, and dorsum of porpodeum largely devoid of setae; pronotum rugose anteriorly, with a series of 5 shallow pron- opes dorsomedially (Fig. 3); pronotal fur- row foveate medially, smooth ventrally; prosterna with anterior and medial margins rugose, otherwise smooth, ending as smooth rounded flange over base of fore coxa (Fig. 4); mesonotum smooth, without any trace of notauli; mesopleuron mostly smooth, but with oblique foveate sternaulus and foveate posterior margin (Fig. 5); scutellar furrow bifoveate (Fig. 7); scutellar disc smooth; lat- eral subapical scutellar margins rugofov- eolate; apex of scutellum minutely bifov- eate; metanotum irregularly longitudinally costate, posterior rim narrow and smooth (Fig. 8); mesosternum anteriorly densely covered with erect setae; mesosternum and metasternum posteriorly obliquely costate (Fig. 11); propodeum clearly subdivided into anterior and posterior halves by a strong VOLUME 97, NUMBER 1 Figs. 1-6. C. gasseni. 1, Lateral view of head. 2, Anterior view of head. 3, Dorsal view of pronotum and anterior mesonotum. 4, Ventral view of pronotum, prosterna, and anterior coxal cavities. 5, Lateral view of mesosoma. 6, Dorsal view of anterior margin of hindwing and hamulli. transverse carina, surfaces mostly smooth et al. (1987) except pterostigma 3.53 x lon- except medially and laterally rugulose (Figs. ger than maximum width; r-rs vein 0.66 x 9; 10). as long as maximum width of pterostigma; Wings. — Venation asin Fig. 306 ofMarsh length of marignal cell 2R1 along anterior 156 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-12. propodeum. 10, Lateral view of propodeum. 11, Ventral view of posterior mesosternum, metasternum, and associated coxal cavities. 12, Lateral view of metasoma. wing margin equal to length of pterostigma; vein m-cu interstitial with vein Rs; vein M apically represented by a very short scler- otized stub; hindwing with 3 sinuate ha- mules (Fig. 12). Legs.— Moderately setose throughout; surfaces smooth and shining, especially of coxae and femorae; coxae short and com- pact, fore and middle coxal lengths equal to C. gasseni. 7, Dorsal view of scutellum. 8, Dorsal view of metanotum. 9, Dorsal view of trochanter length; hind coxa longer, 1.8 x middle coxa length; hind femora length 5.4 x longer than maximum width; hind tibia length 9x longer than maximum width, apical 30% moderately compressed in dor- sal view; hind tibial spurs moderately long, 0.48 x as long as hind basitarsus; ratio of hind tarsomeres from basitarsus apically 48: 26:22:16:24; tarsal claws large, simple. VOLUME 97, NUMBER 1 Figs. 13-18. sternum. 15, Ventral view of apex of metasoma showing hypopygium and ovipositor sheaths with ovipositor retracted. 16, Lateral view of apex of metasoma showing hypopygium and ovipositor sheaths with ovipositor extended. 17, Lateral view of ovipositor apex. 18, Lateral view of ovipositor sheath apex. Metasoma (Figs. 12—18).—Tergum | ba- sally 0.63 x as wide as apical width, 1.7 x longer than apical width; spiracle of tergum 1 situated on lateral margin near basal quar- ter of segment; sculpture of tergum 1 lon- gitudinally costate laterally, smooth medi- ally and apically (Fig. 13); sternum 1 costate anteromedially, smooth posteriorly (Fig. 14); remainder of metasoma smooth and shin- 157 C. gasseni. 13, Dorsolateral view of first metasomal tergum. 14, Ventral view of first metasomal ing (Fig. 12); hypopygium deeply incised and laterally situated (Figs. 15, 16); ovi- positor sheath short and broad, 3.2 x longer than basal width, densely setose (Figs. 15, 16), terminating in a sharp point formed from a fringe of dense setae (Fig. 18); dorsal and ventral edges of right ovipositor sheath wrapping over edges of left sheath; ovipos- itor smooth and saber-like, dorsal valve with 158 a deep subapical notch, ventral valve with 3 subapical curved incisions (Fig. 17). Color.— Body bright orangish yellow ex- cept flagellum, compound eyes, ocelli, tips of tarsi, and ovipositor sheaths black; ped- icel, mesonotum anteriorly and laterally, wing venation, apical 4 of hind tibia, and remainder of hind tarsus dark brown; wing membrane infumated with light brown. Variation.— Paratype females. Essential- ly as in holotype except body length 2.6-3.7 mm; forewing length 3.0-3.5 mm; 25-26 flagellomeres; color of compound eye vary- ing from black to light gray; color of ocelli varying from black to light reddish brown; ocellar triangle varying from orange to dark brown or black; anterior and lateral meso- notal lobes varying from brown to black; Ovipositor position varying from concealed to fully extended (as Fig. 12). Paratype males.—As female except body length 2.6-3.5 mm; forewing length 2.5-3.0 mm; 24 flagellomeres; apex of metasoma from T3 beyond often infused with dark brown; parameres ovate with ventral mar- gin somewhat flattened, orangish yellow, apically fringed with setae; aedeagus bilo- bate, smooth, yellowish white. Host.— Originally reared from Diabrotica speciosa in Brazil. Reared in quarantine from the southern corn rootworm, Diabrotica un- decimpunctata howardi. Biology.—As far as known, all Centistes species are solitary koinobiont endoparas- itoids of adult beetles, particularly the fam- ilies Curculionidae, Chrysomelidae, Coc- cinellidae, Anthicidae, and Carabidae (Loan 1972, Shaw 1985). Only one other species, Centistes (Syrrhizus) diabroticae (Gahan), is recorded as parasitizing adult stages of the subtribe Diabroticina (Gahan 1922). That species attacks the striped cucumber beetle, Acalymma vittatum (F.). The only other named neotropical species, Centistes epi- caer! Muesebeck, parasitizes a curculionid on coffee (Muesebeck 1958). Some Centistes have been reported to leap onto the host elytra (Gahan 1922) or grasp the host beetle PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with their legs during oviposition (Loan 1964, 1972). The dense layer of mesosternal setae in Centistes gasseni may be an adap- tation to similar host-mounting behavior. More complete reviews of euphorine biol- ogy are given by Shaw (1988, 1994) and Shaw and Huddleston (1991). Discussion. — Centistes gasseni is a very distinctive species because of its bright or- angish yellow coloration, and should not be confused with any of the other known Cen- tistes species. The only other described neo- tropical species, C. epicaeri, has an entirely black body, more coarsely costate tergum 1, and has the posterior face of the propo- deum subdivided by a median carina that is absent in C. gasseni. The other unde- scribed Centistes species that I’ve seen from the neotropics all have mostly black body coloration, and are mostly smaller in size. The other species that parasitizes chryso- melids, C. diabroticae varies in body color from black to orangish yellow, but 1s some- what smaller (2.2 mm), has a distinctive median fovea on the mesopleuron that is absent in C. gasseni, and has most of the Rs+M vein absent basally, thus the 1R1 and IM cells are confluent (these cells are completely separated by the Rs+M vein in C. gasseni). C. gasseni can be distinguished from other known Centistes in the New World by the key given below. This key treats all of the named New World Centistes as reclassified by Shaw (1985). Material examined.—Holotype: female, Brazil, Rio Grande do Sul, Passo Fundo, July 1991, D. N. Gassen, reared from Di- abrotica speciosa, (RMSEL). Paratypes: 2 females, 2 males, same data as holotype; 30 females, 22 males, lab reared, United States, Maryland, Annapolis, Maryland Depart- ment of Agriculture quarantine facility, em. May 1, 1993, R. Schroder, reared from Di- abrotica undecimpunctata howardi; 11 fe- males, 149 males, same data except em. June-July 1993, (RMSEL, USNM, UFSC, EMBRAPA-CNPT). Etymology.— Named for the collector of VOLUME 97, NUMBER 1 the type series, Mr. Dirceu N. Gassen, of EMBRAPA-CNPT, Passo Fundo, Rio Grande do Sul, Brazil. KEY TO THE NEw WORLD SPECIES OF CENTISTES HALIDAY 1. | Forewing vein Rs+M greatly reduced or ab- sent, thus cells 1R1 and 1M forming a single large confluent cell; mesonotum smooth and without any trace of notauli (subgenus Syr- TALZUS) eee ey eT Mess 2 - Forewing vein Rs+M complete, thus cells 1R1 and 1M completely separated; meso- notum variable, but notauli often indicated, as least anteriorly, by a groove or foveation 2(1). Forewing vein Rs+M entirely absent; meso- notum entirely smooth, without any trace of sculpture; mesopleuron smooth, without any trace of a sternaulus; body black; parasitoids of Anthicidae (Notoxus); New York south to Virginia, westwards to Wyoming ...... Se ay Ma A ae BO ee Centistes agilus (Cresson) - Forewing vein Rs+M partly present apical- ly, at least with a distinct stub; mesonotum with a distinctive median fovea, but other- wise smooth; mesopleuron with a small but distinct foveate sternaulus, but otherwise smooth; body color variable, but often ex- tensively marked with reddish to yellowish brown; parasitoids of Chrysomelidae (Aca- lymma), central United States (Illinois, Ohio, Missouri) ...... Centistes diabroticae (Gahan) 3(1). Notauli foveate and distinct throughout the length of the mesonotum; metasoma some- times with small tooth-like structures situ- ated on sternum 5 (subgenus Ancylocentrus) - Notauli distinct only anteriorly, indicated by smooth grooves, or absent; metasoma smooth, without small tooth-like structures situated on sternum 5 (subgenus Centistes) 4(3). Small species 3 mm or less in body size; propodeum areolate posteriorly; metasoma with small tooth-like structures situated on sternum 5; parasitoids of Curculionidae (Si- tona); holarctic (northeastern Canada and EUrOpe) eerre le eer oso ee ete Oe Centistes ater (Nees) [= C. excrucians Haliday] = Robust species 5 mm or greater in body size; propodeum coarsely rugose; metasoma without small tooth-like structures situated 159 on sternum 5; parasitoids of Carabidae (Amara); eastern North America from Que- bec south to Pennsylvania, westwards to South Dakota and Wyoming ............ Le See EE Pas) Centistes laevis (Cresson) 5(3). Body color mostly bright orangish yellow; ovipositor sheath terminating in a sharp point (Fig. 18); parasitoids of Chrysomelidae (Di- abrotica); native to Brazil, introduced to the Wnited’Statesies a eee eer ee ete eats Centistes gasseni Shaw, NEW SPECIES - Not as above; body color mostly black; ovi- positor sheath blunt or rounded, not ter- minatingiinia sharp pomt .2---224--..--- 6 6(5). Mesonotum with notauli indicated by fine grooves; posterior face of the propodeum subdivided by a median carina; tergum | coarsely costate and nearly parallel-sided, only slightly wider posteriorly than basally; moderately large species 3-4 mm in body length; parasitoids of Curculionidae (Epi- caerus) on coffee; El Salvador eee ee Centistes epicaeri Muesebeck - Mesonotum smooth, notauli absent, poste- rior face of the propodeum not subdivided by a median carina; tergum | not so coarsely costate and about 1.5 wider posteriorly than basally; very small species 2 mm or less in body size; hosts unknown; central United States from Michigan south to Arkansas, Washington, D.C. westwards to Wyoming Centistes claripennis (Ashmead) [= C. politus (Ashmead)] ACKNOWLEDGMENTS Specimens of the new species and finan- cial assistance for publication were provid- ed through the courtesy of Dr. Robert F. W. Schroder, USDA-ARS, Insect Biocontrol Laboratory, Beltsville, Maryland. Speci- mens of other species for comparison were borrowed from the U.S. National Museum of Natural History (Dr. P. Marsh & T. Nuhn). Financial support for scanning elec- tron microscopy was provided by National Science Foundation grant DEB 930-0517. Dr. Susan Swapp and Mr. Keith Krugh, of the Department of Geology and Geophys- ics, University of Wyoming, provided crit- ical assistance with scanning electron mi- croscopy and photography. Special thanks to Ms. Nina Zitani for her assistance with photographic development and printing. 160 LITERATURE CITED Achterberg, C. van. 1985. The genera and subgenera of Centistini, with description of two new taxa from the Nearctic region (Hymenoptera: Bracon- idae: Euphorinae). Zoologishche Mededelingen Leiden 59: 348-362. 1992. Centistoides gen. nov. (Hymenoptera: Braconidae: Euphorinae) from Suriname. Zoolo- gishche Mededelingen Leiden 66: 345-348. Gahan, A. B. 1922. A new hymenopterous parasite upon adult beetles. The Ohio Journal of Science 22: 140-142. Goulet, H. and J. T. Huber. 1993. Hymenoptera of the world: An identification guide to the families. Research Branch Agriculture Canada Publication 1894/E: 1-668. Harris, R. A. 1979. A glossary of surface sculpturing. Occasional papers of the Bureau of Entomology of the California Department of Agriculture No. 28: 1-31. Loan, C. C. 1964. Observations on the biology of Centistes excrucians Haliday (Hymenoptera: Bra- conidae). Prodeedings of the Entomological So- ciety of Ontario 94: 56-61. 1972. Parasitism of adult Notoxus anchora Henz. (Coleoptera: Anthicidae) by Syrrhizus agi- lus (Cresson) (Hymenoptera: Braconidae). Pro- ceedings of the Entomological Society of Ontario 103: 76. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Marsh, P. M., S. R. Shaw, and R. A. Wharton. 1987. An identification manual for the North American genera of the family Braconidae (Hymenoptera). Memoirs of the Entomological Society of Wash- ington 13: 1-98. Muesebeck, C. F. W. 1958. New neotropical wasps of the family Braconidae (Hymenoptera) in the U.S. National Museum. Proceedings of the United States National Museum 107: 405-420. Shaw, M. R. and T. Huddleston. 1991. Classification and biology of braconid wasps. Royal Entomo- logical Society of London, Handbooks for the Identification of British Insects 7(11): 1-126. Shaw, S. R. 1985. A phylogenetic study of the sub- families Meteorinae and Euphorinae. Entomog- raphy 3: 277-370. 1988. Euphorine phylogeny: The evolution of diversity in host-utilization by parasitoid wasps (Hymenoptera: Braconidae). Ecological Entomol- ogy 13: 323-335. 1993. Three new Microctonus species indig- enous to New Zealand (Hymenoptera: Braconi- dae). The New Zealand Entomologist 16: 29-39. 1994. The Braconidae. Jn Hanson, P. and Gauld, I. eds., The Hymenoptera of Costa Rica. Oxford University Press. (In press.) PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 161-177 NEW WESTERN NEARCTIC SWELTSA (PLECOPTERA: CHLOROPERLIDAE) REBECCA F. SuRpDICK, PH.D. Entomological Laboratory, Route 2, Box 1072, Front Royal, Virginia 22630. Abstract. —New species Sweltsa resima, Sweltsa cristata, Sweltsa umbonata and Sweltsa adamantea are described and compared with their closest relatives. Key Words: The following four new species of the stonefly genus Swe/tsa Ricker (Plecoptera: Chloroperlidae) were discovered during ex- amination of numerous collections of the genus from throughout western North America. Information on similarities and differences between each new species and the rest of its group accompanies descrip- tions in advance of publication of further revisionary studies. Sweltsa, found in the eastern Palearctic and the Nearctic, is one of three genera in tribe Alloperlini of subfamily Chloroperli- nae. They are small- to medium-sized, yel- lowish stoneflies with a dark abdominal stripe and distinct dark marks on head and nota. The genus is particularly recognized by the recurved epiproct that is composed of the basal bar, widened anteriorly into a cupped anchor, and the hinged, sculptured epiproct tip, extending anteriorly to a trans- verse ridge on deeply incurved tergum nine (Surdick 1985). The subgenital plate of the female is slightly convex and usually basi- cally hexagonal. Surdick (1985) gives a key to Nearctic chloroperlinid genera and a list including western Nearctic Swe/tsa species. Bau- mann, Gaufin, and Surdick (1977) give a key to most Rocky Mountain and Pacific Northwest Swe/tsa and Jewett (1960) gives one for most California Sweltsa. Plecoptera, Chloroperlidae, Sweltsa, Nearctic Sweltsa resima, sp. nov. Figs. 1-6 Adult. — General color darkly-marked tan in alcohol. Head with 3 dark ocellar rings, dark rugulae in dusky environs anterior to and adjacent to ocellar triangle and poste- rior to compound eyes. Pronotum with broad dark explanate margin, dark longi- tudinal rugulae, median longitudinal un- marked area one-fourth width of pronotum. Meso- and metanota dusky, each with dark recurrent scutoscutellar suture. Abdomen with dark median longitudinal stripe from tergum | to 8, single dark lateral stripes from segment | to 5, lateral pectens on segments 7, 8, 9. Macropterous; wings hyaline, dark- veined, representative of genus. Male.—Body length 8 mm; forewing length 9 mm. Epiproct sclerotized, hinged, with tip extending anteriorly to prominent sclerotized deeply crenate transverse ridge on anterior of tergum 9. In dorsal aspect, epiproct tip lozenged, twice as long as wide; verge curved ventrad; median longitudinal furrow ending at recurved apex. In lateral aspect of epiproct tip, distal half of dorsal face declivent 45 degrees; apex recurved, conical; verso of furrow visible beneath verge. Basal anchor of epiproct broadly and deeply scoop-like, notched anteriorly; epi- proct support structures otherwise repre- 162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-9. Sweltsa resima. 1, Male terminalia, dorsal aspect. 2, Epiproct tip, dorsolateral aspect. 3, Epiproct tip, lateral aspect. 4, Subgenital plate of female, lateral aspect. 5, Subgenital plate, ventral aspect. 6, Adult head and pronotum. Sweltsa townesi, epiproct tip. 7, Dorsal aspect. 8, Dorsolateral aspect. 9, Lateral aspect. VOLUME 97, NUMBER 1 sentative of genus as are segment 9, hemi- terga of segment 10. Aedeagus membranous. Female.—Body length 9 mm; forewing length 9 mm. Subgenital plate dusky, entire, crudely hexagonal, as long as wide, based centrally on sternite 8, evenly hirsute with longer hairs posteromedially, distinguisha- ble anteriorly by narrow membranous pe- riphery; posterior half flap-like, extending over most of sternite 9; profile planate but distinctly elevated. Vagina membranous. Material.—Holotype 4, allotype 2 (U.S. National Museum): California, Inyo Co., Whitney Portal 15-VI-1966 D. C. Rentz. Paratypes: 1 4, 1 2(R. F. Surdick) California, Mono Co., Lee Vining Creek, Tioga Pass 15-VII-1979 R. W. Surdick; 2 ¢ (Brigham Young Univ.) California, Inyo Co., Coyote Crk. 11 mi. SSW of Bishop, 1000 ft. 4-VII- 1986 D. Giuliani; 2 4, 2 2 (Brigham Young Univ.) California, Mono Co., Glass Crk. Meadows, Sierra Nevada, 9000 ft. 20-VII- 1990 D. Giuliani; 1 4, 1 2 (Brigham Young Univ.) same locality 28-VI-1990 D. Giuli- ani; 1 6(Brigham Young Univ.) California, Mono Co., White Mountains, Lone Tree Crk., 6400 ft. 23-VII-1985 D. Giuliani. Other material: 2 3, 2 2 (S. G. Jewett, Jr.) California, Inyo Co., Lone Pine 28-VII-1940 L. J. Lipovsky, R. H. Beamer. Compared material.—Sweltsa townesi: holotype 4, allotype 2 (Illinois Natural His- tory Survey); specimens from El Dorado, Nevada, Placer, Plumas, Sierra, Tehama, Tuolumne Cos., California (Brigham Young Univ., Field Museum of Natural History, AOR: Gaufine RYE. Hill, S: G. Jewett, Jr., W. E. Ricker, Royal Ontario Museum, B. P. Stark, R. F. Surdick, Univ. of California at Riverside, Univ. of Kansas, U.S. Na- tional Museum). Etymology.— The adjective resima means turned up and bent back and describes the tip of the epiproct. Diagnosis. — This species is closely relat- ed to Sweltsa townesi (Ricker) 1952 (Figs. 7-9). Cursorily, the dorsal aspects of the epiproct tips of the species look the same. 163 In lateral aspect, the differences are evident: the epiproct tip of S. townesi resembles a partially ballooned version of the epiproct tip of S. resima without the conical, up- turned apex. The subgenital plates of the species are similar. The pronota both bear a distinctive broad dark marginal ring. Sweltsa resima has been found on the Great Basin side of the Sierra Nevada south of the Mono Lake area. Swel/tsa townesi has been found in the northern Sierra Nevada. Sweltsa cristata, sp. nov. Figs. 21-24, 40, 42-44 Adult.— General color dusky-marked light tan in alcohol. Head with 3 dark ocellar rings, small dusky rugulae anterior to and adjacent to ocellar triangle and posterior to compound eyes. Pronotum with thin dark encircling line, thin dusky longitudinal ru- gulae. Meso- and metanota each with dark recurrent scutoscutellar suture, dusky areas. Abdomen with dark median longitudinal stripe from tergum | to 8, single dusky lat- eral stripes from segment | to 3, lateral pec- tens on segments 7, 8, 9. Macropterous; wings hyaline, dusky-veined, representative of genus. Male.—Body length 7 mm; forewing length 7.5 mm. Epiproct sclerotized, hinged, with tip deeply sculptured, crudely pris- moid, extending anteriorly to deeply cre- nate transverse ridge on anterior of tergum 9. In dorsal aspect of epiproct tip, proximal three-fourths a slightly undulating horizon- tal elongately pandurate plane constricted to half width basad of median, obtuse-an- gulate distally, with greatest width half length; middle portion a barely visible pro- jection of bulk, blunt distally, as long as wide; distal fourth filamentous. In lateral aspect, bulk of epiproct tip cuneate, ap- pearing to intersect three-fourths of slightly declivent horizontal plane forming an arced dorsal crest one-fourth height of epiproct tip; slightly excurved anterior profile a con- tinuation of crest; anteroventral filamen- tous projection slightly up-curved, colorless 164 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OG 20 ( d 19 ws 24 Figs. 10-24. Sweltsa gaufini, epiproct tip. 10, Dorsal aspect. 11, Dorsolateral aspect. 12, Lateral aspect. 13, Frontal aspect. Sweltsa albertensis, epiproct tip, showing individual variation. 14, Dorsal aspect. 15, Dorsolateral aspect. 16, Lateral aspect. 17, Dorsal aspect. 18, Dorsolateral aspect. 19, Lateral aspect. 20, Frontal aspect. Sweltsa cristata, epiproct tip. 21, Dorsal aspect and center of transverse ridge of tergum 9. 22, Dorsolateral aspect. 23, Lateral aspect. 24, Frontal aspect. VOLUME 97, NUMBER 1 beyond base. Bulk of epiproct tip concavo- concave; elliptical ventral surface nearly congruent to and aligned with horizontal plane, convex proximally, explanate later- ally. Epiproct support structures, segment 9, hemiterga of segment 10 representative of genus. Aedaegus membranous with light- ly sclerotized striate crudely square lamella bilobate apically, incurved to half length ba- sally, recurved laterally. Female.—Body length 8 mm; forewing length 8 mm. Subgenital plate dusky, crude- ly hexagonal, based centrally on sternite 8, distinguishable anteriorly by narrow mem- branous periphery; posterior half entire, flap- like, extending over most of sternite 9; mid- dle third from base to apex convex, more hirsute than remainder; profile elevated, a flattened curve. Vagina membranous. Material.—Holotype ¢, allotype ¢ (U.S. National Museum), paratype | 4(R. F. Sur- dick): Utah, San Juan Co., Johnson Crk., Tunnel 19 mi. N. of Blanding [no date] Mu- liak. Compared material.—Sweltsa gaufini: holotype 4, allotype 2° (U.S. National Mu- seum); hundreds of specimens from Idaho, Utah. Sweltsa albertensis: topoparatype ° (Illinois Natural History Survey); hundreds of specimens from Alberta, British Colum- bia, Idaho, Montana, Oregon, Wyoming. Sweltsa lamba: topoparatype 2 4, 2 (Illinois Natural History Survey, U.S. National Mu- seum); hundreds of specimens from Colo- rado, Idaho, Oregon, Utah, Wyoming. Sweltsa hondo: topoparatype 6, 2 (R. F. Sur- dick); 2 2 from New Mexico. (For above species, Brigham Young Univ., California Academy of Science, M. Cather, D. Dun- ster, Field Museum of Natural History, A. R. Gaufin, Illinois Natural History Survey, S. G. Jewett, Jr., Kansas State Univ., Lyman Museum, Montshire Museum of Science, Oregon State Univ., W. E. Ricker, Royal Ontario Museum, B. P. Stark, R. F. Surdick, Univ. of Kansas, Univ. of Minnesota, U.S. National Museum, Utah State Univ., Washington State Univ.) 165 Etymology.—The adjective cristata de- scribes the crest on the epiproct tip. Diagnosis.—Sweltsa cristata is most closely related to Sweltsa lamba (Needham and Claassen 1925) (Figs. 25-34, 41, 45- 47), Sweltsa hondo Baumann and Jacobi 1984 (Figs. 35-38), Sweltsa albertensis (Needham and Claassen) 1925 (Figs. 14-20, 39) and Sweltsa gaufini Baumann 1973 (Figs. 10-13). All five species in the Sweltsa lamba group have a sclerotized leaflet on the ae- deagus that is not found in other Sweltsa species. Their elaborately sculptured epi- proct tips and their nearly identical female subgenital plates are variations on basic forms that are unique to the group even though similar in some ways to other Swelt- sa relatives. The epiproct tips are basically concavo-concave rhombohedrons. The up- per surface is a curved plane bisected by a longitudinal carina. The sides curve to meet anteriorly at a right or acute angle bearing a swelling or projection. And, a partially colorless filament extends anteriorly from the ventral surface. In S. gaufini, found only in the Bear River area of the northern Wasatch Mountains of Idaho and Utah, the filamentous part of the epiproct tip is a short, slanted blade. The epiproct tip is more elongate than in the other species; the carina is a prominent dou- ble crest; the anterior angle bears an oblique hastate projection that is triangular in fron- tal aspect; the sides are deeply concave, and the ventral surface is half as wide as the dorsal plane. The leaflet on the aedeagus is longer than wide, lightly sclerotized and has rectangular lobes. In S. albertensis, found in the Rocky Mountains of British Columbia, Alberta, Montana, Idaho and northwestern Wyo- ming, the filament is a long, slanted blade. The dorsal plane is concave and only slight- ly flanged; the carina is minor, either not reaching all the way to the anterior angle or continuing over the anterior angle, and there is no swelling on the anterior angle but the anterior profile is undercut. The sides are 166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON < 2% : 3g 1 35 Sha a7 Figs. 25-38. Sweltsa lamba from Wasatch Range, Virgin River drainage of Utah. 25, Male terminalia, dorsal aspect. 26, Epiproct tip, dorsal aspect. 27, Epiproct tip, dorsolateral aspect. 28, Epiproct tip, lateral aspect. 29, Epiproct tip, frontal aspect. 30, Epiproct tip, frontal aspect. Sweltsa lamba from Colorado, epiproct tip. 31, Dorsal aspect. 32, Dorsolateral aspect. 33, Lateral aspect. 34, Frontal aspect. Sweltsa hondo, epiproct tip. 35, Dorsal aspect. 36, Dorsolateral aspect. 37, Lateral aspect. 38, Frontal aspect. VOLUME 97, NUMBER 1 shallowly concave and the ventral surface is nearly as wide as the dorsal plane and more convex than in the other species. The leaflet on the aedeagus is longer than wide but it is darkly sclerotized and each lobe is rounded. Sweltsa cristata, S. lamba and S. hondo are more similar to each other than to either S. gaufini or S. albertensis. The dorsal plane of each epiproct tip is undulating and broad- ly pandurate; the carina is a single crest; the slight to prominent projection on the an- terior angle is lenticular or diamond-shaped in frontal aspect; the filament is a curled extension on a blade-like base, and the leaf- let on the aedeagus is about as long as wide. The middle of the transverse ridge on ter- gum nine looks like an acutely emarginate triangle in S. gaufini and S. albertensis. It is crenate in the other four species. Characters of the epiproct tip and aede- agus distinctly separate S. cristata from S. lamba and S. hondo and indicate that the latter species are more closely related to each other than to S. cristata. The anterior part of the pandurate plane of the epiproct tip 1s nearly elliptical in S. cristata; slightly more pyriform and acuminate in S. Jamba, and nearly triangular in S. hondo. The projec- tion at the anterior angle of the bulk of the epiproct tip is minimal in S. cristata and visible as a slight swelling in frontal aspect. It is about one-fifth the length of the epi- proct tip in the other two species. The ven- tral surface of the epiproct tip is nearly as wide as and aligned with the dorsal plane in S. cristata, but itis more anteriorly placed and narrower than the dorsal plane in the other two species. The leaflet on the aedeagus of S. cristata is slightly longer than in the other two spe- cies and the broader notch in its distal edge bisects it into two rounded lobes. The nar- rower notch in the leaflets of S. Jamba and S. hondo has only slightly rounded shoul- ders and bisects a sinuate edge. Sweltsa cristata has been found in south- eastern Utah near Blanding. Sweltsa lamba 167 isa Rocky Mountain species or, more likely, a complex of two incipient or perhaps true species that exhibit considerable variety within and between populations, particu- larly in the shape and size of the anterior projection of the bulk of the epiproct tip. Utah populations found in the arc of high- lands stretching from the Wasatch Moun- tains to the Virgin River drainage have an anterior projection that is less than half the width of the dorsal plane. In those northern populations the projection is parallel-sided in dorsal aspect and lenticular in frontal as- pect (Figs. 25, 29) and in those populations from the Virgin River drainage, it 1s en- larged anteriorly and lozenged in frontal as- pect (Figs. 26-28, 30). Populations from Or- egon to Colorado have a projection that is about half to greater than half as wide as the dorsal plane. Again, in those northern populations, the projection is parallel-sided and in Colorado populations, it is enlarged anteriorly (Figs. 31-34). Sweltsa hondo has been found in the Sangre de Cristo Range of northcentral New Mexico and differs lit- tle from some southern Colorado popula- tions of S. lamba. KEY TO SWELTSA MALES WITH A SCLEROTIZED LAMELLA ON THE AEDEAGUS 1. Epiproct tip elongate in dorsal aspect with dou- ble longitudinal carina; front of bulk of epi- proct tip hastate in dorsal aspect, triangular in frontal aspect; lamella on aedeagus longer than wide with rectangular lobes (Figs. 10-13) gaufini — Epiproct tip with dorsal pandurate plane, single longitudinal carina; front of bulk of epiproct tip parallel-sided or anteriorly swollen in dorsal aspect, lozenged or lenticular in frontal aspect or wedge-like; lamella on aedeagus longer than wide with excurved lobes or as long as wide 2 Dorsal plane of epiproct tip slightly concave with carina minor and not usually rising above concavity; front of bulk of epiproct tip per- pendicular or slightly undercut in profile, wedge- like in frontal aspect; lamella on aedeagus lon- ger than wide with excurved lobes (Figs. 14—- DMSO) Wena niet ery St Ome cee canoe, albertensis — Dorsal plane of epiproct tip slightly convex with carina a prominent crest; projected front to 168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ii Figs. 39-47. 39, Sweltsa albertensis, male terminalia and aedeagus, dorsolateral aspect. 40, Sweltsa cristata, lamella of aedeagus. 41, Sweltsa lamba, lamella of aedeagus. Sweltsa cristata. 42, Subgenital plate of female, ventral aspect. 43, Subgenital plate, lateral aspect. 44, Adult head and pronotum. S weltsa lamba. 45, Subgenital plate of female, ventral aspect. 46, Subgenital plate, lateral aspect. 47, Adult head and pronotum. VOLUME 97, NUMBER 1 of bulk of epiproct tip parallel-sided or ante- riorly swollen in dorsal aspect, lozenged or len- ticular in frontal aspect; lamella on aedeagus as long as wide 3. Projection of bulk of epiproct tip as long as wide and visible as slight swelling in dorsal aspect; ventral surface of epiproct tip as wide as and aligned with dorsal plane (Figs. 21-24) b ISSR Pieces 6 Shs Oe eT aes cristata — Projection of bulk of epiproct tip longer than wide and about one-fifth length of epiproct tip in dorsal aspect; ventral surface of epiproct tip narrower than and staggered anteriorly to dor- SaliplanGe fy san oman tye tie © Ghanem ens 4 4. Dorsal plane of epiproct tip pyriform and an- teriorly acuminate (Figs. 25-34) ......... lamba — Dorsal plane of epiproct tip nearly triangular (Rigss35=5.0)) Be een aod eae hondo Sweltsa umbonata, sp. nov. Figs. 48-54 Adult.— General color dusky-marked pale tan in alcohol. Head with 3 dark ocellar rings, dusky rugulae anterior to and adja- cent to ocellar triangle and posterior to com- pound eyes; posterior ocelli connected to anterior ocellus by diffuse dusky V-mark. Pronotum with thin dark encircling line, dusky longitudinal rugulae. Meso- and me- tanota each with dark recurrent scutoscu- tellar suture. Abdomen with dark median longitudinal stripe from tergum | to 8, sin- gle dusky lateral stripes from segment | to 4, lateral pectens on segments 7, 8, 9. Ma- cropterous; wings hyaline, dusky-veined, representative of genus. Female.— Body length 11 mm; forewing length 11 mm. Subgenital plate slightly dusky, crudely hexagonal, umbonate, based centrally on sternite 8, two-thirds as long as wide, distinguishable anteriorly by narrow membranous periphery; posterior half flap- like, extending over most of sternite 9. Emargination on posterior of plate nearly quadrate with slight interior convexity; each consequent lateral flap thin, flat, trapezi- form, narrowed posteriorly, one-fifth length of plate, halfas wide as emargination, point- ing posteromediad. Tumulus centered on triangular convexity that longitudinally spans middle third of plate prominent, de- 169 clivent anteriorly, steeply sloped laterally, precipitous posteriorly. Plate thickly hirsute on tumulus and anterior declivity with hairs longer, more posteriorly directed approach- ing summit; remainder sparsely hirsute with fine, pale hairs. Vagina membranous. Male.—Body length 10 mm; forewing length 10 mm. Epiproct sclerotized, hinged, with tip extending anteriorly to prominent sclerotized shallowly crenate transverse ridge on anterior of tergum 9. In dorsal aspect, epiproct tip 9 times longer than wide, slight- ly widened at base and at two-thirds length, with distal quarter acuminate. In lateral as- pect, epiproct tip 12 times longer than thick, parallel-sided, slightly cernuous at two- thirds length, with apex circular and slightly broader than remainder. Epiproct support structures, segment 9, hemiterga of segment 10 representative of genus. Aedeagus mem- branous. Material.—Holotype ¢?, allotype ¢ (U.S. National Museum), paratype 9 4, 7 2 (U.S. National Museum): California, Shasta Co., Fowlers Campground, McCloud R. 6-VI- 1965 S. G. Jewett, Jr. Paratypes: 4 4, 2 9 (U.S. National Museum) California, Siski- you Co., Mt. Shasta, head Sacramento R. 29-V-1967 S. G. Jewett, Jr.; 7 6, 42 (US. National Museum, R. F. Surdick) same lo- cality 5-VI-1965 S. G. Jewett, Jr.; 2 6, 1 9 (U.S. National Museum) same locality 18- VI-1967 E. Evans. Compared material.—See compared ma- terial of Sweltsa adamantea, sp. nov. below. Etymology.—The adjective umbonata describes the distinctive appearance of the subgenital plate. The tumulus resembles a boss at the center of a shield. Diagnosis.—See diagnosis of Sweltsa adamantea, sp. nov. below. Sweltsa adamantea, sp. nov. Figs. 55-61 Adult.— General color darkly-marked tan in alcohol. Head with 3 dark ocellar rings, dark rugulae anterior to and adjacent to ocellar triangle and posterior to compound 170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 48-54. Sweltsa umbonata. 48, Male terminalia, dorsal aspect. 49, Epiproct tip, dorsolateral aspect. 50, Epiproct tip, lateral aspect. 51, Subgenital plate of female, ventral aspect. 52, Subgenital plate, ventrolateral aspect. 53, Subgenital plate, lateral aspect. 54, Adult head and pronotum. VOLUME 97, NUMBER 1 eyes; most of frons, ocellar triangle and ru- gulose areas dark. Pronotum with thin dark encircling line, dark longitudinal rugulae in dusky environs. Meso- and metanota dusky, each with dark recurrent scutoscutellar su- ture. Abdomen with dark median longitu- dinal stripe from tergum | to 8, single dark lateral stripes from segment | to 3, lateral pectens on segments 7, 8, 9. Wings ma- cropterous to brachypterous, hyaline, dark- veined, representative of genus. Male.—Body length 9 mm; forewing length of holotype 8 mm. Epiproct sclero- tized, hinged, with tip extending anteriorly to prominent sclerotized, shallowly crenate transverse ridge on anterior of tergum 9. In dorsal aspect, epiproct tip thrice longer than wide, an elongate lozenge with obtuse angles rounded, surface slightly depressed central- ly, verge curved ventrad; apex bluntly acute, compressed with dorsal edge appearing as short bisecting line. In lateral aspect, epi- proct tip with thickness less than half width, slightly declivent and thickened medially; apex one-fifth epiproct tip length, a colorless crescent, paddle-like with vertical elliptical blade slightly broader than thickness of bulk of tip, arising anteromedially from dorsal and ventral surfaces of bulk. Epiproct sup- port structures, segment 9, hemiterga of seg- ment 10 representative of genus. Aedeagus membranous. Female.— Body length 11 mm; forewing length 10 mm. Subgenital plate slightly dusky, crudely hexagonal, based centrally on sternite 8, almost as long as wide, dis- tinguishable anteriorly by narrow membra- nous periphery; posterior half flap-like, ex- tending over most of sternite 9. Emar- gination on posterior of plate bisected by slightly obtuse angular projection; each con- sequent lateral flap thin, flat, trapeziform, narrowed posteriorly, pointing posterome- diad, one-sixth length of plate, one-third longer than interposed angle, half as wide as emargination. Median triangular con- vexity longitudinally spanning middle third of plate based anteriorly, terminated pos- 171 teriorly as angular projection, a plateau in profile, thickly hirsute; remainder of plate sparsely hirsute with fine hairs more nu- merous laterally. Vagina membranous. Material.—Holotype 4, allotype 2 (U.S. National Museum), paratype | 6, 1 2(R. F. Surdick): Oregon, Yamhill Co., McMinn- ville, Peavine Ridge 8-VI-1948 K. M. Fend- er. Additional material: 1 ¢ (S. G. Jewett, Jr.) Oregon, Benton Co., Wren; 1 4 (U.S. National Museum) Washington, Grays Harbor Co., Humptulips. Compared material.—Sweltsa borealis: lectotype 2, cotypes (Museum of Compar- ative Zoology); hundreds of specimens from Alaska, Alberta, British Columbia, Califor- nia, Colorado, Idaho, Montana, Oregon, Utah, Washington, Wyoming, Yukon. Sweltsa fidelis: lectotype 2, cotypes (Muse- um of Comparative Zoology); hundreds of specimens from Alberta, British Columbia, California, Colorado, Idaho, Montana, Or- egon, Washington, Wyoming, Yukon. Sweltsa revelstoka: holotype @, allotype 4 (W. E. Ricker); hundreds of specimens from Al- berta, British Columbia, Montana, Oregon, Washington, Wyoming. (Above species, Brigham Young Univ., California Academy of Science, M. Cather, D. Dunster, A. R. Gaufin, Field Museum of Natural History, R. E. Hill, Illinois Natural History Survey, S. G. Jewett, Jr., Michigan State Univ., Montshire Museum of Science, Oregon State Univ., W. E. Ricker, Royal Ontario Mu- seum, B. P. Stark, R. F. Surdick, Univ. of Alberta, Univ. of British Columbia, Univ. of Kansas, Univ. of Minnesota, Univ. of Montana, Univ. of Nebraska, U.S. Nation- al Museum, Utah State Univ., Washington State Univ., R. Wisseman.) Swel/tsa contin- ua: lectoallotype 2, cotypes (Museum of Comparative Zoology); specimens from Los Angeles, Riverside, San Bernardino, San Diego Cos., California (Brigham Young Univ., California Academy of Science, Field Museum of Natural History, S. G. Jewett, Jr.). Sweltsa californica: holotype é¢ (Cali- fornia Academy of Science); specimens from 172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 55-61. Sweltsa adamantea. 55, Epiproct and center of transverse ridge on tergum 9, dorsal aspect. 56, Epiproct tip, dorsolateral aspect. 57, Epiproct tip, nearly lateral aspect. 58, Subgenital plate of female, ventral aspect. 59, Subgenital plate, ventrolateral aspect. 60, Subgenital plate, lateral aspect. 61, Adult head and pro- notum. VOLUME 97, NUMBER 1 Butte, Placer Cos., California (California Academy of Science, R. E. Hill). Etymology.—The adjective adamantea describes the rhomboid dorsal aspect of the epiproct tip. Diagnosis.—Sweltsa umbonata and S. adamantea are closely related to Sweltsa bo- realis (Banks) 1895 (Figs. 62-65), Sweltsa fidelis (Banks) 1920 (Figs. 66-70), and Sweltsa revelstoka (Jewett) 1955 (Figs. 71- 74). All these western Nearctic species have a similarly elongate, club-like or slightly flattened epiproct tip and an emarginate, approximately hexagonal female subgenital plate. They have a median convexity on the subgenital plate that either does not extend posteriorly as far as the emargination or does not project beyond the lateral flaps formed by the emargination. In the latter case, the projected portion, whether acute, obtuse or slight, is about twice as broad as a lateral flap. Sweltsa continua (Banks) 1911 (Figs. 75— 79, 85) and Sweltsa californica (Jewett) 1965 (Figs. 80-84, 86) also have an emarginate subgenital plate and are included in the Sweltsa borealis group but are more dis- tantly related. Both species have a broad median convexity on the subgenital plate that projects beyond the lateral flaps, a dis- tinctly sculptured epiproct tip and a pro- nounced median dark stripe on the prono- tum. The five species that include S. umbonata and S. adamantea differ from each other in the shapes of the median convexity and lat- eral flaps of the subgenital plate, in the se- tation of the subgenital plate, in the location and width of the widest or flattest part of the epiproct tip, and in the prominence of the compressed apex of the epiproct tip. Some species have a consistent color pattern that can be an aid in determining them. The subgenital plate of all five species bears a triangular convexity on the middle third. In S. borealis, S. fidelis and S. ada- mantea, the convexity is shallow and pla- teau-like in profile but it obviously swells 173 above the sparsely haired lateral thirds that terminate in the thin flaps flanking the emargination. The convexity, or at least its anterior two-thirds, is more hirsute than the remainder of the plate. In S. borealis, the convexity is usually only slightly projected as a small curve or very obtuse angle in the emargination between the lateral flaps. In S. fidelis and S. adamantea, the convexity is usually slightly deeper than in S. borealis and, projecting between the flaps as a nearly right angle, extends half as to almost as far as the flaps extend. In S. umbonata, the con- vexity itself is not pronounced and it pro- jects in the emargination as a slight broad curve. But, there is a prominent boss on the center of the convexity. The boss and the anterior of the convexity where it slopes down from the boss is thickly hirsute with long hairs. In S. revelstoka, the convexity is usually pronounced but reaches posteriorly from the base to only about midway on the plate so it does not project in the square emargination. It is thickly hirsute with long hairs. In S. borealis, the lateral flaps that flank the emargination of the subgenital plate are usually more circular or square than tra- peziform and are often slightly notched or scalloped. They range from small out-curves on the posterior margin of the plate to dis- tinct flaps one-fifth the length of the plate. Their shape and length often differ within and between populations. As a pair, they occasionally exhibit asymmetry and aber- rations of shape or development. In S. fidelis and §. adamantea, the flaps are trapezi- form, often point posteromedially and con- stitute about one-fifth the length of the plate. In S. revelstoka, the flaps, although also tra- peziform and about one-fifth the length of the plate, do not arise as distinctly from the lateral margins of the plate. This gives the plate the appearance of an oblate hexagon with rounded angles and with a prominent, round-shouldered, square notch in its pos- terior margin. In all five species, the epiproct tip is to 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 7 | Figs. 62-74. Sweltsa borealis. 62, Epiproct tip, dorsal aspect. 63, Epiproct tip, lateral aspect. 64, Subgenital plate of female and variations, ventral aspects. 65, Subgenital plate and variation, lateral aspects. Sweltsa fidelis. 66, Epiproct tip, dorsal aspect. 67, Epiproct tip, dorsolateral aspect. 68, Epiproct tip, lateral aspect. 69, Subgenital plate of female, ventral aspect. 70, Subgenital plate, lateral aspect. Sweltsa revelstoka. 71, Epiproct tip, dorsal aspect. 72, Epiproct tip, lateral aspect. 73, Subgenital plate of female, ventral aspect. 74, Subgenital plate, lateral aspect. VOLUME 97, NUMBER | 175 ib | Figs. 75-86. Sweltsa continua. 75, Epiproct tip, dorsal aspect. 76, Epiproct tip, dorsolateral aspect. 77, Epiproct tip, lateral aspect. 78, Subgenital plate of female, ventral aspect. 79, Subgenital plate, lateral aspect. Sweltsa californica. 80, Epiproct tip, dorsal aspect. 81, Epiproct tip, dorsolateral aspect. 82, Epiproct tip, lateral aspect. 83, Subgenital plate of female, ventral aspect. 84, Subgenital plate, lateral aspect. 85, Sweltsa continua, adult head and pronotum. 86, Sweltsa californica, adult head and pronotum. 176 some extent flattened for most of its length and compressed at its partly colorless tip. In S. borealis, it is one-tenth to one-eight as wide and as thick as long, only barely flat- tened anywhere along its length, slightly de- clivent, and nearly parallel-sided with its apical quarter acuminate in dorsal aspect. Its apex is rounded in profile and as thick as or slightly thicker than the bulk of the epiproct tip. Its shape differs within and be- tween populations. The epiproct tip of S. umbonata is similar to that of S. borealis, but it increases slightly in width at two-thirds its length, is more declivent and has an apex slightly larger in profile. In S. fidelis, S. revelstoka and S. ada- mantea, the epiproct tip is wider and slight- ly thicker with a more pronounced apex than in the above two species. It widens and nar- rows along its length. Because the com- pressed apex arises from the dorsal and ven- tral surfaces, the epiproct tip resembles a large horizontal rhomboidal disk intersect- ed on its apex by a small vertical elliptical disk for half the diameter of the small disk. In S. fidelis, the epiproct tip reaches its greatest width, almost one-fourth its length, at about three-fifths its length. Its sides are slightly incurved from base to widest part and it is acuminate beyond. In S. revelstoka, the epiproct tip reaches its greatest width, about one-sixth its length, at three-fourths its length. Its sides are also slightly incurved from base to widest part and it is acuminate beyond. In S. adamantea, the epiproct tip reaches its greatest width, about one-third its length, at about half its length. Its sides are slightly incurved only adjacent to the base; beyond that, its sides are slightly ex- curved and it ends acutely. Sweltsa borealis is usually the largest of the Chloroperlinae and macropterous but its body length and wing length differ within and between populations. The crenulations on the transverse ridge are often compara- tively very shallow. Sweltsa revelstoka, al- though other macropterous, commonly ex- hibits different degrees of brachyptery. The PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON S. adamantea examined are macropterous and brachypterous. Swel/tsa fidelis is usually macropterous and the S. umbonata exam- ined are macropterous. Sweltsa borealis differs, sometimes great- ly, in adult coloration and distinctness of color pattern within and between popula- tions. But, whether pale or dark overall, the rugulae of the head and pronotum are dark- ened, the posterior ocelli are usually con- nected to the anterior ocellus by a dark V and the pronotum is ringed by a dark line. Sweltsa umbonata, S. revelstoka and S. ada- mantea are usually darkly and distinctly colored in the same pattern as S. borealis. In S. revelstoka and even darker S. ada- mantea, however, a broad dark area on the head usually extends from the ocellar tri- angle to the clypeus. Sweltsa fidelis differs in coloration within and between popula- tions but it is usually pale overall. The ru- gulae are barely noticeable or darkened, the ocelli are usually not connected by a dark V and the dark ring on the pronotum is often interrupted, forming a pair of lateral cres- cents. Sweltsa borealis is a common, wide-rang- ing species. It has been found from Alaska to New Mexico and to California. Sweltsa fidelis ranges from Alaska to Utah and to California and S. revelstoka ranges from Al- berta and British Columbia to Wyoming and Oregon. Swe/tsa adamantea has been found in areas of the Coast Ranges of northern Oregon and Washington. Swe/tsa umbonata has been found in the Mount Shasta area of California between the southern Cascade Range and northern Sierra Nevada. KEY TO NEARCTIC SWELTSA WITH THE SUBGENITAL PLATE OF THE FEMALE EMARGINATE 1. Median longitudinal dark stripe on pronotum; in dorsal aspect, epiproct tip narrowed to less than basal width at one-third length, abruptly widened to wider than base in middle third then abruptly and sharply acuminate; median convexity on female subgenital plate projecting posteriorly between emargination to or beyond VOLUME 97, NUMBER 1 apices of lateral flaps where it is at least half width of posterior margin of plate ......... — Median longitudinal area of pronotum pale; in dorsal aspect, epiproct tip nearly parallel-sid- ed, slightly widened or gradually widened to up to twice basal width then gradually tapered to apex or acuminate; median convexity on female subgenital plate either not extending posteriorly as far as emargination or not ex- tending beyond apices of lateral flaps where it is less than half width of posterior margin of DIACCH eee eee meee ten eee gceees 3} 2. Dark ring encircling pronotum; apex of epi- proct tip up-curled, hook-like; median con- vexity of subgenital plate blunt, extending slightly beyond apices of lateral flaps (Figs. 80- SAMS 6) tet, See sone se ated mad californica — No dark ring encircling pronotum; apex of epi- proct tip up-curved, blunt; median convexity of subgenital plate rounded, extending beyond apices of lateral flaps (Figs. 75-79, 85) .... continua 3. Epiproct tip nearly parallel-sided, acuminate distally; lateral flaps of subgenital plate more squarcion rounded were ee iseh ates eee 4 — Epiproct tip widened somewhere along its length, acuminate or acute distally; lateral flaps of subgenital plate more trapeziform 4. Median convexity of subgenital plate with prominent central hirsute boss (Figs. 48-54) 56 OR uecict ASHIES ACRrat eCE art aee umbonata — Median convexity of subgenital plate without aUBOSSACHIGS: 02-09) x. cri noc. stan, aon borealis 5. Median convexity of subgenital plate promi- nent, reaching from base to midway on plate; epiproct tip reaching greatest width at about three-fourths its length (Figs. 71-74) ..revelstoka — Median convexity of subgenital plate extend- ing length of plate and projecting as an obtuse angle in the emargination between the lateral flaps; epiproct tip reaching greatest width at about three-fifths or one-half its length 6. Epiproct tip reaching its greatest width, almost one-fourth its length, at about three-fifths its length; epiproct tip acuminate with sides slight- ly incurved; ocelli usually not connected by Garkava(Bigs 466-70)". 2 tne eee 2 ee fidelis — Epiproct tip reaching its greatest width, almost one-third its length, at about half its length; epiproct tip acute apically with sides slightly excurved; ocelli connected by dark V and dark area reaching to clypeus (Figs. 55-61) 177 ACKNOWLEDGMENTS I am grateful to M. S. Kelley of the Mu- seum of Comparative Zoology, N. Adams and O. S. Flint, Jr. of the U.S. National Museum, R. W. Baumann of Bngham Young University, S. G. Jewett, Jr. and A. R. Gaufin for their assistance and loan of specimens and to all the museums and col- lectors noted in the text. LITERATURE CITED Banks, N. 1895. New neuropteroid insects. Trans- actions of the American Entomological Society 22: 313-316. 1911. Descriptions of new species of North American neuropteroid insects. Transactions of the American Entomological Society 37: 335-360. 1920. New neuropteroid insects. Bulletin of the Museum of Comparative Zoology 64: 299- 362. Baumann, R. W. 1973. Studies on Utah stoneflies (Plecoptera). Great Basin Naturalist 33: 91-108. Baumann, R. W. and G. Z. Jacobi. 1984. Two new species of stoneflies (Plecoptera) from New Mex- ico. Proceedings of the Entomological Society of Washington 86: 147-154. Baumann, R. W., A. R. Gaufin, and R. F. Surdick. 1977. The stoneflies (Plecoptera) of the Rocky Mountains. Memoirs of the American Entomo- logical Society 31. 208 pp. Jewett, S. G., Jr. 1955. Notes and descriptions con- cerning western North American stoneflies (Ple- coptera). Wasmann Journal of Biology 13: 145- ISS: 1960. The stoneflies (Plecoptera) of Califor- nia. Bulletin of the California Insect Survey 6: 125- WF 7e 1965. Four new stoneflies from California and Oregon. Pan-Pacific Entomologist 41: 5-9. Needham, J. G. and P. W. Claassen. 1925. A mono- graph of the Plecoptera or stoneflies of America North of Mexico. Thomas Say Foundation of the Entomological Society of America 2. 397 pp. Ricker, W.E. 1952. Systematic studies in Plecoptera. Indiana University Publications Science Series 18. 200 pp. Surdick, R. F. 1985. Nearctic genera of Chloroper- linae (Plecoptera: Chloroperlidae). Illinois Biolog- ical Monographs 54. 146 pp. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 178-196 NOTES ON THE MINDARUS SPP. (HOMOPTERA: APHIDIDAE) OF NORTH AMERICA WITH DESCRIPTIONS OF TWO NEW SPECIES DAVID VOEGTLIN Center for Biodiversity, Illinois Natural History Survey, Champaign, Illinois 61820- 6970. Abstract.—A background of the biology and taxonomy of Mindarus spp. in North America is presented. A previously unused character, wax gland plates on apterous adult female morphs, has been found to be relatively conservative and useful for discriminating between species. An extensive study of specimens from western North America and Europe suggests that M/. abietinus Koch, a European aphid to which most specimens have been assigned, may not be present in the western Nearctic. Two species, Mindarus kinseyi, n. sp. living on Abies concolor and M. remaudierei, n. sp. living on Abies religiosa, are described. The distribution of . victoria Essig is extended to California and Idaho where it lives on A. concolor. Key Words: Aphids of the genus Mindarus are mon- oecious, needle feeders on Abies, Keteleeria or Picea. In this holarctic genus are five fos- sil and five extant species. Heie (1967) dis- cusses the living species in relation to the fossils and provides a key to eight species. The literature reports three species from North America: Mindarus abietinus Koch, M. obliquus (Cholodkovsky) and M. victoria Essig. Mindarus obliquus lives on Picea and the latter two live on Abies. Because of the morphological similarity between obliquus and abietinus they have been identified pri- marily on the basis of the host on which they were collected. Robinson and Chen (1969) found differences between the chro- mosomes of Mindarus from Picea and those from Abies, and Carter and Eastop (1973) found it possible to separate Mindarus col- lected in Britain from these two hosts by plotting the number of sensoria on the third antennal segment against the length of the segment. Mindarus victoria has been re- Aphididae, Mindarus, aphid, new species corded only from Abies grandis in the type locality, Vancouver Island, British Colum- bia, Canada (Smith and Parron 1978). The life cycle of Mindarus abietinus is reduced in number of generations and rel- atively simple. The fundatrices produce e1- ther alate sexuparae or apterous viviparae that produce only sexuparae. These second and third generation sexuparae produce males and oviparae which mate and by early summer eggs have been deposited on the host. A period of 9-10 months is spent in the egg stage. The biology and seasonal his- tory of M. abietinus has been studied in de- tail by Varty (1966, 1968). This cycle has been common to all species studied until recent work by Ehler and Kinsey (in press) who found that Mindarus living on Abies concolor on the west slope of the Sierra Ne- vada, most particularly pest populations in the United States Forest Service Nursery in Placerville, California, had a life cycle and biology which clearly did not match any VOLUME 97, NUMBER 1 published for M. abietinus (Niisslin 1900, Varty 1966, 1968). This aphid is described below. DATA COLLECTION Hundreds of specimens have been pro- vided by Marvin Kinsey (Department of Entomology, University of California, Da- vis) taken from Abies concolor in the Sierra Nevada. In addition he has reared clones from fundatrices through several genera- tions, collecting good series from these clones. Approximately 1500 slides, each containing a minimum of two aphids, have been prepared from this material. Addi- tional specimens were borrowed from sev- eral museums. Measurements were taken of body, wing, hind tibia, antennal segments III, IV, V, VI base and VI process terminalis, ultimate rostral segment, second hind tarsus and counts were made of secondary sensoria on antennal III and IV of alatae, setal counts were made on cauda, ultimate rostral seg- ment, first hind tarsus, subgenital plate, ab- dominal tergite VIII and scape. The distri- bution of wax gland plates was recorded on apterous female morphs. Measurements were made with the use of a drawing tube on a Zeiss® microscope extending out over a Zidas® digitizing pad. With this system it is possible to make measurements accurate to three decimal places using a 16 x or great- er objective. Wax gland plates (WGPs) have not pre- viously been used to discriminate between species of Mindarus. They are found on fun- datrices, apterae, oviparae and nymphs. They are irregularly shaped, heavily scler- otized plates ranging from circular to oval on which are located groups of cells (Fig. 1). The structure of these cells under high mag- nification is very regular, appearing much like a sieve (Fig. 2). Foottit and Richards (1993) call these cells ‘““margined cribriform discs” and WGPs “clusters of emarginate cribriform discs.”> Sometimes the plate 1s only lightly sclerotized (especially in ovi- 179 parae) but a cluster of closely associated clearly defined cells will still define WGPs in such cases. The number of cells ina WGP can vary from one to over a hundred (Fig. 3). Each gland is accompanied by a setae which is located on the anterior half of its margin. Wax gland plates may be found on the head, thorax, abdomen and subanal plate. Typically there are a pair on the front of the head and a pair on the anal plate. Those on the thorax and abdomen may be assigned to paired marginal, dorsolateral and submedian lines (terminology for these lines after Foottit and Richards 1993). In slide- mounted specimens the lateral line often appears to be ventral. These can be ex- pressed as a formula which is maximally 6.6.6:6.6.6.6.6.6.4. The first three numbers refer to number of WGPs on the thoracic segments and the later seven refer to the WGPs on the first seven abdominal seg- ments. Unless otherwise indicated a 2 means that there are 2 marginal WGPs present, a 4 means that there are 2 marginal and 2 dorsolateral WGPs, and a 6 means that there are 2 marginal, 2 dorsolateral and 2 sub- median WGPs on that segment. No speci- mens have been seen with more than 4 WPGs on abdominal segment VII. WGPs on the prothorax are usually easy to see and appear to be located on the head due to the fusion of head and prothorax in Mindarus (Fig. 4). Diagrammatic representations of the wax gland distribution on various spe- cies is shown in Fig. 7. Although there is intraspecific variation, within a range, the distribution patterns seem to hold. The for- mula for M. victoria is almost invariant on specimens seen from several locations in California and British Columbia taken over several years. Oviparae have two large wax producing structures located on the abdominal venter (Fig. 5). These have somewhat different structure than WGPs in that the entire sur- face, with the exception of a central pore, is glandular with fine lines creating a matrix of cells of variable shape. 180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. sclerotized outer ring. 2, High magnification of wax gland plate of aptera of M. kinseyi showing sieve-like structure within cells. Siphuncular pore and associated sclerite shown at top of photo. 3, A small wax gland plate containing only three cells. 4, Wax gland plate on prothorax posterior to eye of M. kinseyi aptera. 5, Ovipara of M. kinseyi showing paired ventral wax glands found on abdominal venter of all Mindarus oviparae. 6, Abdominal tergite VI of M. kinseyi alata showing membranous wax gland completely embedded in sclerite. Note adjacent glandular areas. Most of the sclerotization on the dorsum of the body in both apterae and alatae con- tains less organized glandular areas usually surrounding a seta. These areas are most likely wax producing glands without the dis- crete structure of the WGPs discussed above. Although present in alatoid nymphs, wax gland plates as described above are not pres- ent in alatae. On the abdomen of alatae at the locations where WGPs would be ex- pected in apterae, there are membranous areas with a dark central spot and a seta (Fig. 6). In some cases under high magni- 1, Abdominal tergites V, VI and VII of M. kinseyi aptera showing wax gland plates with distinct fication fine radial lines can be seen around the central spot. The submedian pair of these membranous areas 1s often embedded in the posterior edge of the sclerotic bands on each tergite (Fig. 8). The dorsolateral and mar- ginal rows are indicated only by the dark- ened pore and associated seta and are often very difficult to see. Since alatae have some abdominal wax, usually appearing as trans- verse bands, and these structures are located where WGPs are in nymphs and apterae, it may be that they are wax producing glands. I have been unable to find a name for these VOLUME 97, NUMBER 1 Bigs. and submedian lines of wax gland plates as observed in the following species: A & B, minimum and maximum distribution seen on M. kinseyi fundatrices; C & D, minimum and maximum distribution seen on M. kinseyi apterae; E, distribution seen on M. remaudierei apterae; F, distribution seen on M. victoria apterae; G & H, minimal and maximal distribution seen on M. abietinus apterae. structures so will call them membranous wax glands (MWGs). Mindarus Koch 1857 Detailed generic descriptions are given in Heie (1980) and Foottit and Richards (1993) and will not be repeated here. Mindarus is the only genus in the subfamily Mindarinae. It is separated from most other aphids by the presence of ventral wax glands in ovip- arous females and the pterostigma in alatae 181 Diagrammatic representation of apterous morph of Mindarus showing paired marginal, dorsolateral extends to the tip of the wing. Males and Oviparae are mostly much smaller than vi- viparous morphs. In apterous morphs there is no visible separation between the head and prothorax. Eyes in fundatrices, ovipar- ae and males consist only of triommatidia while apterae have compound eyes. Only two named species, Mindarus ja- ponicus Takahashi (1931) and M. keteleer- ifoliae Zhang and Zhong (1984), have not been recorded from North America and will not be covered in this paper. 182 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 8-11. Mindarus abietinus Koch Except for Mindarus victoria, specimens of Mindarus from Abies spp. in North America have been identified and referred toas M. abietinus. Based on the morphology of the alatae there is little reason to doubt this. A comparison of the fundatrices and apterae suggests otherwise. Apterae and fundatrices from Europe have a limited dis- tribution of wax gland plates. Dr. G. Re- maudiére (Museum National D’Histoire Naturelle, Paris) and Dr. R. Blackman (Nat- ural History Museum, London) kindly ex- amined additional fundatrices and apterae in their collection, and confirm this general 8, Abdomen of M. kinseyi alata showing maximum level of sclerotization and also membranous wax glands associated with posterior edge of sclerites on anterior segments. 9, Dorsal abdominal sclerite of M. kinseyi alata showing irregular transverse sculpturing. 10, Cauda shape and size typical of M. kinseyi alata. 11, Subgenital plate of M/. kinseyi alata showing paired setae located on approximate midline of plate. pattern of reduction in WGPs in M. abie- tinus. The formula for WGPs ranged from 0:0:0:0.0.0.0.0.2:2 to'0,0:0:0:0.2:2:2:3:47 Dr. Remaudiére has two apterae from Greece with two “‘very small’ WGPs on the pro- thorax which have the formula 2.0.0: 0.0.0.2.2.2.2. This is in considerable con- trast to the patterns seen in fundatrices and apterae from North America (Fig. 7). I have examined several hundred fundatrices and apterae from multiple years, hosts, distri- butions and elevations and have yet to find one without at least one pair of lateral WGPs on the prothorax and, with the exception of one specimen, all have a complete paired lateral line on the first seven abdominal seg- VOLUME 97, NUMBER 1 ments. The paucity of apterae of abietinus in European collections is also in contrast to the abundance of apterae here. With the exception of the aphids studied by Varty (1966, 1968) the number of offspring of the fundatrices (based on collections I have made and those sent to me) which are ap- terous is significant, reaching nearly 100% in some cases. Much additional work needs to be done but I believe that none of the material I have seen, at least from western North America, is M. abietinus. Mindarus kinseyi, n. sp. Fundatrix Color in life: Pale green, mature speci- mens covered with filamentous wax, es- pecially laterally and posteriorly. Color of fundatrices and other forms from Ehler and Kinsey (in press). Morphology: Apterous. Eyes only triom- matidia, however some specimens exam- ined that are considered fundatrices have a few facets in addition to the triommatidia. Body oval, enlarging gradually from head to abdominal segment IV then rounding to caudal region. Head, scape and pedicel smooth, antennal segments IIJ-VI with gradually increasing density of spiculose imbrications. Head and prothorax fused. Rostrum reaching abdominal segment II, ultimate rostral segment gradually tapering to tip with convex sides. Dorsum of thorax and abdomen smooth with sclerotized areas appearing grandular, on posterior segments with irregular transverse lines of spicules. Trochanters and femora fused, joint indi- cated only by a slight ventral crease. Legs smooth through femora, tibiae lightly spi- culose on distal half, second tarsal segment with spiculose imbrications. Cauda slightly raised, transverse ridge. Subgenital plate with spiculose imbrications. Without si- phunculi. Setation: Setae very sparse, and short (<0.015 mm). Dorsal setae on a scler- ite or edge of a gland. Ultimate rostral seg- ment with 0-2 accessory setae. Cauda with 2 setae. Subgenital plate with 2 setae on 183 anterior margin and 7-14 scattered along posterior, some specimens with 2 additional setae on midline of plate. Sclerotization (Fig. 12): Intensity of scle- rotization varies from pale to medium tan. Head with one central sclerite covering middle of vertex and front but not reaching antennal sockets or eyes. Antennae with scape, pedicel, V and VI and distal '4 of segment IV darker than segment III and basal 73 of IV. Paired sclerites on prothorax extend onto posterior of head, mesad of triommatidia. Prothorax with two pairs of small submedian sclerites. Meso- and meta- thorax, and abdominal tergites I-III (IV) each with a row of small, irregularly shaped sclerites. Abdominal tergites (IV), V-VII with sclerotized areas which are highly var- ied in size and usually extending across mid- dorsal line especially on the latter two seg- ments. Tergite VIII with a narrow sclerotic band most often divided into a median and two lateral sections. Cauda, subanal plate and subgenital plate lightly sclerotized. Legs more or less evenly sclerotized throughout. Wax gland plates: Formula ranging from 2.2.2:0.2.2.2.2.4.4 to 2.2.2:4.4.4.4.4.6.4 (Fig. 7A & B). Size: Summary of measurements given in Table 1. Apterae and alatae are divided into two categories in the description below. These categories, based on date of collection, roughly correspond to observed differences in biology noted between spring and later generations of both morphs (Ehler and Kin- sey 1n press). Aptera (second and third generation) Description and measurements based on specimens collected in May and June. Color in life: Body pale green covered with a light wax pulverulence which increas- es with age. Posterior and lateral areas of abdomen can be covered with long wax fil- aments extruded from wax glands. Morphology: Compound eyes present. Body enlarged gradually from narrow head 184 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 12-17. 12, Fundatrix of M. kinseyi. 13, Aptera of M. kinseyi showing close to maximum level of sclerotization. 14, Alata of M. kinseyi showing a moderate level of sclerotization on abdomen. 15, Alata of M. victoria. 16, Fundatrix of M. victoria. 17, Aptera of M. victoria. to mid abdomen rounding smoothly to blunt caudal region; dorsum smooth with faint network of spiculose lines on abdominal tergite VIII. Head and thorax fused, ap- pearing as one unit. Rostrum reaching ab- dominal segment III, ultimate rostral seg- ment gradually tapering to blunt tip: Antennae six-segmented without secondary VOLUME 97, NUMBER 1 sensoria, scape and pedical smooth, flagel- lum increasingly spiculose distally. Legs smooth, second tarsi with light spiculose imbrications. Cauda slightly elevated, transversely elongate. Siphunculi small rimmed pores sometimes surrounded by a small sclerotic area. Subgenital plate usually rugose and always spiculate. Setation: Setae on body sparse, and short (<0.015 mm). Dorsal setae all on a sclerite or on the edge of a gland. Ultimate rostral segment with 2 accessory setae, rarely 1 or 0. Subgenital plate with 2 setae near middle of anterior margin and (5) 8-14 scattered along pos- terior margin. Some specimens with an ad- ditional smaller pair in the middle of the plate or an additional smaller setae on either side of the middle pair on the anterior mar- gin. Sclerotization (Fig. 13): On body from medium to light tan. Head with wide, par- allel sided, median sclerite covering vertex and front, median suture indicated on both vertex and front. Irregularly shaped scle- rotic areas on posterior of head and pro- thorax. Sclerites on meso- and metathorax and tergites I and II small, scattered across segment in an irregular line, gradually in- creasing in size on tergites III and IV and tergites V-VII each with transverse band; tergite VIII with narrow band divided into submedian and two lateral sections. Scape and pedicel concolorous with central head sclerite, distal tip of II] and 4 of IV slightly darkened, V and VI darker than head. Legs evenly tanned, coxae and trochanters light- Sr. Wax gland plates: Formula commonly ranging from 2.0.0:2.2.2.2.4.5.4 to 2.0.0: 4.4.4.4.6.6.4 (Fig. 7C & D). One specimen seen without a complete paired marginal line of glands and one specimen seen with thoracic formula 2.2.2. Size: Summary of measurements given in Table 1. Alata (second and third generation) Descriptions and measurements based on specimens collected in May and June. 185 Color in life: Body light to medium green, shiny in recently molted adults becoming covered with a light pulverulence in mature specimens. Morphology: Head clearly separate from narrow prothorax, ocelli on vertex located on anterior-mesal margin of compound eyes, ocellus on front located at base of indistinct median suture. Rostrum reaching to ab- dominal segment II, ultimate rostral seg- ment with slightly convex sides gradually tapering to rounded tip. Antennae six-seg- mented with 8—23 (mean = 14.5, var. = 6.6) secondary sensoria on segment III and 0-4 (mean = 1.9, var. = 0.7) on segment IV, scape pedicel and III smooth, IV—VI with spiculose imbrications. Abdomen gradually widening from pterothorax to abdominal segment V then tapering to more pointed abdomen than apterae, prothorax and membranous areas of dorsum smooth, dor- sal lobes of pterothorax with reticulate sculpturing, sclerotized bands on abdomi- nal segments I-VIJ with irregular transverse sculpturing (Fig. 9). Legs smooth except for anterior surface of femora and second tarsi which have spiculose imbrications. Siphun- culi rimmed pores, rarely absent, some- times surrounded by a small sclerite. Cauda rugose, subtriangular with rounded tip, usu- ally wider than long (Fig. 10). Subgenital plate spiculose. Setation: On body sparse as in apterae and very short (<0.01 mm). Ab- dominal segments with a row of setae on the sclerotic band. Setae on tibiae < half tibial diameter. Ultimate rostral segment with 2, rarely 1 or O, accessory setae. Subgenital plate usually with 2 large setae located near midline and 8-15 scattered along posterior margin (Fig. 1 1). Some spec- imens with a second smaller pair between midline and posterior group or one on each side of median pair. Sclerotization (Fig. 14): Head and pter- othorax heavily sclerotized, prothorax pale. Scape and pedicel concolorous with head and darker than flagellum which is evenly sclerotized. Coxae and trochanters pale, femora gradually darkening distally, tibiae 186 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Measurements for Mindarus spp. For each character the range is given, the mean is in boldface followed by the variance. The n values for each morph vary because on some of the specimens certain characters were not measurable. Length of antennal segment VI is not separated into base and process terminalis for the Oviparae and males. All measurements are in mm. Antennal Segments Body Wing Hind Tibia Ill IV M. kinseyi Fundatrices 1.42-1.80 0.253-0.431 .115-.198 0.054—0.096 n= 17-23 1.62, 0.009 0.306, 0.003 0.149, 0.000 0.073, 0.000 Apterae May-June 1.35-2.27 0.319-0.560 0.174-0.332 0.087-0.170 n= 116-119 1.84, 0.027 0.468, 0.002 0.275, 0.001 0.136, 0.000 July—Sept. 1.09-2.14 0.292-0.577 0.155-0.330 0.074—0.168 n= 129-131 1.56, 0.042 0.416, 0.003 0.243, 0.001 0.118, 0.000 Alatae May-June 1.28-2.90 1.61-3.47 0.453-0.925 0.195-0.475 0.102-0.242 n= 199-212 2.08, 0.115 2.58, 0.160 0.704, 0.008 0.356, 0.002 0.187, 0.001 July—Nov. 1.02—2.83 1.30-3.28 0.346-0.864 0.173-0.439 0.098-0.231 n= 154-156 1.79, 0.090 2.26, 0.147 0.601, 0.010 0.307, 0.002 0.169, 0.001 Oviparae 0.074-1.00 0.131-0.196 0.040-0.066 0.016-0.043 n = 68-69 0.88, 0.002 0.161, 0.000 0.051, 0.000 0.031, 0.000 Males 0.52-0.66 0.126-0.174 0.050-0.090 0.029-0.051 = 35-36 0.60, 0.001 0.150, 0.000 0.065, 0.000 0.040, 0.000 M. remaudierei Apterae 2.01-2.34 .510-.550 0.295-0.314 0.107-0.127 n=4 2.16, 0.019 0.528, 0.000 0.302, 0.000 0.116, 0.000 Alatae 2.05-3.00 2.91-3.48 0.706-0.851 0.449-0.510 0.171-0.210 n= 15 2.43, 0.063 3.23, 0.033 0.808, 0.002 0.490, 0.000 0.192, 0.000 M. victoria Fundatrices 1.67-2.20 0.382-0.565 0.207-0.312 0.108-0.169 n= 40 1.96, 0.018 0.512, 0.001 0.274, 0.000 0.140, 0.000 Apterae 1.76-2.47 0.537-0.688 0.287-0.415 0.158-0.218 n = 32-33 2.08, 0.031 0.620, 0.002 0.368, 0.001 0.187, 0.000 Alatae 1.67-2.85 2.13-3.52 0.562-0.870 0.333-0.484 0.168—-0.260 n = 47-50 2.36, 0.070 3.01, 0.106 0.785, 0.005 0.419, 0.001 0.218, 0.000 Oviparae 1.11-1.94 0.307-0.494 0.159-0.307 0.076-0.152 n= 39-40 1.53, 0.033 0.410, 0.002 0.241, 0.001 0.119, 0.000 Males 0.769-0.889 0.226-0.275 0.09 1-0.139 0.05 1-0.064 n= 11 0.808, 0.001 0.252, 0.000 0.114, 0.000 0.058, 0.000 and tarsi evenly sclerotized. Abdominal ter- Membranous wax glands: The most gites I-VII with transverse bands, decreas- ing 1n size on anterior segments. Tergite VIII with narrow band which is sometimes in- complete forming a median and two mar- ginal sections. Cauda and anal region darker than unevenly sclerotized subgenital plate. common arrangement seen is two on ab- dominal segment VI within the sclerotic band and two on segment VII that are not contained completely within the band. Fig. 8 shows the maximum development of these glands in spring alatae. VOLUME 97, NUMBER 1 187 Table 1. Extended. Aatennal Seements Ultimate Second Antennal Vv VI base VI pt Rostral Seg. Hind Tarsus Flagellum M. kinseyi 0.083-0.108 0.091-0.114 0.020-0.030 0.048-0.070 0.102-0.137 0.386-0.540 0.092, 0.000 0.101, 0.000 0.026, 0.000 0.057, 0.000 0.118, 0.000 0.440, 0.002 0.095-0.168 0.092-0.152 0.024-0.039 0.05 1-0.080 0.128-0.194 0.484-0.832 0.136, 0.000 0.127, 0.000 0.032, 0.000 0.069, 0.000 0.168, 0.000 0.706, 0.005 0.087-0.174 0.088-0.168 0.023-0.042 0.051-0.083 0.119-0.199 0.336-0.873 0.127, 0.000 0.122, 0.000 0.031, 0.000 0.066, 0.000 0.159, 0.000 0.638, 0.007 0.124-0.243 0.1 10—-0.203 0.023-0.043 0.058-0.087 0.125-0.227 0.554-0.150 0.192, 0.001 0.158, 0.000 0.032, 0.000 0.071, 0.000 0.187, 0.000 0.925, 0.111 0.114—0.231 0.104—0.190 0.021-0.040 0.056-0.08 1 0.125-0.208 0.513-1.076 0.178, 0.001 0.149, 0.000 0.031, 0.000 0.067, 0.000 0.172, 0.000 0.834, 0.011 0.032-0.063 0.071-0.099 0.034-0.049 0.057-0.077 0.160-0.267 0.050, 0.000 0.082, 0.000 0.044, 0.000 0.067, 0.000 0.214, 0.000 0.036-0.069 0.073-0.104 0.030-0.039 0.056-0.071 0.198-0.303 0.050, 0.000 0.086, 0.000 0.033, 0.000 0.064, 0.000 0.241, 0.001 M. remaudierei 0.150-0.166 0.141-0.158 0.031-0.040 0.088-0.096 0.157-0.179 0.732-0.805 0.157, 0.000 0.146, 0.000 0.36, 0.000 0.091, 0.000 0.168, 0.000 0.757, 0.001 0.197-0.240 0.162-0.190 0.031-0.042 0.087-0.102 0.172-0.193 0.930-1.164 0.218, 0.000 0.174, 0.000 0.037, 0.000 0.093, 0.000 0.183, 0.000 1.098, 0.004 M. victoria 0.119-0.192 0.117-0.190 0.021-0.029 0.060-0.078 0.141-0.219 0.576-0.857 0.168, 0.000 0.164, 0.000 0.025, 0.000 0.072, 0.000 0.198, 0.000 0.772, 0.003 0.187-0.231 0.173-0.207 0.022-0.034 0.070-0.083 0.207-0.247 0.850-1.079 0.207, 0.000 0.190, 0.000 0.028, 0.000 0.077, 0.000 0.234, 0.000 0.980, 0.003 0.186-0.276 0.166-0.242 0.026-0.041 0.067-0.085 0.1840.266 0.879-1.247 0.237, 0.000 0.209, 0.000 0.032, 0.000 0.077, 0.000 0.234, 0.000 1.116, 0.007 0.099-0.183 0.127-0.205 0.061-0.079 0.140-0.199 0.478-0.830 0.147, 0.000 0.170, 0.000 0.068, 0.000 0.178, 0.000 0.676, 0.000 0.079-0.093 0.115-0.135 0.040-0.047 0.108-0.127 0.364-0.402 0.085, 0.000 0.124, 0.000 0.045, 0.000 0.121, 0.000 0.381, 0.000 duce viviparous offspring and those pro- ducing sexuales. Description and measurements made from specimens col- lected July through November. Color in life: Similar to spring apterae. Morphology. Similar to spring apterae. Eye size is highly variable in this group. Size: Summary of measurements given in Table 1. Aptera (late season viviparae and sexuparae) It is not possible to discriminate mor- phologically between those apterae that pro- 188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 18-21. 18, Compound eye of M. kinseyi aptera showing reduction in facets common in summer apterae. 19, Magnification of ventral wax gland of M. kinseyi ovipara showing combination of mostly triangular and square cells. 20, Genital structure of M. kinseyi ovipara. 21, Genital structure of M. kinseyi male. There seems to be a general decrease in number of facets as the season progresses. Not all specimens exhibit this but late sum- mer apterae with only a few facets in ad- dition to the triommatidia are common (Fig. 18). Siphuncular pores generally without a sclerite. Setation: As in spring apterae. Sclerotization: Pattern of sclerotization similar to the apterae described above, how- ever, there is a gradual reduction in size and VOLUME 97, NUMBER 1 intensity of tanning of sclerites as the sum- mer progresses. Specimens collected during the winter show increased darkness and size of sclerites so this variability may be in part a function of temperature. In the most re- duced form the entire body is almost com- pletely pale with only the wax glands visible on the abdomen. Sclerites are first lost on anterior segments then bands on tergites V, VI and VII begin to disintegrate into indi- vidual pale sclerites, finally becoming very small and almost invisible. Wax gland plates: Formula ranging from 2.0.0:2.2.2.2.4.6.4 to 2.0.0:4.4.4.4.4.6.4. Size: Summary of measurements given in Table 1. Alata (late season viviparae and sexuparae) It is not possible to discriminate mor- phologically between those alatae that pro- duce viviparous offspring and those that produce sexuales. Descriptions and mea- surements made from specimens collected July through November. Color in life: Similar to spring alatae. Morphology: Similar to spring alatae. There is a gradual decrease in size of alatae throughout the season (Table 1). Siphun- cular pores may not have an associated sclerite, sometimes absent. Setation: As in spring alatae. Sclerotization: This group shows tre- mendous range of presence and intensity of sclerotization. Specimens developing in July and August range from the sclerotization of spring alatae to a point where virtually none of the sclerotic bands on the abdominal seg- ments are visible and the pterothorax and appendages are pale. Specimens collected in September through February are more like- ly to show sclerotization intensity of spring generations. This seasonal variation in scle- rotization is similar to that seen in apterae. Wax glands: Wax production areas ap- pear to be associated with sclerites and since the sclerites are greatly reduced or absent the amount of glandular area on the dorsum 189 appears to be much reduced in some spec- imens. The paired MWGs on abdominal segments VI and VII are usually not con- tained within the sclerotized bands. Size: Summary of measurements given in Table 1. Ovipara Color in life: Entire body yellow to gold. Not as covered with wax as are other ap- terous morphs. Morphology: Apterous. Eyes only triom- matidia. Body elongate oval appearing par- allel sided from mesothorax to abdominal segment V, dorsum smooth with faint spi- culose lines visible on posterior abdominal segments. Head and prothorax fused, ap- pearing as one unit. Rostrum reaching ab- dominal segment IV, ultimate rostral seg- ment blunt with slightly convex sides. Antennae six-segmented without secondary sensoria, segments I-IV smooth, V and VI with a few spiculose imbrications. Cauda only slightly elevated. Siphuncular pores absent. Hind tibia with (1) 2-8 (11) scent plaques (Fig. 23). A modified genital struc- ture present (Fig. 20) consisting of a base and two posteriorly directed arms. Setation: Much reduced as in apterae. Ultimate ros- tral segment with 2 sometimes | or 0 ac- cessory setae. Subgenital plate with 6-10 se- tae along posterior margin, 2 near anterior margin. Sclerotization (Fig. 23): Head with me- dian parallel-sided sclerite extending over vertex and front but not enclosing base of antennae or triommatidia. Median suture visible on front but not vertex. Front with a pair of wax gland plates. Antennae with scape, pedicel, segments V and VI some- what darker than head, III and IV pale. Legs evenly sclerotized and only slightly darker than antennae. Hind tibiae with 4-13 irreg- ularly shaped scent plaques. No dorsal scle- rites visible on thorax or abdomen. Cauda light tan. Subgenital plate at most lightly sclerotized, indicated by setal arrangement. Wax gland plates: Formula ranging from 190 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 23 Figs. 22-25. 22, Ovipara of M. victoria. 23, Ovipara of M. kinseyi. 24, Male of M. victoria. 25, Male of M. kinseyi. 222 0:2.232.2.3.4 to 2.2.22 .4:4.4.4:6-4 more variable than in fundatrices and ap- terae. When present on the first abdominal tergite the two glands are in the dorsolateral line, not the marginal as in apterae. Two large, asymmetrical glands on the abdomi- nal venter (Fig. 5) produce elongated, straight wax filaments which protrude from the glands perpendicular to the body. These glands are granular in appearance and have a lattice like substructure creating numerous triangular and square cells (Fig. 19). There is also a central pore in which (0) 2-4 (5) cells are visible. Size: Summary of measurements given in Table 1. Male Color in life: Entire body blue to blue green. Morphology: Apterous. Eyes only triom- matidia. Small, elongate oval body almost VOLUME 97, NUMBER 1 parallel sided from mesothorax to abdom- inal segment V. Head and prothorax fused, appearing as one unit. Rostrum reaching mesocoxae, ultimate rostral segment short, blunt with convex sides. Antennae 6 seg- mented with secondary sensoria on seg- ments IV (1-2), V (1-3) and VI (2-4); seg- ments I-IV (I-III) smooth, (IV) V-VI with some spiculose imbrications. Trochanter- femoral joint fused, indicated only by faint suture. Legs smooth throughout. Thoracic sclerites smooth, abdominal sclerites in- creasingly spiculose on posterior segments. Cauda not developed but indicated by two setae. Siphuncular pores absent. A genital structure present (Fig. 21). Setation: Very reduced. Ultimate rostral segment without accessory setae. Sclerotization (Fig. 25): Head sclerotized throughout, thoracic and abdominal seg- ments each with wide transverse band. Scape and pedicel, distal '2 of III, IV, V and VI concolorous with head. Legs evenly dark throughout and only slightly darker than body. Wax glands: served. Size: Summary of measurements given in Table 1. Types: Holotype, morphotypes and all paratypes collected on Abies concolor in California by Marvin Kinsey unless other- wise noted. Holotype: Aptera, top specimen circled on slide #92-63-12 with three other apterae. Abies concolor, USFS Nursery, Pla- cerville, Eldorado County, California, 14 June 1992, M. Kinsey, deposited at the II- linois Natural History Survey, Champaign, Illinois. Morphotypes: Fundatrix, Cold Springs, Toulumne County, 18 June 1992, M. Kinsey & D. Voegtlin (top specimen cir- cled on slide #92-55-22 with one aptera); alata, Westpoint, Calaveras County, 24 June 1991 (on slide #92-6-1 with aptera); ovipara and male, USFS Nursery, Placerville, El- dorado County, 7 June 1992 (circled on slide #92-61-40 with one other male and ovi- para); all deposited at INHS. Paratypes: Ca- No glandular areas ob- 191 laveras County: Sonora, 26 June 1991, 10 ap., 20 al.; 3 July 1991, 2 fund. West Point, 24 June 1991, 12'aps 13\al 10 July 1991, 5 ap., 19 al. Eldorado County: Camino, 15 May 1990, 5 ap., 1 al., 13 June 1991, 20 ap., 10 al.; 11 September 1990, 16 ap., 18 al. Morgan, 6 June 1991, 10 fund., 16 ap., 15 al. USFS Nursery, Placerville, 24 August 1989, 22 ap., 22 al., L. Ehler; 30 August 1989, 19 ap., 14 al., L. Ehler; 21 June 1990, 11 ap., 11 al.; 30 July 1990, 12 ap., 12 al.; 28 August 1990, 5 ap., 19 al.; 11 September 1990, 12 ap., 8 al.; 28 August 1991, 6 ap., 18 al.; 2 October 1991, 12 ap., 12 al. Sly Park, 25 June 1991, 16 ap., 16 al.; 10 July 1991, 6 ap., 10 al. 40 km S. E. of Sly Park, Eldorado Nat. For., 7 July 1992, 3 ap., 26 al., 12 ovip., 12 male. Plumas County: Grey Eagle, 7 July 1991, 4 ap., 5 al. Siskiyou County: Mt. Shasta, 20 June 1991, 3 ap., 3 al. Tulare County: Peppermint Camp Ground, nr. Johnsondale, 17 June 1992, 2 fund., 4 ap., 29 al., 3 ovip., 3 male, M. Kinsey & D. Voegtlin. Sequoia Kings Can- yon N. P., 27 July 1992, 11 ap., 14 al. Toul- umne County: Yosemite Institute, 18 June 1992, 7 al., M. Kinsey & D. Voegtlin. Cold Springs, 18 June 1992, 2 fund., 22 ap., 21 al., 5 ovip., 1 male, M. Kinsey & D. Voeg- tlin. Hardin Flat, 18 June 1992, 2 fund., 5 ap., 16 al., 2 ovip., 4 male, M. Kinsey & D. Voegtlin. Strawberry, 19 June 1992, 2 fund., M. Kinsey & D. Voegtlin. Paratypes will be deposited at the following museums: Unit- ed States National Museum, Beltsville; The Natural History Museum, London; Muse- um National D’Histoire Naturelle, Paris; Canadian National Museum, Ottawa, all re- maining slides are on deposit at INHS. Discrimination: The biology of this spe- cies separates it from all other Mindarus species. Fundatrices and apterae can easily be separated from M. abietinus by the WGPs (see discussion of M. abietius above). Many alatae have more than one sensoria on an- tennal segment IV, a condition uncommon in M. abietinus but common in M. obliquus. The tremendous variabilities in size and 192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 26-31. 26, Holotype of M. remaudierei. 27, Morphotype of M. remaudierei. 28, Head of M. remaudierei aptera showing few eye facets in addition to the triommatidion. 29, Subgenital plate of M. remaudierei aptera showing paired setae on anterior margin and cluster of setae at midplate. 30, Dorsal abdominal sclerite of M. remaudierei aptera showing paired setae on anterior margin and cluster of setae at midplate. 31, Antennal segments III and IV of M. remaudierei alata showing large number of secondary sensoria on III. pigmentation of the alatae make it virtually and field work detailed much of the biology impossible to separate this morph from MM. — of this species (Ehler and Kinsey in press) abietinus or M. obliquus. and who collected and reared the majority Etymology: This species is named after of specimens on which this description is Marvin Kinsey whose careful observations — based. VOLUME 97, NUMBER | Mindarus remaudierei, n. sp. Aptera Color in life: Not seen alive. Morphology: Body oval, increasing in width to abdominal segment IV then round- ing to caudal region. Eyes only a few facets in addition to the triommatidia (Fig. 28). Head, scape and pedicel smooth, spiculose imbrications gradually increasing from an- tennal segments III-VI. Two of the four ap- terae seen with one secondary sensoria on antennal segment III but they have no other indications of alatoidy such as small wing buds and large compound eyes. Rostrum reaching abdominal segment II, ultimate rostral segment with gradually tapering slightly convex sides and rounded tip. Head and prothorax fused, smooth as is the re- mainder of the dorsum. Legs smooth to sec- ond tarsi which have spiculose imbrica- tions. Subgenital plate spiculose. Cauda slightly elevated. Siphuncular pore rimmed. Setation: Very sparse and short <0.015 mm. Ultimate rostral segment with 2 accessory setae; cauda with 2 setae; subgenital plate with 20-23 setae arranged with 2 on the anterior half, from 2-6 in the central area and 14-17 along the posterior edge. Sclerotization (Fig. 26): Head with front and much of vertex sclerotized, enclosing antennal sockets but not eyes. Irregular pale sclerotic band extending across prothorax and abdominal segment VIII. Remainder of dorsum clear. Antennae slightly darker than head with only basal '3-'2 of segment III lighter. Legs medium brown, darker than antennae with base of femora and trochan- ters paler. Cauda and subanal plate only slightly darkened. Subgenital plate with ir- regular dark areas. Wax gland plates: Formula ranging from 4.0.0:5.6.6.6.6.6.4 to 4.0.1:6.6.6.6.6.6.4 (Fig. 7E). Without a distinct sclerotized ring around each plate. The four on the protho- rax are on the submedian and marginal lines. Three of the four specimens have an addi- tional WGP on the vertex. 193 Size: Summary of measurements given in Table 1. Alata Color in life: Not seen in life. Morphology: Head smooth with area on vertex posterior to eyes with longitudinal ridges. Antennal scape, pedicel and III smooth, IV-VI imbricated, most imbrica- tions on segments IV and V without spic- ules, those on VI spiculose. Antennal seg- ment III with 45 to 54, and segment IV with 1-5 secondary sensoria (Fig. 31). Ptero- thorax rugose. Anterior surface of femora and second tarsi with spiculose imbrica- tions, remainder of legs smooth. Non-scler- otized regions of the abdomen smooth. Sclerotic bands on abdominal segments with limited sculpturing, much less than seen in other Mindarus spp. (Fig. 30). Siphuncular pore usually surrounded with a small, pale sclerotized area. Cauda subtriangular with rugose knob at tip, wider than long. Subgen- ital plate spiculose. Setation: Sparse and short <0.015 mm. Tibial setae 0.22 mm, sclerotic bands on ab- domen very pale not enclosing membra- nous wax glands and the secondary sensoria on antennal segment III often not parallel sided, rather appearing as irregular polygons especially on the distal half of the segment (Fig. 32). In apterae the dorsum 1s free of sclerotization, length of second hind tarsal segment >0.20 mm and wax gland plates are large and lack a distinct outer sclerotized ring (Fig. 33), WGP formular is 2.0.0: 4.4.4.4.4.4.6(5).4 (Fig. 7F). The oviparae are much larger than seen in any other species (compare Figs. 22 and 23) and mounted specimens have been seen with as many as five eggs. Mindarus obliquus (Cholodkovsky) I have examined only a limited number of specimens labeled as M. obliquus. The few apterae seen have a WGP formula sim- ilar to M. kinseyi so they are easily separated from M. abietinus. The alatae are small and often have more than one secondary sen- soria on antennal segment IV. In these char- acters it overlaps with M. kinseyi, however, its host specificity (Carter and Eastop 1973) and differences in chromosomes (Robinson and Chen 1969) make it clear that it is dis- tinct from the species feeding on Abies spp. At present there is no morphological basis for distinguishing between apterae or alatae of M. obliquus and M. kinseyi. DISCUSSION Heie (1967) noted that the Asian species M. japonicus Takahashi overlaps in counts VOLUME 97, NUMBER | aN ms) A é 195 Figs. 32, 33. 32, Antennal segment III from two alatae of M. victoria showing distinct polygonal shape of sensoria, especially on distal half of segment. 33, Abdomen of aptera of M. victoria showing wax gland plates without a distinct outer sclerotized ring. of secondary sensoria with M. abietinus and suggested that they are only isolated geo- graphical populations of the same species. I think there 1s little to be gained from this concept. Detailed studies such as that car- ried out by Ehler and Kinsey (in press) and additional morphological examination will undoubtedly reveal distinct biological and morphological differences between popu- lations of Mindarus which are great enough to warrant specific standing. Additional work on the Mindarus spp. in North Amer- ica needs to be done and is in progress. This study has focused primarily on western North America but I have collected and ex- amined material from the eastern half of the country and the Rocky Mountain region. Much of this material shows distinctive wax gland patterns which need to be verified with collections over several years from a variety of elevations and locations. Unfortunately the prospects for accurate identification of trap collected alate Min- darus are not good. Limits of variation will have to be developed from material which can be associated with identifiable apterae. Eventually it may be possible to develop discriminant functions for alatae. Unfor- tunately the abbreviated life cycle of most of the species makes gathering this material a long term project. ACKNOWLEDGMENTS The following individuals very kindly ar- ranged for the loan of slides of Mindarus. J. Powell, University of California, Berke- ley; R. Foottit, Centre for Land and Bio- logical Resources Research, Ottawa, Can- ada; G. Remaudiére, Museum National D’Histoire Naturelle, Paris, France; R. Blackman, Natural History Museum, Lon- don, United Kingdom; S. Halbert, Univer- sity of Idaho, Aberdeen; A. Jensen, Uni- versity of Oregon, Corvallis. Unmounted material was supplied by N. Winchester, Victoria, British Columbia, Canada, and M. Kinsey and L. Ehler, University of Califor- nia, Davis. L. Ehler initiated this study by asking me to identify an aphid causing prob- lems in the Placerville Nursery and M. Hav- erty, United States Forest Service, Albany, California provided funding for part of this study. The manuscript has been improved by the review and suggestions of W. LaBerge and D. Webb, Illinois Natural History Sur- vey, Champaign and M. B. Stoetzel, Sys- tematic Entomology Laboratory, Beltsville, Maryland. 196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Carter, C. I. and V. F. Eastop. 1973 (1972). Mindarus obliquus (Chol.) (Homoptera, Aphidoidea) new to Britain and records of two other aphids recently found feeding on conifers. Entomologists Monthly Magazine 108: 202-204. Cholodkovsky, N. 1896. Uber die auf Nadelhélzern vorkommenden Pemphigiden. Zoologischer An- zeiger 19: 257-260. Ehler, L. E. and M. G. Kinsey. Ecology and manage- ment of Mindarus kinseyi Voegtlin (Aphidoidea: Mindaridae) on white fir seedlings at the Placer- ville Nursery. Hilgardia (In press). Essig, E.O. 1939. A new aphid of the genus Mindarus from white fir in British Columbia. The Pan-Pa- cific Entomologist 15: 105-110. Foottit, R.G. and W.R. Richards. 1993. The genera of the aphids in Canada. Homoptera: Aphidoidea and Phylloxeroidea. Part 22 of The Insects and Arachnids of Canada. Centre for Land and Bio- logical Resources Research, Ottawa, Ontario Pub- lication 1885. 765 pp. Heie, O. E. 1967. Studies on fossil aphids (Homop- tera: Aphididae). Spolia Zoologica Musei Hau- niensis 26, Copenhagen. 273 pp. . 1980. The Aphidoidea (Hemiptera) and Fen- noscandia and Denmark. I. General Part. The Families Mindaridae, Hormaphididae, Thelaxi- dae, Anoeciidae, and Pemphigidae. Fauna Ento- mologica Scandinavica 9: 1-236. Koch, C. L. 1854-1857. Die Pflanzenlause Aphiden, getreu nach dem Leben abgebildet und geschrie- ben. Niirnberg. 336 pp. Niisslin, O. 1900. Zur biologie der Schizoneuriden— gattung Mindarus Koch. I. Die eiblage und das auskommen der fundatrix aus dem winterei. Biol- ogisches Zentralblatt 20: 479-485. Robinson, A. G. and Y. Chen. 1969. Cytotaxonomy of Aphididae. Canadian Journal of Zoology 47: 511-516. Smith, C. F. and C. S. Parron. 1978. An annotated list of Aphididae (Homoptera) of North America. North Carolina Agricultural Experiment Station Technical Bulletin No. 255. 428 pp. Takahashi, R. 1931. A new Mindarus from Japan. Transactions of the Natural History Society of Formosa 21(114): 137-139. Varty, I. W. 1966. The seasonal history and popu- lation trends of the balsam twig aphid, Mindarus abietinus Koch in New Brunswick. Internal Report M-12. Fredericton, New Brunswick, Canada, For- est Research Laboratory, Department of Forestry and Rural Development. 21 pp. . 1968. The biology of the balsam twig aphid, Mindarus abietinus Koch, in New Brunswick: Polymorphism, rates of development, and season- al distribution of populations. Internal Report M-24. Fredericton, New Brunswick, Canada, For- est Research Laboratory, Department of Forestry and Rural Development. 64 pp. Zhang, G. and T. Zhong. 1984. A new species of Mindarus Koch from Yunnan, China. Entomo- taxonomia 6(2-3): 235-236. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 197-224 THE NEW WORLD GENUS CHROMOLEPIDA COLE (DIPTERA: THEREVIDAE: THEREVINAE) DONALD W. WEBB AND MICHAEL E. IRWIN Illinois Natural History Survey, 605 East Peabody Drive, Champaign, Illinois 61820. Abstract. —Species of the genus Chromolepida Cole are confined to western North Amer- ican and northern South America. The species are revised, a phylogeny is hypothesized, and a key to the species is provided. Two species from México, Chromolepida clavitibia Webb and Irwin and C. nigra Webb and Irwin, are described as new to science. Key Words: Diptera, stiletto flies, Therevidae, Chromolepida, New World revision, phy- logeny, species keys, descriptions, distributions This paper is the fifth of a series (Webb and Irwin 1988, Webb and Irwin 199 1a, b, c) revising the species of the genera treated in the monograph of the Nearctic genera of Therevidae. A key to the species of Chrom- olepida Cole is provided along with species descriptions and distributions. The morphological terminology used for the male terminalia was originally defined and described by Lyneborg (1968) and sub- sequently modified by Lyneborg (1972, 1976, 1978) and Irwin (1977a, b). The fe- male terminalia terminology was defined and described by Irwin (1976). Other mor- phological features are described with ter- minology set out in the “Manual of Nearctic Diptera” (McAlpine 1981). The range for each setal count is followed by the mode. Setal counts were taken from the left side of the specimen and from the entire pos- terior margin of the scutellum. Each specimen was assigned a unique THEREVIDAE/M. E. IRWIN/SPECI- MEN number. This number is used to as- sociate the ecological and label data with a given specimen, and when referred to here- in, is printed in italics. The data are in- tended to be incorporated into an auto- mated data management system originally designed by Rauch (1970). To conserve space and include as much information as possible about each speci- men, a format adopted from Irwin (1983) is used in the ““Specimens Examined” sec- tion of each species. Many of the terms used there were defined by Stuckenberg and Ir- win (1973). In the presentation of repetitive locality data, a semicolon terminates one series of specimens and signals the begin- ning of the next, thus, data not repeated in a series are the same as those of preceding series. PHYLOGENETIC RELATIONSHIPS The genus Chromolepida belongs to the Cyclotelus group of therevids, along with Cyclotelus Walker, Ozodiceromya Bigot, and an undescribed genus from Chile. This group is characterized by rather bulbous gonocox- ites. The black, glossy calli on the face ven- tral to the antennal bases and the appressed, iridescent, silver green scales on the thorax and abdomen separate Chromolepida as a monophyletic genus within this group. The cladogram (Fig. 1) represents the re- lationship we hypothesize for the five spe- 198 Fig. 1. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Primitive Ancestor bella pruinosa nigra epidajowom clavitibia mexicana QUT Cladogram showing the hypothesized phylogeny of the species of Chromolepida. Numbers on the branches refer to apomorphic alternatives of characteristics listed in Table 1. (R) indicates a reversal in a character State. cies of Chromolepida based on character- istics given in Table 1. Although no hypothesized generic-level phylogeny has been attempted for the Therevidae, our work to date strongly suggests that the Cyc/lotelus group is more derived than members of the genus Phycus (Phycus group) and the genus Dialineura (Thereva group). The determi- nation of the plesiomorphic condition, then, was based on an examination of the char- acteristics found in New World species of Dialineura and Phycus. In these genera, the plesiomorphic con- dition for Characters 1, 4, and 5 was found in all species. Character 2. The plesio- morphic state of a narrow, simple gono- stylus was found in most species. Only in the one species of Dialineura was a lateral projection found. Character 3. The anterior margin of the male dorsal apodeme of the aedeagus generally has a single transverse attachment to the anterior margin of the hypoproct. Only in Phycus frontalis is the dorsal apodeme bifurcate, forming a double attachment to the hypoproct. The character matrix is given in Table 2. A heuristic search of this matrix using PAUP (Version 3.1.1, Swofford 1993) resulted in a single tree of seven steps (consistency in- dex = 0.714, retention index = 0.750). This cladogram was rooted from a hypothetic an- cestor possessing the primitive state of all characteristics. Chromolepida bella formed a clade pos- sessing a simple gonostylus (Fig. 4) without a lateral projection, although it does possess the most specialized and distinctive disti- phallus which is more elongate and re- VOLUME 97, NUMBER 1 Table 1. Character states for Chromolepida. Plesiomorphic Apomorphic — . Female frontal setae present . Male gonostylus simple . Male dorsal apodeme truncate or slightly emarginate anteriorly . Male ventral apodeme truncate or slightly emarginate anteriorly . Endophallus simple a wh A) Female frontal setae absent Male gonostylus with lateral projection Male dorsal apodeme deeply emarginate anteriorly Male ventral apodeme pointed, extended anteriorly Endophallus recurved curved than all other species of Chromole- pida. This species currently ranges from California to southern Washington, east to Idaho, northern Utah, and Nevada. Chromolepida pruinosa formed a south- erm, neotropical clade separated from C. bella by the possession of a lateral projec- tion on the male gonostylus and from C. nigra, C. clavitibia, and C. mexicana in re- taining the more primitive characteristics of the male dorsal and ventral apodemes of the aedeagus. It currently ranges from Guy- ana to Colombia, north to Nicaragua. Chromolepida nigra, C. clavitibia, and C. mexicana form a clade defined by the male dorsal apodeme of the aedeagus being deep- ly emarginate anteriorly and the anterior apodeme of the aedeagus being pointed and extended anteriorly. These species are cen- tered in Mexico and the southwestern Unit- ed States with C. nigra found in southern Mexico, but not sympatric with C. pruinosa. Chromolepida clavitibia and C. mexicana form the final clade based on the presence of frontal setae in the females of both spe- cies. Chromolepida clavitibia apparently Table 2. Character matrix for Chromolepida. Character matrix Taxa l 2 3 4 5 Chromolepida bella Chromolepida clavitibia Chromolepida mexicana Chromolepida nigra Chromolepida pruinosa NNNNE KBNNN= EBENNNE =e eK NWN NNN =) NO separated from C. mexicana and 1s cur- rently restricted to the southern tip of Baja California de Sur, with C. mexicana ranging from northern Mexico to southern Califor- nia, Arizona, New Mexico, western Texas, and southern Colorado, but not sympatric with C. bella. Chromolepida may be separated from other Nearctic therevids by using the keys to the genera of Nearctic Therevidae in Ir- win and Lyneborg (198 1a, b). No key to the genera of Neotropical Therevidae is avail- able at this time; however, all species of Chromolepida have a pair of shiny, black, raised calli on the upper face below the an- tennal bases, which is unique to species of this genus. Chromolepida Cole Chromolepida Cole (1923: 23). Type spe- cies: Psilocephala pruinosa Coquillett (1904: 91) by original designation. Derivation of name.—chroma (Greek) = color; /epido (Greek) = scale. Diagnosis.— Chromolepida is a member of the Cyclotelus group of therevid genera. Beyond Chromolepida, this group includes Cyclotelus Walker, Ozodiceromya Bigot, and an undescribed genus from Chile. Species in this group of genera can be characterized by the following: middle coxae without se- tae on posterior surface; male gonocoxites often strongly fused ventrally, forming a rather compact capsule; dorsal apodeme of male aedeagus weakly to strongly joined to internal dorsal portion of gonocoxite (= par- 200 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-11. view. 3. Male sternite 8, gonocoxites and gonostylus, ventral view. 4. Male gonocoxites and gonostylus, dorsal view. 5. Male gonocoxite, lateral view. 6. Male aedeagus, dorsal view. 7. Male aedeagus, ventral view. 8. Male aedeagus, lateral view. 9. Female terminalia, dorsal view. 10. Female terminalia, ventral view. 11. Female furca, dorsal view. Scale = 0.1 mm, unless otherwise indicated. ameral process); male hypandrium small or absent; male gonocoxites usually strongly fused with lateral margins of epandrium; posterior parameral processes composed of pair of nonarticulated projections; and male distiphallus of aedeagus often curved, ex- tending posteroventrally beyond gonocox- ites. Chromolepida is separated from the other genera of the Cyclote/us group in hav- ing a pair of shiny, black, raised calli on the face just below the antennal bases, and the thorax and abdomen clothed with ap- Chromolepida bella (9177, 9179) 2. Male tergite 8, epandrium, cercus, and hypoproct, dorsal pressed iridescent silver green scales in ad- dition to normal setae. Small to medium-sized flies. Description of male and female. — Head: Eyes of male holoptic, ommatidia on ven- tral third smaller; eyes of female dichoptic, all ommatidia of similar size; antenna (Fig. 42), length 0.9-1.7 times head length; scape cylindrical; pedicel cylindrical to ovate; fla- gellum awl-shaped, tapered apically; style subapical, two-segmented, including min- ute apical spine; frons in female (Fig. 52) VOLUME 97, NUMBER 1 18 Figs. 12-21. Chromolepida clavitibia (8812, 8805) 12. Male tergite 8, epandrium, cercus, and hypoproct, dorsal view. 13. Male sternite 8, gonocoxites and gonostylus, ventral view. 14. Male gonocoxites and gonostylus, dorsal view. 15. Male gonocoxite, lateral view. 16. Male aedeagus, dorsal view. 17. Male aedeagus, ventral view. 18. Male aedeagus, lateral view. 19. Female terminalia, dorsal view. 20. Female terminalia, ventral view. 21. Female furca, dorsal view. Scale = 0.1 mm, unless otherwise indicated. broad, lateral margins convergent dorsally; frontal calli glossy, with distinct area of black pile dorsolateral to antennal bases; parafa- cial broad with enlarged, glossy parafacial calli; maxillary palpus one-segmented (Fig. 43), cylindrical to slightly clavate, rounded apically. Setae of two types (filiform, and appressed, iridescent, silver green scale-like), scattered filiform on female frons; short, fi- liform on antenna, abundant, filiform on maxillary palpus, absent on eyes, parafacial, clypeus and generally on male frons. Genal setae white, filiform and silver green scale- like, ventral projection of gena with con- centration of short, black, filiform setae. Macrosetae thick, filiform, moderately long, scattered on apex of scape. Thorax: Macrosetae: np 3-4, sa 1, pa 1, de 0, sc 4. Postpronotal lobe concolorous with thorax. Vittae indistinct. Setae of 2 types: filiform, on prosternum, mesonotum, propleuron, anepisternum, and dorsal third of katepisternum, and appressed, iridescent, silver-green scale-like on mesonotum, pro- 202 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON P| 28 Figs. 22-31. Chromolepida mexicana (8814, 9025) 22. Male tergite 8, epandrium, cercus, and hypoproct, dorsal view. 23. Male sternite 8, gonocoxites and gonostylus, ventral view. 24. Male gonocoxites and gonostylus, dorsal view. 24. Male gonocoxite, lateral view. 26. Male aedeagus, dorsal view. 27. Male aedeagus, ventral view. 28. Male aedeagus, lateral view. 29. Female terminalia, dorsal view. 30. Female terminalia, ventral view. 31. Female furca, dorsal view. Scale = 0.1 mm, unless otherwise indicated. pleuron, anepisternum and dorsal third of katepisternum, absent on remaining pleural sclerites. Wing (Fig. 44) hyaline to opaque, often patterned; veins generally pale brown; pterostigma indistinct to brown; setulae ab- sent; length of R, 0.8-1.3 times R,;; length of cell r, 1.7-4.0 times width; veins M,, Mp, and M, originate separately from apical margin of discal cell; cell m, open; discal cell acute basally; cell cup closed with short petiole; m-cu/r-m subequal. Legs. Coxae moderately long. Setae of two types: fili- form, and appressed, iridescent, silver-green scale-like, on anterior half of fore and mid- dle coxae and over entire hind coxa, over entire femora, filiform on tibiae. Macrose- tae dark brown, apical, 2 on fore and middle coxae, 3 on hind coxa. Abdomen: Rather narrow, gradually ta- pering from base to apex, male dorsum flat- tened, female dorsum convex. Setae fili- form, appressed, iridescent, silver green scale-like or lanceolate. Male terminalia (Figs. 2-8, 12-18, 22-28, 32-38, 42-51). VOLUME 97, NUMBER 1 203 Figs. 32-41. view. 33. Male sternite 8, gonocoxites and gonostylus, ventral view. 34. Male gonocoxites and gonostylus, dorsal view. 35. Male gonocoxite, lateral view. 36. Male aedeagus, dorsal view. 37. Male aedeagus, ventral view. 38. Male aedeagus, lateral view. 39. Female terminalia, dorsal view. 40. Female terminalia, ventral view. 41. Female furca, dorsal view. Scale = 0.1 mm, unless otherwise indicated. Tergite 8 large, about as wide as epandrium, posterior deeply emarginate. Sternite 8 somewhat reduced, posterior emarginate. Epandrium about two-thirds as long me- dially as wide, posterolateral corners round- ed; cercus free, well sclerotized, ending be- fore or slightly beyond hypoproct; hypoproct attached to membranous subepandrial plate, which extends anteriorly and attaches to an- terior margin of aedeagal dorsal apodeme. Hypandrium greatly reduced, narrow. Gon- ocoxite bulbous, sides broad in lateral view: in ventral view, gonocoxite with medial Chromolepida nigra (8815, 9239) 32. Male tergite 8, epandrium, cercus, and hypoproct, dorsal patch of short setae. Parameral apodeme reduced. Gonostylus elongate, with or with- out lateral projection. Aedeagus attached to parameres and gonocoxites; a long, weak, membranous bridge stretching from distal corners of dorsal apodeme to midsection of paramere and a stronger attachment be- tween ventral lobes of gonocoxites and ven- tral surface of aedeagus; distiphallus scler- otized, variable; ventral apodeme extends parallel to dorsal apodeme, anterior margin variable; ejaculatory apodeme about as long as ventral apodeme, slender. Female ter- 204 minalia (Figs. 9-11, 19-21, 29-31, 39-41, 53-55). Tergite 8 subrectangular, longer than wide, with broadly concave posterior mar- gin; setae black, elongate, scattered. Tergite 9+10 fused, with strong acanthophorites. Cercus triangular, membranous, with nu- merous, fine, short setae projecting poste- riorly. Sternite 8 large, longer than wide; posterior emarginate; setae black, elongate, scattered. Sternite 9 greatly modified, in- vaginated above sternite 8 to form internal sclerotized furca which is closed anteriorly; ventral surface of furca covered with lightly sclerotized membrane. Sternite 10 mem- branous, subtriangular with short, thick se- tae. Immature stages.— Unknown. Biology.—Adults of Chromolepida have been collected from flowers of a variety of plants, a reseeded range, a fen area, oak- chapparal habitats, and coastal dunes. Distribution (Figs. 56—58).—Chromolepi- da has been collected from South America (Guyana, Venezuela, Colombia) and in North America from Costa Rica to the state of Washington and eastward to Utah, Col- orado and western Texas. KEY TO SPECIES OF CHROMOLEPIDA lieeaElaltendarksbrowngeeen see eee eee 2) =waplialtersyellow ieee eaten Stine tec one 4 2. Femora dark reddish brown to black ....... 3 — Femora dark yellow .. mexicana Cole (females) 3. Fore tibia distinctly clavate -........:;....00 ... Clavitibia Webb and Irwin, n. sp. (males and fe- males) — Fore tibia cylindrical Sean bette: nigra Webb and Irwin, n. sp. (females) 4. Wing membrane without apical band; male distiphallus elongate, distinctly recurved (Fig. 8); male gonostylus lacking lateral projection (Fig. 4); male frontal setae absent ys ten ay 5 ext tal bella Cole (males and females) — Wing membrane (Fig. 44) with smoky brown Figs. 42-55. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to black apical band; male distiphallus short to moderately long, if recurved, only slightly (Figs. 18, 28, 38, 51); male gonostylus with lateral projection (Figs. 14, 24, 34, 47); frontal setae’ DIACKS. ‘tsp aengsi ee ee Oe eae one 5 5. Costal and subcostal cells of wing dark yellow; male distiphallus short, sinuate (Fig. 51) ... os ee pruinosa (Coquillett) (males and females) — Costal and subcostal cells of wing hyaline; dis- tiphallus elongate, bent at right angle to plane of aedeagus (Figs. 28, 38) 6. Gonocoxite in lateral view (Fig. 25) with distal half broad; length of fore femur to length of scape 2.4 + 0.2; length of fore tibia to length of scape 2.5 + 0.2; length of hind femur to length-of Scapers:5"-E 1054) 2) eae ee PRR yas Mer ee mexicana Cole (males) — Gonocoxite in lateral view (Fig. 35) with distal half reduced, tapered apically; length of fore femur to length of scape 1.7 + 0.2; length of fore tibia to length of scape 1.8 + 0.2; length of hind femur to length of scape 2.4 + 0.3 ... Saori Red bs nigra Webb and Irwin, n. sp. (males) Chromolepida bella Cole Chromolepida bella Cole (1923: 24): Cole (1965: 350); Irwin and Lyneborg (198 la: 260). Derivation of name.—bdel/us (Latin) = beautiful. Diagnosis.—Females of Chromolepida bella, like C. nigra and C. pruinosa, lack frontal setae. The species is separated from C. nigra in having the wing membrane pale yellow, concolorous; the anterior margin of the dorsal apodeme of the male aedeagus truncate; and the females have yellow hal- teres and femora. Chromolepida bella, un- like C. pruinosa, has pale yellow, concol- orous wing membranes; the male gonostylus lacks a lateral projection; and the male dis- tiphallus is elongate and sinuate. Redescription of holotype male (9/77).— Body length 4.0 mm. — Chromolepida pruinosa (9218, 8316) 42. Male antenna, lateral view. 43. Male maxillary palpus, lateral view. 44. Male wing, dorsal view. 45. Male tergite 8, epandrium, cercus, and hypoproct, dorsal view. 46. Male sternite 8, gonocoxites and gonostylus, ventral view. 47. Male gonocoxites and gonostylus, dorsal view (aedeagal complex removed). 48. Male gonocoxite, lateral view. 49. Male aedeagus, dorsal view. 50. Male VOLUME 97, NUMBER 1 205 (eae ee aedeagus, ventral view. 51. Male aedeagus, lateral view. 52. Female head, frontal view. 53. Female terminalia, dorsal view. 54. Female terminalia, ventral view. 55. Female furca, dorsal view. Abbreviations: Cercus (C); Distiphallus (Dp); Dorsal apodeme (DA); Ejaculatory apodeme (EA); Epandrium (Epa); Furca (F); Gonocoxite (Gx), Gonostylus (Gs), Hypoproct (Hyprct); Parameral apodeme (PA); Sternite 8 (S,); Sternite 10 (S10): Tergite 8 (T,); Tergite 9+10 (Ts+10)5 Ventral apodeme (VA). Scale = 0.1 mm, unless otherwise indicated. 206 Head: Length 0.8 mm. Ocellar tubercle dark reddish brown, silver pruinose; setae dark reddish brown. Eyes dull reddish brown; medial margin rounded. Frons dark reddish brown, glossy, with lateral silver pile and black dorsal pile, dorsal pile becoming silver dorsally. Antenna dark reddish brown, pruinose; setae dark reddish brown; macro- setae dark brown; length of antenna 1.4 times head length; length of scape 0.75 mm, 9.3 times width, 4.4 times length of pedicel; length of pedicel 0.11 mm, 1.1 times width; length of flagellum 0.34 mm, 3.4 times width, 0.4 times length of scape; length of basal stylomere 0.05 mm. Parafacial dark reddish brown, glossy with silver pile along lateral margins. Maxillary palpus pale brown; length 0.28 mm, 2.8 times width; setae pale yellow. Thorax: Macrosetae: np 3. Dark reddish brown, pruinose. Postpronotal setae white, filiform and scale-like. Prosternal setae white. Pleura dark reddish brown, pruinose, anepisternum and ventral half of katepi- sternum glossy; setae white, filiform, scat- tered on propleuron and dorsal half of kat- episternum, pale yellow, filiform, over entire anepisternum, scale-like over propleuron, anepisternum, and dorsal half of katepi- sternum. Scutellum dark reddish brown; se- tae white, filiform, and scale-like. Latero- tergite dark reddish brown, pruinose; setae dark white to pale yellow, filiform. Wing. Length 3.2 mm, 2.7 times width; pale yel- low, opaque; pterostigma indistinct. Halter dark yellow, stalk darker. Legs. Coxae dark reddish brown, pruinose. Femora dark yel- lowish brown. Fore tibiae dark reddish brown, middle and hind tibia dark yellow- ish brown. Tarsi dark brown. Coxal and femoral setae white, filiform and scale-like; tibial setae dark reddish brown. Abdomen: Dark reddish brown; setae white, filiform, more abundant laterally than dorsally, and white, appressed, lanceolate, more abundant along posterior margins. Terminalia (Figs. 2-8). Dark yellow. Ster- nite 8 moderately reduced, posterior margin slightly concave. Gonostylus lacks lateral PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 56. The distribution of Chromolepida bella in the western United States. projection. Dorsal apodeme of aedeagus broad anteriorly, wider than ventral apo- deme, anterior slightly emarginate; disti- phallus elongate, sinuate. Variation in males: Body measurements for males are provided in Table 3. Consid- erable variation in coloration was noted be- tween the holotype and most males of this species. Specimens in the San Francisco area and from Inglenook Fen appeared to be smaller and more slender in size than spec- imens inland from the coastal dune habitats where the holotype was collected. Head. Antenna reddish brown, pruinose, paler on basal half of scape; setae reddish brown. Maxillary palpus dark reddish brown. In one specimen (9045), a few frontal setae were noted at base of antenna. Wing. Membrane VOLUME 97, NUMBER 1 Fig. 57. in the southwestern United States and México. pale yellow. Halter dark yellow. Legs. Fem- ora dark reddish brown, apical half dark yellow. In one specimen (9037), the legs were entirely dark reddish brown, subshiny. Ab- domen. Tergite | dark reddish brown, ter- gites 2-8 dark yellow; setae white, lanceo- late, dense dorsally on tergites 1-3. Terminalia. The contour of the lateral mar- gin of the epandrium varies from specimen to specimen. Description of female.—Similar to male with following exceptions. Body measure- ments for females given in Table 3. 207 The distribution of Chromolepida clavitibia (Squares), C. mexicana (Circles) and C. nigra (Triangles) Head: Frons dark reddish brown, glossy dorsad of antennal bases, with dark gray pile lateral to antennal bases with silver, medial, pruinose band and a circular area of black pile dorsolateral to antennal bases; setae ab- sent. Thorax: Macrosetae: np 3. Setae white, elongate, with scattered reddish brown se- tae. Legs. Femora dark yellowish brown, be- coming dark reddish brown basally on mid- dle and hind femora. Abdomen: Dark yellowish brown, sub- shiny; setae dark reddish brown, filiform PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 208 a 7 €€-0'¢ ce b'y-6'7 adeos jo yisua]/inuiay puly Jo ysuIT 91 9°I-8'0 81 CCH Inway pury jo wus] 8 PCG C7 se=0ic adeos jo yIBua]/eIqn a10j Jo YIsUIT CA Gi=lal €1 I-01 RIQH) d10} JO YSU] Lal y7'l v7 8°7-8'1 adeos jo yisua[/Inulay I10J JO YIBUIT LT GaAlslel (GA bI-0'1 INWIgyJ I10J JO YISUIT 19 0'8-7'P C9 ¢'8-8'p pS pL-T'€ 19 ¢°9-8'¢ Tes €°9-8'7 yipim/yisuag] sndyjed Areyixeyy 6r0 O90-%h0 LeO 9S0-LE0 v0 70-00 670 cSO-9r0 080 80-870 yisug] sndyed Arey[rxey 90°0 90°0 soo 86366900700 Ss (S0'0—«é*N'0-E'0—COs«éO'| r0'0 ¢00 ~=—- 800-700 yisua] J1IWOJAIS 9°0 L'0-S'0 9°0 L:0-S'0 80 6'0-9'0 L0 20-220 b'0 L:0-€'0 yisua] odeos/yisus] winqjase|{ 67 hn 4 e€ Gic- we Ce O'b-L'7 2 9'P-S'€ Ie (SS IL76 yIpIm/yisug] winyjase|{ C60. S60-+50 SEO O”F0-SE0 980 “crO-Lc70 [vor 9P0-RE0 CEO! 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Costa Rica py @ <— ane et oo Sw 58 ioe te Fig. 58. Guyana. and scale-like. Terminalia (Figs. 9-11). Sternite 8 (Fig. 10) with posterior margin bilobed medially. Furca (Fig. 11) subrec- tangular; length 0.5 mm; anterior margins broadly rounded; posterior margin bicon- cave; lateral margins sinuate. Variation in females. —In specimen (9047) scutellum with 5 macrosetae. In specimen (9036) femora dark reddish brown, apical half of fore and middle femora dark yellow. Specimens in the San Francisco area and from Inglenook Fen appeared to be smaller and more slender in size than specimens collected inland and away from coastal dunes. Seasonal activity and distribution In the specimens examined, adults were collected between 16 April and 27 Septem- ber. The following plant taxa have been re- corded as sources from which specimens were collected: Achillea Millefolium L., Apocynum sp., Atriplex sp., Daucus pusillus Michx., Descurainia Sophia (L.) Webb, Lactuca pulchella (Pursh) DC., Lupinus sp., Phacelia ciliata Benth. Specimens have also been collected from a reseeded range, fen areas, an oak-chapparal zone, and in flight and light traps. Chromolepida bella have PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Venezuela P Guyana ye c The distribution of Chromolepida pruinosa in Nicaragua, Costa Rica, Colombia, Venezuela, and been collected from southern California to southeastern Washington eastward into Utah and Nevada (Fig. 56). Specimens examined Type material.—The holotype male of Chromolepida bella Cole (U.S. Museum of Natural History, Type No. 25929) (M. E. Irwin Therevidae Specimen Number 9/77) was collected near San Francisco, California on 6 June 1920 by F. R. Cole. Other material.—UNITED STATES. CALIFORNIA. Fox Valley, 8-VII-1935, J. Schuh, | @ (FSCA). Alameda County: Mid- way Road, 16-VIII-1957, W. W. Middle- kauff, 1 @ (UCB); Arroyo Mocho, Mines Road, 28 km NE Livermore, 18-VI-1967, P. H. Arnaud, 1 4 (CAS); Berkeley, 27-V- 1933, G. E. Bohart, 1 ¢@ (UCB); 22-V-1907, M. A. Cazier, 2 6 (AMNH); Patterson Re- serve, Del Valle Lake, 20-VIII-1973, J. Powell, 3 ¢ (UCB). Calaveras County: Big Trees, 7-VI-1931, E. C. Van Dyke, 1 2 (CAS). Contra Costa County: 6-VI-1912, 1 ¢(UCB); 11-VIII-1982, D. G. Denning, 1 ¢ (CAS); Orinda Village, San Pueblo Ridge below Eu- reka Peak, 10-VIII-1969, E. I. Schlinger, 1 2? (MEI), oak-chapparal zone; Lafayette, 25- VI-1984, E. I. Schlinger, 1 ¢ (UCB); Mount VOLUME 97, NUMBER 1 Diablo, 16-VII-1933, R. H. Beamer, 1 4 (SEM); Danville, 10-VIII-1949, F. X. Wil- liams, 1 6 (UCB); 11-VIII-1951, 2 2 (UCB); Clayton, 20-VI-1992, E. I. Schlinger 1 ° (MEI); 15-VII-1992, 2 2 (MEI). Fresno County: Silver Creek, 16 km W Mendota, 10-VI-1962, P. F. Torchio, 1 2? (OSM); Sel- ma, 4-VI-1929, R. L. Usinger, 1 ¢ 1 2(UCB). Kern County: Lost Hills, 13-VI-1957, P. Opler, 1 2 (UCB); 16 km W Kramer Junc- tion, 16-IV-1974, J. Wilcox, 1 2 (CAS). Lake County: Bogg’s Lake, 10 km SSE Kelsey- ville, 838 m, 20-VIII-1966, P. H. Arnaud, 2 6 (CAS). Lassen County: Susan River Camp, 9-VII-1949, A. T. McClay, 1 ¢ (UCD); 10-VII-1949, 1 ¢ (UCD); Bridge Creek Camp, 12-VII-1954, R. C. Bechtel, 1 6 1 2 (MEI); 40 km S Adin, 20-VI-1963, J. Wilcox, 1 6 (UCB). Los Angeles County: 27 km E Gorman, 16-IV-1962, J. A. Litsinger, 1 ¢ (UWisc); Chuchapate Ranger Station, 22-N-1959-4E* 1, Schlinger, 2-2: (UCD): Palmdale, 1-V-1968, J. Powell, 2 6 (MEI, UCB). Marin County: 3 km N Point Reyes light, 4-VI-1956, A. M. Barnes, 1 6 (UCB); Lagunitas, 11-V-1924, E. C. Van Dyke, 1 ¢ (UCB); 25-VI-1924, E. H. Nast, 1 6 1 9 (UCB). Mendocino County: Alpine Lake, 4-VITI-1955, C. D. MacNeil, 1 ¢ (UCB); Inglenook Fen, fen area, 9-15 m, 13-VI- 1973, E. I. Schlinger, 2 ¢ 2 2 (MEI, UCB); Inglenook Fen, dunes, 6-24 m, 21-VII-1972, P. A. Rauch, 2 2 (MEI); Inglenook Fen Re- serve, Ten Miles dunes, 24-VI-1982, M. Buegler, | ¢ (MEI), on sand dunes. Modoc County: 24 km E Cedarville, 3-VII-1935, J. Schuh, 2 @ (CSU, FSCA); Newell, 31-VII- 1963, J. Schuh, 1 2? (UCB), on Lactuca pul- chella. Monterey County: near Pacific Grove, 8-VII-1935, J. Schuh, 1 2 (UCM); U.N. Lanham 1 2?(UCM); Arroyo Seco, 21- V-1955, D. Ribble, 1 ¢ (SEM); Arroyo Seco Camp, 11-V-1958, R. M. Bohart, 1 ¢(UCD). Napa County: Oakville, 55 m, 6-IX-1970, P. H. Arnaud, 7 ¢ 2 2? (CAS, INHS). Nevada County: Sagehen Creek, near Hobart Mills, 25-VI-1954, J. A. Powell, 2 ¢ (UCB); 5-VII- 211 1962, M. E. Irwin, 2 6 2 2 (MEI). Placer County: Carnelian Bay, Lake Tahoe, 13-VI- 1959, R. M. Bohart, 2 6 3 2 (INHS, UCD); 24-VI-1973, 2 6(INHS, UCD); 8-VII-1955, 26 3 2 (UCD). Sacramento County: Galt, 27-VII-1952, E. I. Schlinger, 1 @ (UCD); Sacramento, 1-VIII-1955, E. A. Kurtz, 1 (UCD). San Francisco County: San Fran- cisco), 2 l=V=1922.. C. E> Boxals (UCB): sand dunes; 23-V-1927, 2 6 (CAS); 13-V- 1960, D. C. Rentz, 3 6 1 ¢ (CAS); 14-V- 1960, 2 6 (CAS); 6-VI-1920, E. P. Van Du- zee, 46 1 2(MCZ, UCB, USNM); San Fran- cisco, Laguna Puerca, 1 1-V-1960, J. Powell, 2 2 (UCB); 16-V-1960, 9 63 2? (INHS, UCB, UCD); 24-V-1961, 1 6 (UCB); 13-VI-1961, W.E. Ferguson, | ¢ (UCB); 14-VI-1960, G. I. Stage, 2 2 (UCB); San Francisco, Golden Gate Park, 6-VI-1920, F. R. Cole, 9 6 2 2 (ANSP; CNC;. M@Z,;, UCB, WMinn: USNM); 9-VI-1922, 2 6 2 2 (MEI, UCB); 15-VI-1921, 4 6 (MEI, UCB); Golden Gate Dunes, 25-VI-1931, C. H. & D. Marrin, 2 6 (OSU); Lobos Creek, 10-V-1979, J. Pow- ell, 5 6 3 2 (UCB); Stanford University, 21- VI-1910, 1 ¢(USNM). Santa Clara County: 11 km S San Antonio Ranger Station, 27- VI-1953, G. A. Marsh, 1 @ (UCB); 11 km S San Antonio Ranger Station, 27-VI-1953, G. A. Marsh, | 2 (UCB); San Jose, 6-VIII- 1956, D. Ribble, 1 @ (SEM); Smith Creek Ranger Station, 640 m, 20-21-VII-1968, P. H. Arnaud, | 2 (CAS). Santa Cruz County: Felton, Santa Cruz Mountains, 91-152 m, 20-25-V-1907, Bradley, 5 65 2(CU). Shasta County: 21 km NW McArthur, 3-VI-1964, J. Schuh, 1 ¢(UCB), on Apocynum sp.; Jones Valley, Shasta Lake, 25-VIII-1959, W. G. Iltis, 1 ¢ 1 (UCD). Sierra County: Sattley, 13-VII-1962, R. M. Bohart, 2 2 (INHS, UCD); 26-VI-1964, 6 6 1 2 (INHS, UCD); 26-VI-1964, M. E. Irwin, 10 6 4 2 (INHS, MEI, UCD), swept from pasture of bloom- ing Daucus pusillus, 28-V1-1964, M. E. Ir- win, | 6(UCD); 26-VI-1964, C. R. Kovacic, 6 6(UCD); Sierra Valley, 3-VII-1980, L. S. Kimsey, | 6 (UA, UCD). Siskiyou County: 212 10 km S Macdoel, 2-VII-1956, J. Schuh, 1 6 1 2 (UCB). Solano County: Solano Lake, 14-V-1971, E. E. Grissell, 1 2 (UCD). Son- oma County: Windsor, 3-VIII-1956, A. M. Barnes, | ¢ (INHS); Cloverdale, 24-VIII- 1953, E. I. Schlinger, 1 @ (UCB). Stanislaus County: Del Puerto Canyon, Frank Rains Park, 335 m, 27-IX-1969, P. H. Arnaud, 1 2 (CAS). Trinity County: Hayfork, 22-V- 1973, J. Chemsak, 1 2? INHS, UCB); 23-V- 1972,246192(MEI, UCB), flight trap; Moun- tain Meadow Ranch, head of Coffee Creek, 1555 m, 8-10-VII-1969, C. Slobodchikoff, 1 6 (UCB); W. G. Goodman, | ¢ (UCD); Hayfork Ranger Station, 22-V-1973, J. Chemsak, 3 ¢ 4 2 (MEI, UCB), light trap. Ventura County: Wagon Road Number 2 Campground, 29 km WSW Gorman, 1585 m, 4-VII-1968, P. H. Arnaud, | ¢ (CAS); Lockwood Valley near Stauffer Post Office, 5-V-1959, P. D. Hurd, | 6 (UCB), on Pha- celia ciliata; 7-V-1959, 1 6 (UCB), on Des- curainia Sophia; 3-V-1959, C. W. O’Brien, 2 62 2(UCB), on Phacelia ciliata; J. Powell, 1 6(UCB). Yolo County: 13 km NW Win- ters, 10-VIII-1959, J. Fowler, 1 2 (UCD); 6 km SW Dunnigan, 8-IX-1959, 1 @ (UCD), light trap; Rumsey, 6-VIII-1955, E. A. Kurtz, 1 @ (UCD); Davis, 30-VI-1936, R. M. Bohart, 2 2? (UCD); 11-VII-1936, 2 ¢ (UCD); 3-VIII-1955, 2 ¢ (UCD); 14-VIII- 1969, 1 2 (UA); 11-VIHI-1982, D. G. Den- ning, 1 6 (CAS); 25-V-1962, M. E. Irwin, 1 2? (MEI); 27-V-1962, 1 2? (MEI); 4-VIII-1962, 1622(MEI); 11-VIII-1962, 2? (MEI, UCD); 4-VI-1965, C. R. Kovacic, | 4, 4-VITI-1965, 1 6 (UCD); 18-V-1950, A. T. McClay, 2 (UCD); 9-VI-1950, 2 (UCD); 22-VII-1956, 4 2 (UCD); 20-VIII-1955, 2 6 (UCD); 21- VIII-1955, 2 2 (UCD); 28-VII-1955, 2 6 (UCD); 2-IX-1956, 2 ¢ (INHS, UCD); 8-VITI-1953, W. D. McClellan, 2 6 (UCD); 18-VIII-1964, F. D. Parker, 2 2 (UCD); 26- IX-1965, 2 2 (INHS, UCD); 16-VII-1952, E. I. Schlinger, 2 6 (UCD); 17-VII-1959, 2 6 (UCD); 15-VII-1962, R. O. Schuster, 8 4 4 2 (UCD); 23-IV-1959, F. E. Strong, 2 6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (UCD); 21-VI-1959, 1 6 (UCD); 29-VI- 1959, 1 6 (UCD); 8-VIII-1959, 4 6 (UCD), light trap; 10-VIHI-1959, 2 ¢ (UCD); 23- VII-1959, 2 6 2 2 (UCD), light trap. IDA- HO. Eastern Idaho, 11-VII-31, D. E. F., 1 6 (USNM). Boise County: 14 km E Garden Valley, 3-VII-1979, M. W. Hanks, 2 6 (UIda). Camas County: 16 km NW Hill City, 15-VII-1967, A. R. Gittins, 1 ¢ 1 2 (Ulda), Lupinus sp. Fremont County: 11 km NW Anthony, 16-VII-1965, R. L. Westcott, 2 2 (UlIda). Gooding County: Wendell, 9-VI-33, Norta-Salsola, 1 ¢ (USNM); Bliss, 1000 m, 1-VI-1926, R. W. Haegele, 2 2 (UIda). Jef- ferson County: 7 km NW Terreton, 25-VII- 1957, W. F. Barr, 2 2 (UIda). Latah County: Idlers Rest Creek near Moscow, 12-VII- 1964, R. L. Westcott, 2 2 (UIda). Minidoka County: Adelaide, 14-VI-1929, 3 ¢ (UlIda). Oneida County: Stone Reservoir, 31-V- 1969, G. F. Knowlton, 1 ¢(USU); Pocatello Valley, 11-VI-1973, G. F. Knowlton, 1 4 (USU); 3.2 km S Roy Summit, 17-VIH-1972, G. F. Knowlton, | 2 (USU). Owyhee Coun- ty: 21 km S Grasmere, 7-VII-1969, A. R. Gittins, 2 @ (UIda); 27 km S Grasmere, 7-VII-1968, A. R. Gittins, 2 2? (UIda), Achil- lea Millefolium. Twin Falls County: Castle- ford, 28-VI-28, 1 ?(USNM); Castleford, So- phia #3, 15-VI-34, 1 2(USNM); Roseworth, 16-VII-1963, O. O. Fillmore and G. B. Hewitt, 2 6(UIda); 11 km S Kimberly, 8-VI- 1967, M. A. Brusven, 2 ° (UIda); Hollister, 13-VI-1931, D. E. Fox, 2 6 (MEI, UCD); 26-VI-1931, D. E. Fox, 1 ¢é (UCD). Valley County: Camp Creek, South Fork Salmon River, 14-VI-1966, W. F. Barr, 2 2 (UIda). Washington County: Midvale, 782 m, 20- VI-1926, R. W. Haegele, 2 ¢ (UIda). NE- VADA. Elko County: Carlin, 25-VI-1935, R. H. Beamer, 5 ¢ (INHS, SEM); 8 km NE Lamoille, 19-VI-1958, R. C. Bechtel, 2 6 (MEI, UCD); Harrison Summit, Ruby Mountain, 2208 m, 28-VI-1960, R. E. Blackwelder, J. C. Downey, 2 6 2 2 (UCD); 11 km S Carlin, 27-VI-1955, J. C. Downey, 2 612 (UCD); Wells, 12-VII-1911, J. M. VOLUME 97, NUMBER 1 Aldrich, 1 @ (USNM). Humboldt County: Orovada, 14-VII-1962, F. D. Parker, 2 6 2 ? (UCD); Winnemucca, 23-V-1960, T. R. Haig, 2 6 (UCB). Lander County: Austin Summit; 92255m, 27-Vi-1962, L. R: O’Brien, 2 2 (UCB) Ormsby County: Orms- by, 6-VII-, Baker, 1 ¢(USNM); Carson City, 26-VII-1929, E. P. Van Duzee, 1 62 9(CAS, UCB). Washoe County: Reno, 19-V-1963, R. C. Bechtel, 2 @ (UCD); Patrick, 16-VI- 1964, J. E. Slanky, 2 2 (MEI). OREGON. Critterion, 2-VII-1935, R. H. Beamer, 2 ¢ (SEM). Baker County: Sparta, 3-VII-1922, E. C. Van Dyke, 1 ¢ (UCB). Gilliam County: 3 km W Arlington, 29-VII-1964, J. W. MacSwain, 7 2 (MEI, UCB). Grant County: Seneca, 11-VII-1935, J. Schuh, 1 ¢ (CSU); Keerin’s Ranch, Izee, 25-VI-1935, J. Schuh, 1 @ (FSCA). Harney County: Antelope Mountain, 11-VII-1931, D. K. Frewing, 1 2 (OSM); Roaring Spring Ranch, Steens Mountains, 1370 m, 7-VII-1927, H. A. Scullen, 1 ¢(OSM). Josephine County: 6 km E Grants Pass, 17-VII-1969, R. L. Westcott, 1 6 (OSDA). Klamath County: Klamath Falls, 13-V-1924, C. L. Fox, 1 6 (MEI); 21- VII-1973, J. Schuh, 1 ¢ (FSCA); Worden, 1-VII-1935, R. H. Beamer, 2 2?(INHS, SEM); Algoma, Klamath Falls, 18-VII-1955, J. Schuh, 2 6 (UCB); 21-VIII-1973, 1 @ (FSCA). Malheur County: 3 km S Vale, 28-V-1969, K. Gordon, | 2 (OSDA), reseeded range. Wasco County: Shaniko, 17-VII-1935, J. Schuh, 1 2(FSCA); 13 km N Warm Springs, 2-VI-1968, K. Goeden, 1 6(OSDA). UTAH. Lampo, 22-V-1932, G. V. Knowlton, | 4 (SEM); 31-V-1939, 1 2; Locomotive Spring, 9-VI-1930, 1 6 (AMNH), Atriplex sp. Box Elder County: Snowville, 24-VI-1931, G. F. Knowlton, 1 @ (USU); 5-VI-1934, W. L. Thomas, | 2 (USU); 8 km W Snowville, 6-VI-1969, G. F. Knowlton, 1 @ (USU); Bothwell, 22-VI-1938, D. E. & H. T. Hardy, 1 2 (USU). Cache County: Hyrum, 22-VI- 1938, D. E: Harax, W. P. Nye, 1 2 (INHS); Logan, South Farm, 9-VII-1948, B. A. Haws, | 6(USNM). WASHINGTON. Ben- 213 ton County: 3 km W West Richland, 7-VI- 1973, N. E. Woodley, 1 2 (WSU). Klickitat County: 40 km N Goldendale, 26-VI-1969, R. L. Westcott, 1 2° (Ulda). Chromolepida clavitibia Webb and Irwin, NEw SPECIES Derivation of name.—c/ava (Latin, fe- male) = club; tibia (Latin) = shinbone. Diagnosis. — Males of Chromolepida clay- itibia differ from the males of all other spe- cies of Chromolepida in having dark brown halteres, dark reddish brown to black fem- ora, and the fore tibia distinctly clavate. Fe- males of Chromolepida clavitibia, like C. nigra, have dark brown to black halteres and femora. The species is separated from C. nigra in having the fore tibia distinctly clavate. Description of holotype male (88/2).— Body length 5.3 mm. Head: Length 0.9 mm. Ocellar tubercle black, pruinose; setae black, filiform. Eyes dark reddish brown; medial margin round- ed. Frons with medial callus dark reddish brown, glossy, dorsal and lateral pile silver; setae black, filiform. Antenna black, prui- nose; setae black, filiform, short; macrose- tae black; length of antenna 1.3 times head length; length of scape 0.56 mm, 5.6 times width, 4.7 times length of pedicel; length of pedicel 0.12 mm, 1.2 times width; length of flagellum 0.46 mm, 4.6 times width, 0.8 times length of scape; length of stylomere 0.04 mm. Parafacial dark reddish brown, glossy with silver pile along lateral margins. Maxillary palpus black; length 0.46 mm, 5.8 times width; setae white, filiform. Thorax: Macrosetae: np 3. Black, prui- nose; setae white and dark brown filiform and silver-green scale-like. Postpronotal se- tae white filiform and silver-green scale-like. Prosternal setae white filiform. Pleura dark reddish brown to black, pruinose, with pos- terior three-fourths of anepisternum glossy; setae pale yellow filiform on propleuron, anepisternum and dorsal third of katepi- 214 meron, silver-green scale-like on anepister- num and dorsal third of katepisternum. Scutellum black, pruinose; setae silver lan- ceolate in band across posterior margin. La- terotergite black, subshiny; setae white to pale yellow filiform. Wing. Length 4.0 mm, 3.3 times width; opaque, pale yellow, pale smoky black band across wing apex and apex of discal and posterior basal cells; ptero- stigma pale brown. Halter brown. Legs. Coxae dark reddish brown to black, prui- nose. Femora dark reddish brown to black, subshiny. Tibiae dark reddish brown; fore tibia distinctly clavate. Tarsi dark reddish brown. Coxal setae white filiform and sil- ver-green scale-like; femoral setae white fi- liform and silver-green scale-like; tibial se- tae dark reddish brown filiform. Abdomen: Tergites 1-2 dark brown, pru- inose, tergites 3-8 yellowish orange, sub- shiny; dorsal setae white lanceolate, densely appressed over tergites 1-2 and medially on tergite 3, lateral setae pale yellow filiform and silver-green scale-like. Terminalia (Figs. 12-18). Yellowish orange, subshiny. Ster- nite 8 moderately reduced, posterior margin moderately concave. Gonostylus with short basilateral projection. Dorsal apodemes of aedeagus broad, anterior margin deeply concave; ventral apodeme broad, anterior margin pointed; distiphallus elongate, sin- uate. Variation in males: Body measurements for males are provided in Table 3. Description of female.—Similar to male with following exceptions. Body measure- ments for females given in Table 3. Head: Frons black, pruinose with medial callus dark reddish brown, subshiny, with dorsolateral circle of black pile and narrow band of white pile lateral to antennal bases; setae black. Thorax: Macrosetae: np 3. Wing. Pale smoky brown, margin of veins darker; pter- ostigma dark brown. Abdomen: Dark reddish brown to black, subshiny; dorsal setae white, lanceolate, across posterior margin of tergites 1-3, lat- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON eral setae pale yellow filiform and silver- green scale-like on tergites 1-2. Terminalia (Figs. 19-21). Sternite 8 (Fig. 20) with pos- terior margin bilobed. Furca (Fig. 21) sub- rectangular; length 0.34 mm; posterior and anterior margins truncate; lateral margins parallel. Seasonal activity and distribution In the specimens examined, adults were collected during April, October, and De- cember. Chromolepida clavitibia has been collected only from the southern tip of Baja California Sur, México (Fig. 57). Specimens examined Type material.—The holotype male of Chromolepida clavitibia (deposited in the California Academy of Sciences, Type No. 17029, on permanent loan from the Uni- versity of California, Riverside) (M. E. Ir- win Therevidae Specimen Number 88/2) was collected at Los Frailes, Baja California Sur, MEXICO, by E. M. Fisher, on 25-26- IV-1975. Other material.—ALLOTYPE: In copulo with holotype, same data as holotype (CAS). PARATYPES: MEXICO. Baja California Sur. Los Frailes, 25-26-IV-1975, E. M. Fisher, 2 6 2 2 (INHS, UCR); 16 km NW La Paz, 6-X-1941, Ross, Bohart, | ¢ (CAS); La Paz, 19-XII-1973, W. Middlekauff, 1 2 (UCB). Chromolepida mexicana Cole Chromolepida mexicana Cole (1923: 460): Cole (1965: 350); Irwin and Lyneborg (1981la: 260). Derivation of name.— mexicana = of or from México. Diagnosis.— Males of Chromolepida mexicana, like C. nigra, have the disti- phallus elongate and bent at a right angle to the plane of the aedeagus. The males differ from C. nigra in having the apical half of the gonocoxite broad in lateral view. Fe- VOLUME 97, NUMBER 1 males of Chromolepida mexicana, like C. clavitibia and C. nigra, have dark brown to black halteres. The females are separated from C. clavitibia and C. nigra in having the femora dark yellow. Redescription of holotype female (9025).—Body length 7.0 mm. Head: Length 1.0 mm. Ocellar tubercle dark reddish brown, pruinose; setae black. Eyes dull reddish brown; medial margin rounded. Frons dark reddish brown, with glossy callus dorsad of antennal bases, pru- inose over remainder of frons, with silver, lateral pile and circular area of black pile dorsolateral to callus; setae black, scattered. Antenna reddish brown, pruinose; setae dark reddish brown; length of antenna 1.2 times head length; length of scape 0.66 mm, 6.6 times width, 4.4 times length of pedicel; length of pedicel 0.15 mm, 1.5 times width; length of flagellum 0.39 mm, 3.9 times width, 0.6 times length of scape; length of stylomere 0.04 mm. Parafacial dark reddish brown, glossy with silver pile along lateral margins. Maxillary palpus dark reddish brown; length 0.48 mm, 4.8 times width; setae white, filiform. Thorax: Macrosetae: np 3. Dark brown; setae white, filiform, short, scattered scale- like. Postpronotal setae white and scale-like. Prosternal setae white. Pleura dark reddish brown, pruinose, anepimeron glossy; setae white, filiform, on propleuron, anepister- num and dorsal third of katepisternum and silver, scale-like over entire anepisternum and dorsal third of katepisternum. Scutel- lum dark reddish brown, pruinose; setae white, lanceolate, in broad band across pos- terior margin. Laterotergite dark reddish brown; setae dark white. Wing. Length 4.7 mm, 3.1 times width; opaque, pale smoky brown with apical band, apex of discal and posterior basal cells, and along posterior margin of wing pale brown; pterostigma brown. Halter dark brown. Legs. Coxae dark reddish brown, pruinose. Femora dark yel- low, subshiny. Fore tibia dark reddish brown, middle and hind tibae dark yellow, 215 subshiny. Tarsi dark reddish brown. Setae white, filiform and, scale-like setae on fore coxa and femora, dark reddish brown, fili- form on tibae. Abdomen: Dark reddish brown, subshi- ny; dorsal setae scale-like, and silver, ap- pressed, lanceolate across posterior margin tergites 1-4, lateral setae white, filiform on tergites 1-2. Terminalia (Figs. 29-31). Ster- nite 8 (Fig. 30), posterior margin with deep central notch; apical fourth of sternite flat- tened. Furca (Fig. 31) subrectangular; length 0.4 mm; posterior and anterior margins broadly rounded; lateral margins parallel. Variation in females: Body measure- ments for females are provided in Table 3. Thorax. Macrosetae: np 2-3, 3, sa 1, pa 1, dc 0, sc 4. Description of male.—Similar to female with following exceptions. Body measure- ments for males given in Table 3. Head: Ocellar tubercle dark reddish brown. Frons dark reddish brown, glossy, with silver pile dorsad and laterad. Maxil- lary palpus dark brown, pruinose. Thorax: Macrosetae: np 3-4, 4. Vittae faintly separated by gray pruinose areas. Wings. Hyaline to pale yellow with apical band and apex of discal and posterior basal cells pale smoky brown; pterostigma slightly darker than membrane. Halter pale yellow, stalk darker yellow. Legs. Tibiae dark brown, pruinose. Length of fore femur to length of scape 1.8-2.9, 2.4 + 0.2 (N = 30); length of fore tibia to length of scape 2.0-3.1, 2.5 + 0.2 (N = 30); length of hind femur to length of scape 2.4-4.4, 3.5 + 0.4 (N = 30). Abdomen: Tergites 1-4 dark reddish brown, tergites 5-8 dark yellowish orange; setae white, filiform and dense, silver, lan- ceolate, appressed dorsally on tergites 1-4. Terminalia (Figs. 22-28). Dark yellowish orange. Sternite 8 moderately reduced, pos- terior margin slightly concave. Gonostylus with lateral hook-like projection. Dorsal apodeme of aedeagus broad anteriorly, an- terior margin deeply emarginate, ventral apodeme attenuate, pointed anteriorly; dis- 216 tiphallus moderately long, bent at right an- gle to plane of aedeagus. Seasonal activity and distribution In the specimens examined, adults were taken from 31 January to 29 September. Specimens have been collected on Acacia sp., Baccharis sp., B. glutinosa Pers., Bail- eya sp., B. pleniradiata Harr. and Gray, Chenopodium album L., Euphorbia albo- marginata T. and G., Happlopapus tenu- isectus (Greene) Blake, Lepidium Thurberi Woot, Senecio sp., Solanum sp., swept al- falfa, and from a dry wash. Chromolepida mexicana has been collected from northern México north to Arizona, southern Califor- nia, central Colorado, southwestern New Mexico, and western Texas (Fig. 57). Specimens examined Type material.— The holotype of Chrom- olepida mexicana Cole (CAS) (M. E. Irwin Therevidae Specimen Number 9025) was collected at Guaymas, Sonora, México, on 10 April 1921 by E. P. Van Duzee. Other material.— MEXICO. Chihuahua: 14.4 km W El Sueco, 1-VII-1972, R. Man- ga, T. Sluss, 1 ¢ (USNM); Chicuahua, 13- VII-1938, L. J. Lipovsky, 1 6 (SEM); 24 km S Camargo, 1310 m, 6-IX-1962, R. H. & E. M. Painter, | 6 (KSU). Sonora: Nogales, 28-ITI-1950, 1 ¢(USNM); 19 km N Ciudad Obregon, 24-IV-1961, R. H. & E. M. Paint- er, 1 2 (KSU); 132 km N Hermosillo, 7-II- 1964, M. E. Irwin, 1 2? (MEI); Guaymas, 10- IVal92 1 CE PS Vani Duzee,~1 “2 (GAS): UNITED STATES. ARIZONA. Southern Arizona, 30-VII-1959, F. M. Hull, 3 6192 (CNC). Cochise County: 32 km W Benson, 4-IX-1968, D. R. Miller, J. E. Lauck, 1 ¢ (UCD); Peloncillo Mountains, 16 km E Apache, 5-VIII-1972, R. F. Denno, K. Yeargan, J. R. Benedict, 1 ¢ (UCD); Carr Peak, Huachuca Mountains, 16-VIII-1966, R. L. Westcott, 2 2 (UIda); Douglas, 23-III- 1933, W. W. Jones, 1 ¢ (UCB); 2-IV-1933, 1 ¢ (UCB); 2 km E Wilcox, 26-VII-1973, J. D. Pinto, 1 ? (UCR), Chenopodium album; 8 km E Lowell, 15-VIII-1958, P. M. Marsh, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1 @ (UCD); Chiricahua Mountains, 5-IX- 1947, D. J. & J. N. Knull, 1 2 (UA); South- west Research Station, 8 km W Portal 28- IX-1965, V. D. Roth, 1 @ (MEI); 16 km E Sierra Vista, 15-IX-1974, A. E. & M. M. Michelbacher, 1 2 (UCB), Happlopapus ten- uisectus;, 4.8 km E Douglas, 1-IX-1984, R. M. Bohart, 1 2 (UCD); 5 km E Douglas, 1216 m, 12-VIII-1962, H. A. Scullen, 1 4 1 2 (OSU); San Bernardino Ranch, Douglas, 1143 m, -VIII-, F. H. Snow, 3 4 1 2 (INHS, SEM); 48 km NE Douglas, 1417 m, 1-VIII- 1946, H. A. Scullen, 1 ¢ (OSU); Skeleton Canyon, 10 km SE Apache, 1-IX-1958, E. G. Linsley, | 6 (UCB); 7 km W Montezuma Pass, 9-IX-1965, C. W. O’Brien, | ¢ (UCB); Bruno Canyon, Chiricahua Mountains, 1525 m, 28-VIII-1965, G. R. Ballmer, 1 ¢(UCR); 4 km NE Portal, 30-VIII-1959, E. G. Lins- ley, 1 6 (UCB), Acacia sp.; 3 km NE Portal, 2-4-VIII-1960, M. A. Cazier, 1 2 (AMNH); 17-I[X-1961, 1 ¢ (AMNH); 6 km E Portal, Chiricahua Mountains, 13-VIII-1965, G. R. Ballmer, 1 ¢ (UCR); 11 km E Portal, 23- VIII-1963, M. A. Cazier, 1 ¢ (UCB); 13- VIII-1962, J. Wilcox, 1 @ (UCB); 18-IX- 1962, 1 6 2 2 (UCB); Wilcox, 2-VIII-1975, J. D. Pinto, 1 ¢ (UCR); 18-VIII-1958, R. M. Bohart, 1 2 (UCD); 14-IX-1935, F. H. Parker, 1 6(USNM); Huachuca Mountains, 1 2 (USNM); 12-VIII-1950, D. J. & J. N. Knull, 2 6 1 (OSM); San Bernadino Ranch, 1145 m, F. H. Snow, 1 ? (SEM); 15.4 km N Apache, 14-VIII-1959, E. G. Linsley, 1 2 (UCB); 6 km S Apache, 1-IX-1959, H. E. Evans, 1 2 (CU), Baccharis glutinosa; 8 km S Apache, 11-VIII-1958, P. D. Hurd, 1 2 (UCB); 8 km SW Apache, 1311 m, 12-VIII- 1959, H. E. Evans, 1 2 (CU), Baccharis glu- tinosa; 19-VIII-1959, E. G. Linsley, 1 @ (UCB), Baccharis sp.; 14 km N Apache, 14- VIII-1959, E. G. Linsley, 2 ¢ (UCB); 21 km SW Apache, 24-VIII-1980, J. G. Rozen, 1 6 (AMNH); 2 km E Apache, 9-VIII-1973, J. G. Ehrenfeld, 2 2 (AMNH), Euphorbia albomarginata; 10-VIUI-1973, J. G. Ehren- feld, 1 2 (AMNH), Euphorbia albomargin- ata; Portal, 1525 m, 7-[X-1959, H. E. Evans, 1 6, 1 2 (CU). Gila County: base of Pinal VOLUME 97, NUMBER 1 Mountains, -III-, D. K. Duncan, | 4, 1 2 (FSCA); Mendoza Canyon, west slope Coy- ote Mountains, 1-VIII-1968, M. L. Lindsey, 1 (UA); Globe, 15-III-1936, F. H. Parker, 1 2 (UA); 25-III-1938, 2 ¢ (USNM); 2-IV- 1935, 1 6 1 ? (NHS, USNM)); 5-IV-1935, 3 6 3 2 (INHS, UA, USNM, WSU); 8-IV- 1935, 1 9(UA); 21-IV-1937, 1 9(UA). Mar- icopa County: Canon Lake, 2-IX-1935, F. H. Parker, | ¢ (USNM); Kyrene, 20-VI- 1956, G. D. Butler, 1 (UA); Theba, 2-VIII- 1954, G. D. Butler, 1 6 1 @ (INHS, UA), swept alfalfa; Chandler, 26-V-1955, G. D. Butler, 32 (UA), swept alfalfa; 2-VIII-1955, O. L. Barnes, 24 12 (UA), swept alfalfa. Pima County: Baboquivari Mountains, 28-IV- 1935, F. H. Parker, 1 ¢(USNM); Santa Rita Range Reserve, 1219 m, 12-VIII-1949, F. Werner, W. Nutting, 1 ¢ (USNM), mes- quite-desert grassland; Santa Cruz Village, Cobabi Mountains, 945 m, 10-12-VIII- 1916, 1 2 (CNC); Sasabe, 15-II-1997, O. Peck, 1 65 9(ASU, CNC, USU); Pena Blan- co, Arivaca, 3-I[X-1963, E. R. Burdien, | (USNM); Quijotoa, 28-VIII-1927, 1 ¢ (USNM); 48 km E Quiotoa, 28-29-VIII- 1927, 1 2 (CU); Santa Catalina Mountains, 19-VIII-1968, R. M. Bohart, 1 @ (UCD); 26 km E Tucson, -III-1935, J. A. Griswold, 1 2 (MCZ); 76 km SW Tucson, 20-III-1961, R.H. & E. M. Painter, 1 ¢(KSU), on flowers of Baileya sp.; Tucson, 5-III-1964, G. Dan- ila, 1 6 1 @ (USNM); 28-III-1967, D. M. Wood, | 2 (CNC); 25-III-1956, G. D. But- ler, 1 2, swept alfalfa; 28-VIII-, F. M. Car- penter, 1 2 (MCZ); 29-IX-1940, E. L. Pe- terson, 1 2 (UA); south of Tucson, 25-III- 1970, O. R. Taylor, 1 ?(SEM); Madera Can- yon, 18-VIII-1963, V. L. Vesterby, 2 2 (MEI, UCD); 2-[X-1973, L. Bezark, M. Ebertz, C. Katayama, | 2 (UCB); Schaeffer Cafion, Ba- boquivari Mountains, 1573-1676 m, 18-IX- 1924, R. & H., 1 6 (ANSP); 16 km NW Tucson, 732 m, R. H. & E. M. Painter, | ¢ (KSU); Baboquivari, 19-VII-1932, R. H. Beamer, | ¢ (SEM); 35 km E Ajo, 610 m, 31-III-1965, Bollinger, 2 ¢ (ASU, OSU); Rincon Mountains, 19-IX-1937, R. S. Beal, 1 6 (UA); Continental, 22-III-1956, F. W. 217 Werner, G. D. Butler, 1 ¢ 1 2 (INHS, UA), swept alfalfa; Sierrita Mountains, 915-1220 m, 10-VIII-1924, A. A. Nichol, 1 ¢ (UA); Brown’s Canyon, Baboquivari Mountains, 18-VIII-1956, G. D. Butler, F. G. Werner, 2 6 12 (UA); 21-VITI-1957, C. W. O’Brien, 1 6 (UA); Summerhaven, Santa Catalina Mountains, 2345 m, 20-VIII-1934, I. Moore, | 2?(SDNHM); 5 km N Madera Can- yon, 10-VIII-1979, G. Forbes, 1 2?(NMSU), in mesquite wash; 2 km W Robles Junction, 26-VII-1973, E. M. Fisher, 2 ¢ 2 ° (UCR); Box Canyon, 25-VIII-1975, L. Bezark, G. Nishida, C. Kitayama, B. Tilden, | @ (UCB); 10-VIII-1977, D. K. Faulkner, 1 64 (SDNHM). Pinal County: Aravaipa Can- yon, 9-IX-1986, F. Parker, T. Griswold, 1 2 (USU); Coolidge, 19-VII-1956, C. Wil- liams, 1 6 (INHS); Casa Grande, 26-V-1955, G. D. Butler, 2 ¢ (UA), swept alfalfa; 20- VI-1956, 1 ¢ (UA), swept alfalfa; Oracle, 1370 m, 11-VIII-1950, R. S. Beal, 1 ¢(UCB); 25-VIII-1934, I. Moore, 1 ¢ 1 2 (SDNHM). Santa Cruz: Parkers Ranch, Santa Rita Mountains, 1981 m, 21-VII-1937, H. Ruckes, | 2 (USNM); 19 km N Nogales, 1-VIII-1966, C. R. Kovacic, 1 @ (UCD); Santa Rita Range Reserve, 1219 m, 12-VIII- 1949, F. G. Werner, W. Nutting, 1 ¢ (USNM), mesquite-desert grassland; 8 km E Nogales, 1-IX-1970, G. E. & R. M. Bo- hart, 1 6 1 2(UCD, USU); Santa Rita Moun- tains, 6-IV-1937, W. Benedict, | 62 9 (SEM); 1-VIII-1941, R. H. Beamer, 1 ¢ (SEM); Cal- abasas, 9.6 km W Nogales, 1-VIII-1961, F. G. Werner, W. Nutting, 1 ¢(INHS); Madera Canyon, 12-VIII-1965, D. N. Harrington, 1 6 (MEI); 3 km SW Patagonia, 9-III-1963, J. C. Bequaert, 1 2 (UA); Patagonia Moun- tains, 7-VIII-1950, D. J. & J. N. Knull, 1 ¢ (OSM); Patagonia, 22-III-1956, G. D. But- ler, F. G. Werner, 1 2 (WSU), Senecio sp.; 19-IV-1956, G. D. Butler, 1 2 (UA), swept alfalfa; 2-VIII-1924, E.P. Van Duzee, 19 (UCB); 5-VIII-1937, R. S. Beal, 1¢ (UA); 5 km W Sonoita, 15-VIII-1966, R. L. West- cott, 1 6, 1 @ (UIda). Yavapai County: Cot- tonwood, 29-II-1978, R. C. Miller, 1 ¢ (UCD). CALIFORNIA. San Diego County: 218 Campo, 18-VII-1940, R. H. Beamer, 1 ¢ 1 ? (INHS). COLORADO. El Paso County: Foster Ranch, T15S R65W Sec 23 N'A, 1737 m, F. M. Brown, 1 2 (CSU). NEW MEXI- CO. Grant County: Silver City, 10-III-1934, R. T. Kellog, 3 6 2 2? (MEI, OSM). Hildago County: Skeleton Canyon, Rodeo, 10-VIII- 1955, R. R. Dreisbach, | 6, Skeleton Can- yon, 16-VIII-1968, J. B. Heppner, 1 3 1 2 (MEI), sweeping Lepidium Thurberi; 4 km N Rodeo, 23-VIII-1958, E. G. Linsley, 1 2 (UCB), Baileya pleniradiata; Rodeo, 1219 m, 21-VIII-1958, P. M. Marsh, | 62 9 (MEI, UCD); 28-VIII-1959, H. E. Evans, 1 ¢3 ¢ (CU, USNM); 31-VII-1951, 1 ¢ (USNM); 5-IX-1959, 1 (CU); 5 km NW Rodeo, 16- IX-1960, M. A. Cazier, 1 ¢(AMNH); 2 km N Rodeo, 29-VII-1959, E. G. Linsley, 2 6 (UCB); 16-VIII-1963, M. A. Cazier, Mor- tenson, | 6 1 2 (MEI); Cotton City, 19-VITI- 1979, R. M. Bohart, 1 2 (UCD). TEXAS. Jeff Davis County: Fort Davis, 22-VI-1947, R. H. Beamer, | 2 (INHS). Reeves County: Texas A & M Research Station, Pecos, 17- VITI-1976, S. J. Merritt, 1 9 (TAMU), sweeping Solanum sp. Chromolepida nigra Webb and Irwin, NEw SPECIES Derivation of name.—nigra (Latin) = black. Diagnosis.— Males of C. nigra, like C. mexicana, have the distiphallus moderately long and curved and the costal and subcos- tal cells of the wing hyaline. The males are separated from C. mexicana in having the distal half of the gonocoxite reduced and tapered apically in lateral view. Females of Chromolepida nigra, like C. mexicana and C. clavitibia, have dark brown to black hal- teres. The females may be separated from C. mexicana in having the femora dark brown to black and from C. clavitibia in having the fore tibia cylindrical rather than distinctly clavate. Description of holotype male (88/5).— Body length 4.7 mm. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Head: Length 0.9 mm. Ocellar tubercle black, pruinose; setae black. Eyes dark red- dish brown; medial margin rounded. Frons with medial callus black, glossy, surrounded by narrow band of silver pile; setae dark brown. Antenna dark brown, pruinose; se- tae black; length of antenna 1.3 times head length; length of scape 0.7 mm, 7.9 times width, 4.7 times length of pedicel; length of pedicel 0.15 mm, 1.4 times width; length of flagellum 0.37 mm, 3.4 times width, 0.5 times length of scape; length of stylomere 0.06 mm. Parafacial dark reddish brown, glossy with silver pile along lateral margins. Maxillary palpus dark reddish brown, pru- inose; length 0.37 mm, 7.4 times width; se- tae white. Thorax: Macrosetae: np 3. Black, prui- nose; pronotal setae pale yellow, filiform and scale-like. Postpronotal setae pale yellow, filiform and scale-like. Prosternal setae white. Pleura dark reddish brown, pruinose; setae white, filiform on propleuron, anepi- sternum and dorsal third of katepisternum, and scale-like on propleuron, anepister- num, and dorsal third of katepisternum. Scutellum black; setae silver, lanceolate, in band across posterior margin. Laterotergite dark reddish brown, subshiny; setae pale yellow. Wing. Length 3.6 mm, 2.9 times width; dark yellow, opaque, with smoky brown apical band, and at apex of discal and posterior basal cells; pterostigma ab- sent. Halter yellow. Legs. Coxae, tibiae, and tarsi dark reddish brown, pruinose; femora dark yellow, base dark reddish brown, sub- shiny. Length of fore femur 1.2 mm, 1.6 times length of scape; length of fore tibia 1.2 mm, 1.7 times length of scape; length of hind femur 1.5 mm, 2.1 times length of scape. Setae white, filiform, and scale-like on coxae and femora; dark reddish brown, filiform on tibiae. Abdomen: Tergites 1-3 dark reddish brown, pruinose; tergites 4-7 yellowish or- ange, subshiny; dorsal setae silver, lanceo- late on tergites 1-3, lateral setae white, fi- liform on tergites 1-2 and scale-like. VOLUME 97, NUMBER 1 Terminalia (Figs. 32-38) yellowish orange, subshiny. Sternite 8 moderately reduced, posterior margin slightly concave. Gono- stylus with lateral hook-like projection. Dorsal apodeme of aedeagus broad anteri- orly, anterior edge deeply emarginate, ven- tral apodeme large, attenuate, pointed an- teriorly; distiphallus moderately long, bent at right angle to plane of aedeagus. Variation in males: Body measurements for males are provided in Table 3. Length of fore femur to length of scape 1.4—2.4, 1.7 + 0.2 (N = 20); length of fore tibia to length of scape 1.5-2.4, 1.8 + 0.2 (N = 20); length of hind femur to length of scape 2.0-3.3, 2.4 + 0.3 (N = 20). In some males the pos- terior margin of sternite 8 is more deeply emarginate than normal; the lateral projec- tion on the gonostylus is generally slender, although it is short and hook-like in some specimens; and in specimen (8825) the an- terior margin of the ventral apodeme of the aedeagus is broadly rounded, rather than attenuated and pointed. Description of female.—Similar to male with following exceptions. Body measure- ments for females given in Table 3. Head: Ocellar tubercle dark reddish brown, pruinose; setae dark brown. Frons with medial callus black, glossy, with dor- solateral circle of black pile and narrow band of silver pile lateral to antennal bases; setae absent. Maxillary palpus dark reddish brown to dark brown, pruinose. Thorax: Macrosetae: np 3. Dark brown to black, pruinose; setae white and black, filiform, short and scale-like. Pleura dark reddish brown, subshiny, propleuron, kat- episternum, and meron pruinose. Wing. Pterostigma brown. Halter black. Legs. Dark reddish brown, pruinose. Abdomen: Dark reddish brown, subshi- ny; dorsal setae silver, lanceolate, across posterior margin of tergite 1, lateral setae white, filiform on tergites 1—2 and scale-like. Terminalia (Figs. 29-31). Sternite (Fig. 30) with narrow, deep median notch posteri- orly. Furca (Fig. 31) oval length 0.38 mm; 219 posterior margin truncate; anterior margins rounded; lateral margins curved. Seasonal activity and distribution In the specimens examined, adults were collected throughout the year, only from the southern half of México (Fig. 57). Specimens examined Type material.—The holotype male of Chromolepida nigra Webb and Irwin (de- posited in the Illinois Natural History Sur- vey) (M. E. Irwin Therevidae Specimen Number 8815) was collected 34 km E Villa Union on | February 1964 by M. E. Irwin. Other material.—PARATYPES: MEXI- CO. Hildalgo: Zimpan, 8-VII-1968, M. W. Wasbauer, J. E. Slansky, 1 2(UCD). Jalisco: Jocotepec, 12-13-IIJ-1972, R. Lavigne, 1 2 (UWyo); 19-III-1972, R. Lavigne, 1 2 (UWyo); Chamela, 26-30-IX-1985, F. D. Parker, T. L. Griswold, 1 @ (USU); 18 km SE Tequila, 20-XII-1963, M. J. Tauber, C. A. Toschi, 1 @ (UCB); Plan de Barrancas, 24-III-1962, F. D. Parker, 2 ? (UCD). Mo- relos: Canon de Lobos, Yautepec, 1219 m, 7-III-1959, H. E. Evans, 2 2 (CU); Xochi- calco Pyramid, 29-IJI-1962, F. D. Parker, L; A; Stange, 2 ¢ (MEI, UCD): Yautepec; 28-III-1962, F. D. Parker, L. A. Stange, 1 6 (UCD). Oaxaca: 37 km S Matias Romero, 6-IV-1962, F. D. Parker, 1 ¢ (MEI). Puebla: 5 km NW. Petlalcingo, 2-IV-1962, L. A. Stange, 1 ¢ (UCD); 4-III-1972, F. D. Parker, D.R. Miller, 11 646 2?(INHS, MEI, USNM); 14km W Izucar de Matamoros, 16-IX-1972, W. Hanson, J. Poff, 1 6 (MEI). Queretaro: Alvarado [Queretaro], 15-II-1954, R. R. Dreisbach, | @ (MEI). Sinaloa: 18 km N Culiacan, 20-V-1962, F. D. Parker, 1 92 (UCD); 34 km E Villa Union, 1-II-1964, E. I. Schlinger, 1 ¢ (UCR); 87 km S Culiacan, 185 m, 23-IV-1969, M. E. Irwin, | 6 (MEI), dry wash; 21 km N Mazatlan, 30-I-1964, M.E. Irwin, | 6 (MEI); 24 km S Villa Union, 31-I-1964, 1 2 (MEI); 31 km S Villa Union, 31-I-1964, M. E. Irwin, 2 6 (MEI). Yucatan: 220 Puerto Sisal, 25-VII-1964, J. C. & D. Pal- lister, 1 2 (AMNH). Chromolepida pruinosa (Coquillett) Psilocephala pruinosa Coquillett (1904: 91): Irwin and Lyneborg (1981a: 260). Chromolepida pruinosa (Coquillett); Irwin and Lyneborg (1981la: 260). Derivation of name.—pruina (Latin) = hoarfrost, rime. Diagnosis.—The males of Chromolepida pruinosa, like C. mexicana and C. nigra, have black frontal setae, the wing mem- brane with brown to black band across apex of wing and apex of discal and posterior basal cells and the gonostylus with a lateral projection. The males of C. pruinosa can be separated from C. mexicana and C. nigra in having the costal and subcostal cells of the wing dark yellow and the distiphallus very short. Females of Chromolepida prui- nosa, like C. bella, have dark yellow hal- teres. The females may be separated from C. bella in having a smoky brown to black band across the apex of the wing and the apex of the discal and posterior basal cells. Redescription of holotype male (92/8).— Body length 5.7 mm. Head: Length 0.9 mm. Ocellar tubercle black, pruinose; setae black. Eyes dark red- dish brown; medial margin sinuate. Frons with medial callus black, glossy, dorsal and lateral pile silver; setae black. Antenna (Fig. 42) dark yellowish brown, flagellum dark brown; setae black; length of antenna 1.1 times head length; length of scape 0.5 mm, 5.0 times width, 4.2 times length of pedicel; length of pedicel 0.12 mm, 1.2 times width; length of flagellum 0.34 mm, 2.8 times width, 0.7 times length of scape; length of stylomere 0.06 mm. Parafacial black, glossy with silver pile along lateral margins. Max- illary palpus (Fig. 43) black; length 0.46 mm, 5.8 times width; setae white. Thorax: Macrosetae: np 3. Dark black, subshiny; setae dull yellow, filiform and scale-like. Postpronotal setae white, filiform PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and scale-like. Prosternal setae white. Pleu- ra black, glossy, propleuron and posterior two-thirds of katepisternum pruinose; setae whitish yellow, filiform on propleuron, an- episternum and dorsal third of katepister- num, scale-like on propleuron, anepister- num, and dorsal third of katepisternum. Scutellum black, posterior margin pruinose; setae silver, appressed, lanceolate in band across posterior margin. Laterotergite black, subshiny; setae white. Wing (Fig. 44). Length 4.2 mm, 3.2 times width; pale yeliow, with smoky brown to black apical band, basal fourth of cell r;, and apex of discal and pos- terior basal cells smoky brown, costal and subcostal cells dark yellow; anterior veins dark yellow, posterior veins brown; pter- ostigma pale brown. Halter dark yellow. Legs. Coxae black, pruinose. Femora dark yellow, subshiny. Tibiae dull brown. Tarsi dark brown. Setae white, filiform and scale- like setae on coxae and femora, dark reddish brown, filiform on tibiae. Abdomen: Tergites 1-5 black, tergites 6— 8 dark yellow, subshiny; dorsal setae white, lanceolate, dense, appressed over tergites |- 5, lateral setae white, filiform and scale-like. Terminalia (Figs. 45-51). Dark yellowish brown, subshiny. Sternite 8 moderately re- duced, posterior margin moderately con- cave. Gonostylus with thin lateral projec- tion. Dorsal apodemes of aedeagus broad anteriorly, anterior margin truncate; ventral apodeme broad, anterior margin broadly rounded, anterolateral border with ventral projection; distiphallus short, reduced, sin- uate. Variation in males: Body measurements for males are provided in Table 3. Description of female.—Similar to male with following exceptions. Body measure- ments for females given in Table 3. Head: Ocellar setae absent. Frons (Fig. 52) with black, subshiny, medial callus with dorsolateral circle of black pile and narrow band of white pile lateral to antennal bases; setae absent. Parafacial callus broader than in male. VOLUME 97, NUMBER 1 Thorax: Macrosetae: np 3. Wing. Mem- brane pale, opaque brown, much darker than in males. Abdomen: Black, subshiny; dorsal setae whitish gray, filiform across posterior mar- gin of tergites 2-3, black, filiform, moder- ately long on tergites 4-8 black, and scale- like on tergites 1-3, lateral setae white, fi- liform on tergites 1-3, black, filiform on ter- gites 3-8. Terminalia (Figs. 53-55). Sternite 8 (Fig. 54) rather long and narrow, posterior margin with deep median notch posteriorly. Furca (Fig. 55) subrectangular; length 0.38 mm; posterior and anterior margins broadly rounded; lateral margins rounded. Seasonal activity and distribution In the specimens examined, adults were collected from 8 October to 6 May with the majority of specimens collected during Feb- ruary, March, and April. Specimens have been collected on bean leaves, but most specimens were collected by Malaise traps. Chromolepida pruinosa has been collected from Nicaragua south to Colombia and east to Guyana (Fig. 58). Specimens examined Type material.—The holotype male of Psilocephala pruinosa Coquillett (U.S. Mu- seum of Natural History, Type No. 7795) (M. E. Irwin Therevidae Specimen Number 9218) was collected at Granada, Nicaragua by Baker. Other material. -COLOMBIA. Magda- lena. 3 km E Santa Marta, 1-VIII-1973, P. A. Rauch, | @ (MEI); Santa Marta, 8-X- 1OWAGNG. FE: Bohart,1.2.(INHS). ‘COSTA RICA. Alajuela. Biyyagua, 7-IIJ-1991, F. D. Parker, | 2 (DEI). Guanacaste. 10 km SW San Miguel, 21-II-1991, D. W. Webb, M. E. Irwin, 11 6 16 g (CSDA, CSIRO, DZSA, FMNH, INHS, INIA, IOC, LACM, MEI, MPM, MSU, NSDA, PSU, SDNHM, UBC, UCVM),; 22-IT-1991, 20 46 192(CNC, DZSA, EEA, IML, INHS, IOC, LMM, MEI, UG, UNLP, WSU); 23-II-1991, 6 4 16 2 (INHS, 221 MEI); 25-II-1991, F. D. Parker, 1 ¢ (INHS); 26-II-1991, 1 2 (INHS); 27-II-1991,232¢ (INHS); 28-II-1991, 3 6 1 @ (INHS); 1-III- 1991, 1 6 1 e@ (CSDA, INHS); 10-IIT-1991, 6 69 2(CNC, FMNH, INHS, MEI); 12-IT- 1991, 1 6 1 9 (INHS, INIA); 13-III-1991, 3 6 5 2 (BMH, CSDA, INHS, IRSN, NHW, NMP, NRS, PAS, UG); 14-17-III-1991, 8 6 7 2 (ASU, BMNH, BYU, INHS, MCZ, MNH, MNHN, NMB, SMN, UTA, UZM, ZIR); 18-III-1991, 1 6 1 2 (UCM, UCR); 19-20-III-1991,6332(AMNH, ASU, BYU, CAS, CU, MCZ, NMB, UCM, UTA); 21- 22-III-1991, 7 6 4 2 (ANSP, CAS, INHS, UCB, UCR, Ulda, UWisc); 23-24-III-1991, 3 62 29 (INHS, OSM, SEM, UA); 25-27-III- 1991, 8 649; 29-ITI-1991, 1 6 4 2 (FSCA, INHS, IowaSU, KSU, NMSU, OSDA, OSU, TAMU, UA, UCYM, UMinn, UWyo); 31-III-1991, 2 6 8 e (CUS, FSCA, INHS, IowaSU, KSU, NMSU, OSDA, SEM, UCVM); 1-IV-1991, 3 2 (EI, INHS, RNHL); 2-IV-1991, 3 6 1 @ (DEI, DSIR, MNHN, NMP); 3-IV-1991, 2 6 (IRSN, PAS); 4-5-IV-1991, 1 6 3 2 (BMNH, CSI- RO, INHS, MNH); 6-IV-1991, 1 6 1 2 (MMB, NRS); 7-IV-1991, 1 6 1 2 (RNHL, SMN); 8-12-IN-1991, 1 ¢2°(EI, UTI, ZSI); 15-IV-1991, 1 6 1 ? (AMS, MMB); 17-IV- 1991, 1 ¢ (IAS); 19-IV-1991, 3 6 1 2 (AMS, IAS, KIK, ZIR); 19-20-III-1991, 1 9; 22- IV-1991, 2 ?(KUK, UZM); 24-IV-1991, 1 6 (ZSI); 2-V-1991, 1 6 (UTI); Palo Verde National Park, 34 km SW Canas, 2-II-1984, S. A. Cameron, | 2 (SEM); South of Canas, 26-31-I-1989, W. L. Rubink, 3 65 2@(USU); 1-3-II-1989, 2 6 1 2 (USU); 9-14-II-1989, 4 63 2? (INHS, USU); 7-10-III-1989, 1 3 1 2? (INHS, USU); 21-25-I-1989, F. D. Parker, 2642 (INHS, USU); 25-31-I-1989, 5 ¢ 1 2? (INHS, USU); 9-14-II-1989, 3 68 2 (INHS, USU); 16-20-II-1989, 4 64 2 (INHS, USU); 22-24-II-1989, 2 6 1 2 (USU); 25-II-8-III- 1989, 1 9(USU); 25-28-II-1989, 1 9 (INHS); 1-5-1989, 1 6 2 2 (INHS, USU); 7-10-III- 1989, 3 6 2 2 (USU); 18-22-III-1989, 1 9 (USU); 4-6-V-1989, 2 2 (INHS); La Taboga Forest Reserve, 9 km SW Cajfias, 17-27-II- 222 1987, W. L. Rubink, 3 6 3 2 (INHS, USU); 18-20-III-1987, 1 ¢ (USU); 17-27-II-1987, 1 69 2 (INHS, USU); 14 km S Cajias, 11- 31-1 1990, F.. D:: Parkers Ic 1-e(INES, USU); 1-11-II-1990, 1065 ¢ (INHS, USU); 15-22-I-1990, 1 ¢ (USU); 15-24-II-1990, 1 2? (INHS); 1-12-III-1990, 1 ¢ (DSIR); South Canas Experiment Station, 8-18-III-1988, F. D,; Parker, o71"2 (OSU): E:. JN. Agri- cultural Station, 11 km S Canas, 11-31-I- 1990, F. D. Parker, 1 2? (USU); 14-ITI-1990, 1 2 (USU); 10 km S, 10 km W Canas, Ta- boga 10° 19’ N, 85° 09’ W, 3-12-II-1967, 1 ? (USU); Hacienda La Pacifica, 6 km N Cafias, 20-21-III-1987, R. W. Thorp, 1 ¢ (UCD); 22-23-III-1987, 1 6 1 ¢(UCD). San Jose. Escazu, 24-30-I-1988, F. D. Parker, 1 6 1 9 (USU); 1-7-II-1988, 4 6 4 2 (INHS, USU); 7-14-II-1988, 2 6 7 2 (INHS, USU); 14-21-II-1988, 1 9(USU); 22-II-3-II-1988, 3 6 1 2 (USU); 3-III-1988, 1 ¢ (USU); 4-7- III-1988, 1 ¢ 3 2 (USU); 8-18-III-1988, 2 ¢ (USU); 21-27-III-1988, 1 ¢ (BMH); 19-24- III-1988, 1 @(USU); 26-ITI-1988, 1 ¢(USU); 29-III-3-IV-1988, 1 63 2 (INHS, USU); 1-8- IV-1988, 1 6 5 2 (INHS, USU); 4-10-IV- 1988, 1 6 2 2 (USU); 11-18-IV-1988, 1 ¢ (USU); 16-18-IV-1988, 1 6 (USU); 19-24- IV-1988, 1 ¢(USU); 25-30-IV-1988, 1 412 2 (INHS, USU); 2-13-V-1988, 4 2 (INHS, USU),; 23-26-VI-1989, 1 6 3 2 (USU); 8-II- 1987, G. E. Bohart, 1 @ (USU); San Pedro de Montes de Oca, -V-1987, M. Garcia, 1 2 (INHS); San Antonio de Escazu, -IV-1987, W. Everhard, 1 2? (INHS). GUYANA. Up- per Rupupanni, -IJ-III-, Ogilvie, 1 ?(MCZ). NICARAGUA. Granada, Baker, 1 6 (USNM); La Calera, Managua, 1 3-III-1964, L. Saenz, 1 @ (USNM), on bean leaves. VENEZUELA. Aragua: Pozo Diablo Creek, Maracay, 500 m, 10-IV-1961, M. Gelbez, 1 2(UCVM); El Limon, 450 m, 24-IV-1963, E. Osuma, | ¢ (MEI); Puerto de Cata, 10- 11-VI-1976, A. S. Menke, D. Vincent, 1 3 2 2 (MEI, USNM); 2 km N Ocumero de La Costa, 12-VI-1976, A. S. Menke, D. Vin- cent, 1 6 (USNM). Guarico: 12 km S Cal- abozo, 6-12-II-1969, P. & P. Spangler, 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (USNM), blacklight. Zulia: 6 km W La Con- cepcion, 18-VI-1976, A. S. Menke, D. Vin- cent, 1 ¢(USNM); Carrasquero, 15-VI-1976, A. S. Menke, D. Vincent, 1 2 (USNM). ACKNOWLEDGMENTS We thank Drs. G. L. Godfrey, L. M. Page, and B. A. Steinly for reviewing this manu- script. The following institutions and their current or former curators were kind enough to loan us material relevant to this study: Academy of Natural Sciences of Philadel- phia, D. Azuma (ANSP); American Muse- um of Natural History, D. A. Grimaldi (AMNH); Arizona State University, M. A. Cazier (ASU); California Academy of Sci- ences, P. H. Arnaud, Jr. (CAS); Canadian National Collection, H. J. Teskey, D. M. Wood, and J. M. Cumming (CNC); Colo- rado State University, B. C. Kondratieff (CSU); Cornell University, L. L. Pechuman, E. R. Hoebeke (CU); Florida State Collec- tion of Arthropods, H. V. Weems, Jr. (FSCA); Illinois Natural History Survey (INHS); Kansas State University, D. Block- er (KSU); Universidad Central de Vene- zuela, Maracay (UCVM); Michael E. Irwin Collection (MEI); Museum of Comparative Zoology, Harvard University, D. G. Furth (MCZ); New Mexico State University, J. R. Zimmerman (NMSU); Ohio State Museum, C. A. Triplehorn (OSM); Oregon State De- partment of Agriculture, R. L. Westcott (OSDA); Oregon State University, J. D. Lattin (OSU); San Diego Natural History Museum, D. Faulkner (SDNHM);- E. I. Schlinger Collection (EIS); Texas A & M University, E. G. Riley (TAMU); United States National Museum, F. C. Thompson (USNM); University of Arizona, F. G. Wer- ner (UA); University of California, Berke- ley/California Insect Survey Collection, J. Powell, (UCB); University of California, Davis, S. Heydon (UCD); University of California, Riverside, S. Frommer (UCR); University of Colorado, M. J. Weissmann (UCM); University of Idaho, W. F. Barr Entomological Museum, F. W. Merickel VOLUME 97, NUMBER 1 (Ulda); University of Kansas, Snow Ento- mological Museum, G. W. Byers (SEM); University of Minnesota, P. J. Clausen (UMinn); University of Wisconsin, L. E. Bayer, B. J. Harrington (U Wisc); University of Wyoming, S. R. Shaw, R. Lavigne (UWyo); Utah State University, W. J. Han- son (USU); Washington State University, James Entomological Collection, W. J. Tur- ner, R. S. Zack (WSU). Support for this study was provided by the Illinois Natural History Survey, the University of Illinois at Ur- bana-Champaign and the Schlinger Foun- dation. In addition to the museums and collec- tions from which material was borrowed, specimens of Chromolepida pruinosa have been deposited in the following museums and collections: AFRICA AND NEAR EAST: Natal Mu- seum, Pietermaritzburg, South Africa (NMP); University of Tel Aviv, Israel (UTA). ASIA: Institute of Agricultural Sci- ences, Tokyo, Japan (IAS); Kagoshima Uni- versity, Kagoshima, Japan (KUK); Zoolog- ical Survey of India Collection, Calcutta, India (ZSI). AUSTRALASIAN & OCE- ANIA: Australian Museum, Sydney (AMS); Australian National Insect Collection, Commonwealth Scientific and Industrial Research Organization, Canberra, Australia (CSIRO); Bernice P. Bishop Museum, Ho- nolulu, Hawaii (BMH); Department of Sci- entific and Industrial Research, Auckland, New Zealand (DSIR). EUROPE: Deutsches Entomologisches Institut, Berlin, Germany (DEI); Institut Royal des Sciences Naturelle de Belgique, Brussels (IRSN); Istituto di En- tomologia, Bologna, Italy (EI); Moravske Museum, Brno, Czechoslovakia (MMB); Musei Nationalis Hungarici, Budapest, Hungary (MNH); Museum National d’His- toire Naturelle, Paris, France (MNHN); Natural History Museum, London, England (NHML); Naturhistorisches Museum, Ba- sel, Switzerland (NMB); Naturhistorisches Museum, Wien, Austria (NHW); Naturhis- toriska Riksmuseet, Stockholm, Sweden 223 (NRS); Polish Academy of Sciences, War- saw (PAS); Rijkmuseum van Natuurlijke Historie, Leiden, The Netherlands (RNHL); Staatliches Museum fiir Naturkunde, Stutt- gart, Germany (SMN); Universita di Tori- no, Italy (UTI); Universitets Zoologisk Mu- seum, Copenhagen, Denmark (UZM); Zoological Institute, St. Petersburg, Russia (ZIR); Zoologisches Institut, Berlin, Ger- many (ZIB). NORTH AMERICA: Brigham Young University, Provo, Utah (BYU); California State Department of Agriculture, Sacramento (CSDA); Field Museum of Nat- ural History, Chicago, Illinois (FMNH); In- stituto Nacional de Biodiversidad, Heredia, Costa Rica (INBio); Instituto Nacional de Investigaciones Agricolas, Chapingo, Méx- ico (INIA); Michigan State University, East Lansing (MSU); Milwaukee Public Muse- um, Wisconsin (MPM); Natural History Museum of Los Angeles County, Los An- geles, California (LACM); Nevada State De- partment of Agriculture, Reno (NSDA); Pennsylvania State University, Frost En- tomological Museum, State College (PSU); University of British Columbia, Vancou- ver, Canada (UBC); University of Georgia, Athens (UG). SOUTH AMERICA: Insti- tuto Miguel Lillo, Tucuman, Argentina (IML); Museo de Ciencias Naturales, Uni- versidad Nacional de La Plata, Argentina (UNLP); Departamento de Zoologia Agri- cultura, Sao Paulo, Brasil (DZSA); Instituto Oswaldo Cruz, Rio de Janeiro, Brasil (IOC); Estacion Experimental Argonomica, Uni- versidad de Chile, Maipu (EEA); Coleccién Entomologica “Luis Maria Murillo,” Ti- baitata, Bogata, Colombia (LMM). LITERATURE CITED Cole, F. R. 1923. A revision of the North American two-winged flies of the family Therevidae. Pro- ceedings of the United States National Museum 62(4): 1-140. . 1965. Family Therevidae, pp. 348-354. In Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson, eds., A Catalog of the Diptera of America North of Mexico. Agricultural Handbook 276: 1-1696. 224 Coquillett, D. W. 1904. New Diptera from Central America. Proceedings of the Entomological So- ciety of Washington 6: 90-98. Irwin, M.E. 1976. Morphology of the terminalia and known ovipositing behaviour of female Therevi- dae (Diptera: Asiloidea), with an account of cor- related adaptations and comments on phyloge- netic relationships. Annals of the Natal Museum 22: 913-935. . 1977a. Two new genera and four new species of the Pherocera-group from western North Amer- ica, with observations on habitats and behavior (Diptera: Therevidae: Phycinae). Proceedings of the Entomological Society of Washington 79: 422- 451. 1977b. A new genus and species of stiletto- flies from southwestern North America with close affinities to Chilean and Australian genera (Dip- tera: Therevidae: Therevinae). Pan-Pacific Ento- mologist 58: 287-296. . 1983. The boharti species group of the genus Pherocera (Diptera: Therevidae: Phycinae). Pan- Pacific Entomologist 59: 113-139. Irwin, M. E. and L. Lyneborg. 1981a[1980]. The gen- era of Nearctic Therevidae. Bulletin of the Illinois Natural History Survey 32: 193-277. 1981b. Therevidae, Chapter 37, pp. 513- 5233/7 McAlpine: JoF. Be Vv. -Retersons G: E! Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds., Manual of Nearctic Diptera. Research Branch, Agriculture Canada Monograph 1: 1-674. Lyneborg, L. 1968. A comparative description of the male terminalia of Thereva Latr., Dialineura Rond., and Psilocephala Zett. (Diptera, Therevidae). En- tomologiske Meddeleser 36: 546-559. 1972. A revision of the Yestomyza-group of Therevidae (Diptera). Annals of the Natal Mu- seum 21: 297-376. 1976. A revision of the therevine stiletto- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON flies (Diptera: Therevidae) of the Ethiopian region. Bulletin of the British Museum of Natural History (Entomology) 33: 189-346. 1978. The Afrotropical species of Phycus Walker (Diptera: Therevidae). Entomologica Scandinavica 9: 212-233. McAlpine, J. F. 1981. Morphology and terminolo- gy—Adults, pp. 9-63. Jn McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vock- eroth, and D. M. Wood, eds., Manual of Nearctic Diptera. Research Branch, Agriculture Canada Monograph 1: 1-674. Rauch, P. A. 1970. Electronic data processing for entomological museums, an economical approach to an expensive problem. Ph.D. dissertation in entomology, University of California, Riverside. 78 pp. Stuckenberg, B. R. and M. E. Irwin. 1973. Standards for entomological labels. Bulletin of the Ento- mological Society of America 19: 164-168. Swofford, D. 1990. PAUP: Phylogenetic Analysis Us- ing Parsimony, Version 3.1. Webb, D. W. and M. E. Irwin. 1988. The genera Ataenogera and Phycus in the New World. Pro- ceedings of the Entomological Society of Wash- ington 91: 35-50. 199la. A revision of the Nearctic species of Dialineura Rondani and Pallicephala Irwin and Lyneborg (Diptera: Therevidae: Therevinae). Pro- ceedings of the Entomological Society of Wash- ington 93: 869-898. 1991b. The Nearctic genus Nebritus Coquil- lett (Diptera: Therevidae: Therevinae). Proceed- ings of the Entomological Society of Washington 93: 899-913. 1991c. The North American genus Megal- inga Irwin and Lyneborg (Diptera: Therevidae: Therevinae). Proceedings of the Entomological Society of Washington 93: 914-924. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, pp. 225-226 NOTE Lectotype Designations for Two Lace Bugs Described from the United States (Heteroptera: Tingidae) At the request of J. Péricart, who is pre- paring the Tingidae chapter for the upcom- ing Catalogue of Palaearctic Heteroptera, the National Museum of Natural History col- lection at the Smithsonian Institution was searched for the available specimens in the type series of two species that were de- scribed from the United States but have since proven to be junior synonyms of Old World Species. To be able to maintain the format of the new catalog, it is necessary to designate a lectotype for each of these two species names. Leptobyrsa explanata Heidemann (1908, Proceedings of the Entomological Society of Washington 10: 105) [a junior synonym of Stephanitis rhododendri Horvath, 1905, Annales Historico-Naturales Musei Na- tionalis Hungarici 3: 567, synonymy by Champion, 1916, Entomologist’ Monthly Magazine 52 : 207]. The original description included reference to “several specimens, male and female .. .” from several eastern United States localities, and then followed with a paragraph ““Type (Rock Creek, D.C., July 6, 1987, female and male.— No. 9909, U.S.National Museum.” Here the assump- tion 1s made that Heidemann intended to separate his series into two sorts of types, the two Rock Creek specimens to have dominance. Accordingly, the male labeled “Rock Creek, D.C. 6-7-97”; “‘O. Heide- mann Collector’; ““Leptobyrsa explanata Heidem. on Kamia [sic] latifolia [with O.H. in lower left corner]’’; and “TYPE No. 9909” is here designated the lectotype. The type number label was added by persons un- known. Heidemann’s interpretation of the date must be accepted because he wrote that label. The remaining members of the type series found in the collection are designated as paralectotypes and include specimens la- beled: 1 2, same collecting data as lectotype; 1 3, “8690 on Rhododendron maximum” ‘““Kovcona, N.C., Sanderson, July 29-99”; 1 4, 1 9, “°7531 Hopk., W. Va.” “Tibbs Run, Dellslow, W. Va.’ 1 4, 1 2; 1 2, Black Mts., N.C. VI/24” “Beutenmuller, Am. Mus. Nat. Hist., N.Y.” Several other specimens at hand have the same localities reported with the original description but have different dates. Sphaerocysta Peckhami [sic] Ashmead (1887, Entomologica Americana 3: 155), a junior synonym of Galeatus spinifrons (Fal- lén) (1807, Monographia cimicum Sveciae, page 38); to synonymy under Galeatus an- gusticollis Reuter, 1874, by Drake and Ru- hoff (1961, Bulletin of the Southern Cali- fornia Academy of Sciences 60: 163), then the latter species was made a junior syn- onym of Galeatus spinifrons by Drake and Ruhoff (1962, Studia Entomologica 5: 490). The two specimens referred to in the orig- inal description are present. The male, here designated lectotype, is labeled ““Milwau- kee, Wis.”’; “Collection Ashmead”; “Type”; ‘““Galeatus (Sphaerocysta) Peckhami Ash.”’; “TYPE No. 9910 U.S.N.M.” The first two labels are in print, the next two are in Ash- mead’s handwriting, and the “TYPE No.” label was added subsequently by an un- known person. The second specimen, a fe- male here made a paralectotype, has the two labels in print, “Milwaukee, Wis.” and “Collection Ashmead” and labeled “Type” in Ashmead’s handwriting. Acknowledgments: Appreciated and helpful reviews of the manuscript were made by Thomas J. Henry, Systematic Entomol- 226 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ogy Laboratory, U.S. Department of Agri- Richard C. Froeschner, Department of culture, % National Museum of Natural Entomology, MRC-105, National Museum History, Washington, D.C. and Paul J. of Natural History, Washington, D.C. 20560. Spangler, Department of Entomology, Na- tional Museum of Natural History, Wash- ington, D.C. PROC. ENTOMOL. SOC. WASH. 97(1), 1995, p. 227 NOTE Heinrichiessa sanpetella Neunzig (Lepidoptera: Pyralidae) is an Inquiline in Juniper Midge Galls (Diptera: Cecidomyiidae) Imbricate midge galls we collected from western juniper (Juniperus occidentalis Hook.) in Oregon (Grant Co., Mount Vernon, elev. 1370 m, XI-1992) and “‘fruit”’- like midge galls from alligator juniper [Ju- niperus osteosperma (Torr.)] in Arizona (Pinal Co., Oracle, elev. 1700 m, V-1993) contained larvae of a moth recently de- scribed and known from only five speci- mens collected in central Utah, Heinrichies- sa sanpetella Neunzig (1990. The Moths of America North of Mexico, fasc.15.3: 98) (Pyralidae: Phycitinae). Heretofore the host plant of larvae of this lepidopteran was un- known. Our records extend the moth’s known range from the Utah type locality 900 km northwest to central Oregon and 800 km south to Arizona. The Oregon artichoke-like galls are pre- sumably generated by larvae of a Walsho- myia sp., and probably undescribed (Gagne 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ithaca). Three other Walshomyia spp. are implicated by Gagne in formation of “‘fruit’’- like juniper galls, but he gives no Arizona records and none from alligator juniper. The inquilinous H. sanpetella larvae, found in approximately 20% of the Oregon galls and 4% of the Arizona galls, do not coexist with conspecifics nor with (solitary) gall maker midge larvae. They overwinter as larvae and pupate in early spring. Vouchers of all life stages of this lepidop- teran are held by F.F.P. at Ohio State Uni- versity, Columbus. We thank S. Passoa (USDA/APHIS/PPQ, Reynoldsburg, Ohio) for identifying the moth and H. H. Neunzig (Department of Entomology, North Carolina State Univer- sity, Raleigh) for confirming its identifica- tion. R. J. Gagné, M. A. Solis and C. J. Drake made valuable comments on the manuscript. J. J. Furlow, Curator of the Herbarium, Department of Plant Biology, Ohio State University, Columbus, offered botanical advice. Foster Forbes Purrington, Department of Entomology, Ohio State University, Colum- bus, Ohio 43210; Teal MacKenzie Purring- ton, Bureau of Land Management, U.S. De- partment of the Interior, Prineville, Oregon 97754. 1 ae ei Cpls “40s Gah OR AV ge Ratt iS a Tie aus er ; wa wi tote mee ru tied ting 480 Aaa - at: ba, pied Gis yeh ealloayeh 7 ary algo sean tau robes Gilt Pav agit Sh ala Yrnibariv: <= cre nn | wb nihde dnl (s oy, 0 5,*Lgd tot wie onaet 5 aD, lexan at . . adele tonte eae! as Bn ” ned ALEPSETA ASTD) acne ehh mas ve ‘eianteyl Jer Soon og Seeaigge | one: meni? il Foil) in ileaeeoe Ye his. eT heel: oa > an ) | lex potgeciriatt t{ sieeertif Mo tihap waren 0 -heeey eea ‘ are ahhh ist) baer? peal pes itive \Naalamit 7) iA Se Sex ee repeat te rel marirtiier.. ei ee ea taht (tia Ay A aT pecaia ney fale? % a vy: nal PS bee qlee 24 fee VN A oe) pie oo hee Le woe ran ~ eerie, « iene (ag) ast fie eV Td “or} soit Geren 4 ys ‘* ane ar ye H Srey HG ; ra Wig ) \euera Ge jeseale rity isapee) rT Te aos, ai digest rereay 2 eras i ig wheter roger | Gio 2 hs egwenl yl Tgtivyiag ie nah | = fe | se Sa! etd) hed nicer ype or ay ie rth ; | ay valyy fet liaw? gntilyee'’ trey "S ele ae | Meee. LINEN at gbtorin') Tee) ha | ice mrit | o aeneot\: tela aaa lao alan: | vn ent own ‘cunt 4 eR i | | 41th pe ling ay ow ¢ Pay fF 7 ° d "al “ayn hay PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS Cynipid Galls of the Eastern United States, by Lewis H. Weld. 124 pp. 1959... Cynipid Galls of the Southwest, by Lewis H. Weld. 35 pp. 1960... as Tale TE ae Ber Papers COMIC Y Nie Calls w-2e..06 2 oat Ae eg oat tee a MEA Unusual Scalp Dermatitis in Humans Caused by the Mite sehiumt dont ps Jay R. Traver. PSSA ARSENE USRE URL EEA Sa AEE Oe ee 5 AINE RRND 2 Pd ge OMOEA NIC eh etapa) 2 Eh nae Se TSP Aa A Short History of the Entomological Society of Washington, by Ashley B. Gurney. 15 pp. 1976... Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. Steyskal -Taxonomic Studies on Fruit Flies of the Genus Urophora eee Re oan by George C. Pe USROr Gi pp LUMO he ccattete Mm ker e E dade Se Ne Ted Ome MLR ENO MOL. Aline eM Ute aU RED A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael PC MEE GO LOGO 2. 82 Aree! al a NE Nw GS tel ed sa TON me RL eRe Re MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939... No. 2. A Classification of Larvae and Adults of the Genus edicts wand by Adam G. Boving. SSM fay pe PM SEB MONE Ndi SR RON a 21 NOE Ne oe EN aN Soe sey ee Pa Yoel Bl Bo eat) ick UE SEL No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. ISIS By 1) 0 \S6 C2 2 Rena faa ES a Oa Sn LS Re pee Aan EER Paulie We MnO TORS 3 : 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. Beit oe ee ia a ee eo LP eue eis ek i Ry a ARN CL ie eR ries bere, 00 ae a a eg oe _ No.6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi A) pistes ise SO LOIS R ESC a Tepe aa pa UE ks NY ae Ie dae ae tae SES No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. [WO IFAL GYM eal sea DAU ins A IP ade a eat 4 ee Ra eg AMO) LD ATER SEM Much See MA ihe SAR he Ss Asie WU Ne ea No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Pspicta: Cerato- pozonidae); by/W, E: Grogan, Jr.and W.W.,Wirth. 125 ppv 1979. <2 5-0 o he ed Cee No.9. The Flower Flies of the West Indies (Diptera: Pe by F. Christian ree 200 pp. 1A eS Ea AUR AAS AAS MD ANAS UROL EN can WOOP BARD OKC SPR UET OL Je aL Oh fd No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W. Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982 -.............. No. 11. A Systematic Study of the dries ere: gens by Kenkichi sane 370 pp. LO) SSrsiee ls See MSs ig Bek eT tec ee QV ty Fg eta a. Ms ANNU eid Yl Ae Dee Sel yy Sn ee ee cE No. 12. The Holarctic Genera of kegs Sinan Chalcidoidae), 2 Michael E. Schauff. 5, TGR O18 9 Betas hi MPU dice tats IEG fA Pea ae Renae RAEN HEN Et Prt RUN rae ote Se No. 13. An Identification Manual for the North American Genera of the Family Braconidae (Hy- menoptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987 15.00 11.00 12.00 10.00 11.00 18.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $25.00 per volume to non-members and $13.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10 per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian Insti- tution, Washington, D.C. 20560. CONTENTS (Continued from front cover) MALDONADO CAPRILES, J.—Notes about the Old World genus Hexamerocerus Reuter (Het- eroptera: Reduviidae: Ectrichodiinae) . MILLER, SCOTT E., VITOR O. BECKER, and RAUL VELEZ-ANGEL—Podalia bolivari (Lepidoptera: Megalopygidae): a highly sexually dimorphic neotropical pest POLHEMUS, JOHN T.—The identity and synonymy of Nepa fusca Linnaeus, 1758 (Het- eroptera: Nepidae) POLHEMUS, JOHN T. and PAUL J. SPANGLER—A review of the genus Stridulivelia Hunger- | ford and two new species (Heteroptera: Veliidae) from South America SHAW, SCOTT RICHARD—A new species of Centistes from Brazil (Hymenoptera: Bra- conidae: Euphorinae) parasitizing adults of Diabrotica (Coleoptera: Chrysomelidae), with a key to new world species SURDICK, REBECCA F.—New western Nearctic Swe/tsa (Plecoptera: Chloroperlidae) VOEGTLIN, DAVID—Notes on the Mindarus spp. (Homoptera: Aphididae) of North America with descriptions of two new species WEBB, DONALD W. and MICHAEL E. IRWIN—The New World genus Chromolepida Cole (Diptera: Therevidae: Therevinae) NOTES FROESCHNER, RICHARD C.—Lectotype designations for two lace bugs described from the United States (Heteroptera: Tingidae) PURRINGTON, FOSTER FORBES and TEAL MAcKENZIE PURRINGTON—Heinrichiessa sanpetella Neunzig (Lepidoptera: Pyralidae) is an inquiline in juniper midge galls (Diptera: Cecidomylidae) VOL. 97 APRIL 1995 NO. 2 ~ PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED QUARTERLY CONTENTS CONTRIBUTIONS ON HETEROPTERA DEDICATED TO THE MEMORY OF JOSE CANDIDO DE MELO CARVALHO 1 EL GY ENT ICOY RGM) Wirth AW oe ME ai a aah US RR PU re Thomas J. Henry and A. G. Wheeler, Jr. 229 HOSE, @anGido de WMeloi@arvalhoe h2 nss 2b etter Sak ay dheiecle ed adawen «i ean ctenedorte ae Milza Carvalho 231 Pikeimakins* of an) EntOMmOlogish va. Nasick shoes laa kead sialge tata euleas Marcio Antonio de Carvalho 236 José C. M. Carvalho: A student’s remembrance ..................... Paulo Sérgio Fiuza Ferreira 240 ASQUITH, ADAM—Loulucoris, a new genus, and two new species of endemic Hawaiian plant bugatieteroptera; Mitnidaes Onthotylinae) si. o.oo ale es ns RR otal ae STAs atm aaa oe 241 BRAILOVSKY, HARRY—New genus and new species of Colpurini (Heteroptera: Coreidae) TOR Aes EO DINE REDUD GH seh sehtc ce bam tn 07, eine ASN ULIN Mine NC Mets canine ei iam nit ee ay 250 CASSIS, GERASIMOS—A reclassification and phylogeny of the Termatophylini (Heteroptera: Miridae: Deraeocorinae), with a taxonomic revision of the Australian species, and a review Ghihertnbaluclassiicationon the DeracOcOrmacwy a.) .)5. sean hee weaeen eee oe Marek tent ot ase: 258 FROESCHNER, RICHARD C.—Review of the New World lace bug genera Acanthocheila Stal and Carvalhotingis new genus (Heteroptera: Tingidae) ................ ccc cee ce eee e ee ee eee eee eee 331 HENRY, THOMAS J.—Proboscidotylus carvalhoi, a new genus and species of sexually di- morphic plant bug from Mexico (Heteroptera: Miridae: Orthotylinae) ........................ 340 MALDONADO CAPRILES, J. and GEORGE O. POINAR, JR.—Reuteroscopus carvalhoi n. sp., a new fossil plant bug (Heteroptera: Miridae: Phylinae) ....................0.cc cece eee ee 346 POLHEMUS, JOHN T. and DAN A. POLHEMUS—A phylogenetic review of the Potamobates fauna of Colombia (Heteroptera: Gerridae), with descriptions of three new species ........ 350 SCHAFENER, J. C. and P. S. F. FERREIRA—Carvalhoisca, a new genus of Orthotylini from MERIGO iy M Ar eICLEFODtet a) aides. yaa cen ot eee Lae aia elem ahd outs aielets stud Baro ete andi dataan bilo S73 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1995 JOHN W. NEAL, JR., President NoRMAN E. WOODLEY, Treasurer RALPH P. ECKERLIN, President-Elect NATHAN SCHIFF, Program Chairman DARLENE D. Jupp, Recording Secretary M. ALMA SOoLis, Membership Chairman Ho.uiis B. WILLIAMS, Corresponding Secretary PAUL J. SPANGLER, Past President JAMES PAKALUK, Custodian THOMAS J. HENRY, Editor DavID R. SMITH, Associate Editor Gary L. MILLER, Book Review Editor Publications Committee DONALD R. DAVIS TERRY L. ERWIN F. CHRISTIAN THOMPSON A. G. WHEELER, JR., Special Publications Editor Honorary President CurTis W. SABROSKY Honorary Members LouIsE M. RUSSELL ALAN STONE KARL V. KROMBEIN All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, MRC-168, Smithsonian Insti- tution, Washington, D.C. 20560. 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 8 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, MRC-168, Smith- sonian Institution, Washington, D.C. 20560. Members in good standing receive the Proceedings of the Ento- mological 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 P. 218 OF THE JANUARY, 1991 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. Editor: Thomas. J. Henry, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- West, Beltsville, MD 20705. Special Publications: A. G. Wheeler, Jr., Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, PA 17110-9408 Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 27 June 1995 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA ~ © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 229-230 FOREWORD Beginning students of miridology soon learn that the Carvalho “‘Catalog of the Mir- idae of the World” is the bible of their sci- ence. Long before we ever met José, we knew him through his hundreds of papers, keys to the genera of the world, but, most of all, through his indispensable world catalog. Before Carvalho, the mirid literature had grown enormously over the previous cen- tury and a half and was inaccessible to most researchers, except for a few specialists (e.g. H. H. Knight and E. Wagner), who de- scribed large numbers of taxa in many small, scattered papers. The appearance of the Carvalho catalog opened up a new era in mirid research, providing new students a quick grasp of the world literature. Scientists such as José Carvalho come along only once in a great while. His more than 500 published papers and descriptions of nearly 400 genera and more than 2000 species reflect an unusual enthusiasm and self determination. To put things in a little better perspective, José described more spe- \ \ Fig. 1. Heteropterists in attendance at XV International Congress of Entomology, Washington, D.C., 1976. [top row, left to right] Reece I. Sailer. José C. M. Carvalho. John D. Lattin. Peter D. Ashlock. Merrill H. Sweet. Cecil L. Smith. B. Jane Harrington. Alfred G. Wheeler, Jr. James A. Slater. [bottom row] René H. Cobben. Joseph C. Schaffner. Randall T. Schuh. Thomas J. Henry. Ryuichi Matsuda. Harry Brailovsky. 230 cies of Miridae than are known for all of Canada and the United States. Perhaps just as remarkable are José’s numerous activi- ties outside of mirid research. He held many offices and positions—from President of the International Congress of Entomology to Director of the National Museum of Nat- ural History in Rio de Janeiro. One only has to read about his diverse extracurricular activities (see articles in this volume by his wife Milza and brother Marcio) to be in awe of this man’s accomplishments. We first met José at the 1976 Interna- tional Congress of Entomology held in Washington, D.C. (Fig. 1), where he became president-elect for the 1980 congress sched- uled for Kyoto, Japan. Despite his many accomplishments, this renowned scientist had time to talk to two beginning students and share with us his vast knowledge and enthusiasm for Miridae. Throughout the years since that meeting, we grew to admire José for his seemingly endless desire to make the Miridae better known. In 1985, he host- ed one of us (TJH) for six weeks in Brazil. His hospitality and help in making arrange- ments to collect and visit various Brazilian museums made this trip wonderfully pleas- ant and productive. That time spent with José made it easy to see how focused he was on his science. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON In 1992, we were reminiscing about our careers in entomology and our modest be- ginnings in miridology, when José Carval- ho’s name came up. We realized José had been publishing on Miridae for more than 50 years and would turn 80 in June of 1994. That afternoon, we decided to organize a Festschrift in his honor. Unfortunately, it took us a little longer than we anticipated, and sadly he is not here to receive this col- lection of papers written in his honor (José died October 21, 1994). This volume is therefore dedicated to his memory and his great accomplishments. We thank all of the authors who contrib- uted papers for this occasion. We are es- pecially appreciative of Muilza Carvalho, Marcio Carvalho, and S. P. Fiuza Ferreira for their personal tributes to José. Their ar- ticles allow us to see more into the man behind the scientist. We also are grateful to Iara Maria Carvalho for her help in coor- dinating correspondence, editing, and ob- taining photographs for the introductory ar- ticles, and to the Entomological Society of Washington for allowing us to devote this entire issue of the Proceedings to the mem- ory of Dr. José Candido de Melo Carvalho. Thomas J. Henry A. G. Wheeler, Jr. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 231-235 JOSE CANDIDO DE MELO CARVALHO— A WIFE’S VIEW MILzA CARVALHO Rio de Janeiro, Brazil Thomas J. Henry asked me to write a paper for the volume honoring José. After 54 years of marriage, I wondered if I could do justice to him. He was a person of many accomplishments and numerous facts cer- tainly have been left out of this writing. What I have written only touches on his outstand- ing and productive career. Letters from his grandmother (1923) and from his father, which we still keep today, aided me in re- calling much about his early years. Also his personal letters (1938-1973) helped me re- call certain details of his scientific career. José was born on June 11, 1914, ona farm named Saint Joséph in Conceicao Apare- cida Village (the popular name was Barro Preto=Black Mud), a District of Carmo do Rio Claro City in the southern part of Minas Gerais. His parents and grandparents were farmers, and his mother died when he was 5 years old. His father married again some years after. José was raised among eight brothers and sisters. He spent his childhood on the farm, where he learned to love na- ture, animals and, in particular, insects. An oxcart driver and fisherman named Sebas- tiao Pereira, who worked at the farm, was his idol. They used to go together to the virgin forests existing then to watch wild animals and take native honey. José obtained his primary (elementary) education from Lady Lia (Teacher Lia) at the Diocesano School in Minas Gerais. His father had in his library the “‘Natural His- tory of Buffon,” which José loved and read; undoubtedly, this book contributed to his interest in zoology. He attended Champag- nat High School in Franca, Sao Paulo, and during this time, he lived with his uncle Antonio Candido de Melo Carvalho. There he traveled from the farm to school, car- rying his briefcase and books by horse. In high school, he read a lot of ““Tesouro da Juventude = Youth’s Treasure,” an ency- clopedia containing facts and curiosities of the world in general, which had a big influ- ence on him. By 1930, a great recession occurred in Brazil and the situation for farmers, includ- ing his father, became very bleak. Under the advice of his grandmother and step- mother, his father decided to enroll him at the Vicosa Agricultural and Veterinary Uni- versity to study agriculture and learn more about modern farming methods. After fin- ishing this program in 1933, he returned to the farm to work with his father for two years before again entering school at Vicosa to pursue studies in Veterinary Science. There, José was greatly influenced by Ruy Gomes de Morais, a parasitologist, and Joao Moojen, a zoologist. During this time he entered sports, participating in Brazilian and South American championships. In 1936 he took his talents to Paris, representing Brazil in the Youth Olympic Games and won 4th place in the pentathlon. He finished his degree in 1938. During these years he specialized in studying ticks with Henrique de Beaurepair Aragao at the Oswaldo Cruz Institute. The first year after graduating, he taught at Vicosa and during one of the visits to our hometown, he went to my father’s farm to instruct the local peo- ple on how to control some dangerous ants that were plaguing the plantation. In 1939, we married after a long friendship. In August 1940, José received a schol- 232 Fig. 1. lege, Ames, Iowa, 1942. José and Milza Carvalho, Iowa State Col- arship to the University of Nebraska. We traveled to the United States in a cargo ship that carried only a few passengers. It was a hard trip lasting 28 days, with stops at many port cities along the Brazilian and Vene- zuelan coasts. We arrived in New York and took a Greyhound bus to Nebraska. Mrs. Snipes (the mother of Dr. Snipes, a well- known professor from Vicosa) was waiting for us. She arranged for an apartment to rent and gave us a basket full of “Brazilian” and American food, including types we had nev- er seen before. I’ll never forget her kindness. Soon José met with his major Professor, Dr. Harold Manter, a parasitologist. José brought some birds stuffed with straw to give to the university and specimens of Gor- diacea, which were to become his graduate research group. During our stay in Nebraska (1 year) José received from Burbank, California, a letter PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON that had great influence on his life. It was from the ex-director from Vicosa Univer- sity, Dr. John B. Griffing, who said “To nearly all of my friends from Vicosa Uni- versity I advise not to bother about degrees or even grades, but in your case the advice is different. You have the rare gift of re- search and scholarship, so I do hope that the way may be opened for you to take both the Master’s and Ph.D. also. This would enable you to gain much more international recognition for your future in Brazil.” From Nebraska we moved to Ames, Iowa, to pursue graduate work at Iowa State Col- lege (2 years). José met Drs. Carl J. Drake, Harry H. Knight, and Halbert M. Harris. During this time, he received only $100 a month. An encounter with Dr. Knight, who claimed that he could not identify a bug, left José despaired. Dr. Drake came to his res- cue, however, and borrowed some Miridae from the Carnegie Museum in Pittsburgh. As a result, José’s first mirid paper was with Dr. Drake. From then on, the Miridae be- came his life-long pursuit. He successfully defended his thesis on the genus Eimeria, under parasitologist Ellery Becker. Life during José’s graduate years are still vivid. I remember Homecoming Day and all the ceremonies at Iowa State. I also re- member the day that the United States de- clared war on Japan. Finally it came time for us to return to Brazil. On the way, we stopped in Florida to visit Dr. P. H. Rolfs, the founder of Vi- cosa University. It was difficult to get trans- portation to Brazil, especially because of World War II then in progress, but we man- aged to get tickets on a Chilean ship that sailed from New Orleans to Valparaiso, Chile, crossing through the Panama Canal. The return was made by convoy, with our ship sailing in the middle. We had to stay in collective and separate rooms. We wer- en’t allowed to listen to the radio or make noises and at night no lights or cigarettes were allowed because of the bombing threat. On the way, the ship stopped in Guayaquil, VOLUME 97, NUMBER 2 Fig. 2. José Carvalho with Kamaiuna children, Xingu, Para, Brazil, 1947. Lima, and Arica. At Arica the ship stopped for eight days, so we traveled to La Paz and returned to Arica before resuming. Arriving in Valparaiso, we crossed the Andes to Montevideo, and then completed the last part of our trip by train to Sao Paulo. The whole trip lasted three months and 11 days. During this long and sometimes difficult trip José managed to visit several museums and meet with zoologists. Back to Vicosa, José taught zoology, bi- ology, and animal parasitology. Life was quiet. During this time he received from the United States a letter stating that ““You are being considered for a position as a Com- missioned Officer in the Army.” This news made him very happy, although, of course, as a Brazilian citizen, he could not accept. In 1946 we moved to Rio de Janeiro, where José became a zoologist at the National Mu- seum of Natural History. He was motivated to work hard by the lack of a good library, adequate collections, and, above all, by the lack of well-trained staff members. During this time, he received an invitation to be- come a researcher at the North Agronomic Institute, and from Dr. Melo Barreto to take a zoologist’s position at the Rio de Janeiro Zoo (a position he accepted and kept until 1952). Our daughter, Iara, was born in 1947. Two years later, José ventured on an expedition to the Rio Negro (Amazon Region). We were unable to accompany him on this trip be- cause the region was too wild and primitive. Other expeditions to the Amazon Region followed: in 1950 to the Javari, Itacoari, and Jurua rivers, and in 1952 to Paru de Leste River. He traveled extensively throughout 234 the Amazon Region, collecting many bugs and studying zoology. In 1949, he was elected a member of the Brazilian Academy of Science. That same year, he traveled to England to study with W.E. China at the British Museum (Natural History). Again Iara and I weren’t able to follow because life was hard so soon after World War II. In Europe he visited many museums and researchers in Austria, Bel- gium, France, Germany, Italy, Portugal, Scandinavia, and Switzerland. He attended the International Congress of Entomology in Amsterdam and published with Eduard Wagner. Back in Brazil in August 1951, he was offered by O. Monte’s widow the chance to buy her husband’s well-known tingid col- lection, which, of course, he purchased. He worked at the Museum in Rio with bugs until 1953 when he received a Fellow- ship from the Guggenheim Memorial Foun- dation to work on his world catalog in the United States. Iara and I were able to ac- company him on this trip. We traveled on a ship from the Moore McCormick Com- pany, which was much more comfortable than our previous trip in 1940 (only 15 days this time) from Rio de Janeiro to New York. In Washington, José worked with Dr. Reece I. Sailer at the National Museum of Natural History. He also met Harry G. Barber, and Drs. Dave A. Young and R. E. Snodgrass. From Washington, we traveled to Ames, Iowa, where José worked with Dr. Knight for one year and then to Berkeley, Califor- nia, where he worked with Dr. Robert L. Usinger for a few months. Iara attended school in Ames and Berkeley. She learned to read and write first in English, then in Portuguese. During this time, we crossed by car, visiting such National Parks as Yellow- stone, Yosemite, and the Grand Canyon. José studied his bugs, visited museums, and worked with other zoologists the whole time. We spent the final months of 1954 back in Washington, D.C., where José was occupied with activities linked to the completion of his World Mirid Catalog and research with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Reece Sailer on Pentatomidae. That Christ- mas, we bought our first TV set. In March 1955, we took another Moore McCormick ship back to Brazil. In June 1988, José began cooperative work with the Amazon National Institute for Research and became Director of the Goeldi Museum in Belem, Para for several months. Soon after, he was appointed Di- rector of the National Museum in Rio de Janeiro, a position he held for two terms, six years all together. The Museum was al- ready old at that time and in much need of repair. José was instrumental in the Mu- seum’s renovation and the creation of many new and innovative exhibits. He continued to publish many more papers and made nu- merous other expeditions to the interior of Brazil to collect bugs. In 1972, the Brazilian Foreign Affairs Ministry chose José to represent Brazil at the World Conference on Environment in Stockholm. In January 1973, we traveled to College Station, Texas, where José worked with Dr. J. C. Schaffner at Texas A & M University. In late 1973, he received an in- vitation from Dr. D. F. Waterhouse to work at the Division of Entomology, CSIRO, in Canberra, and again José and I traveled afar. We had an exciting trip, visiting museums and the major cities of Australia. In 1978, he was elected President of the Brazilian Academy of Zoology and, from 1980-1984, he served as President of the International Congress of Entomology, held in Kyoto, Ja- pan. As a result of all this travel, he earned the nickname of “globetrotter”’ from his col- leagues! Among his other activities were: Director of the Museum Paraense E. Goeldi (1954— 1955); Director of the National Museum (1955-1961); Naturalist Museum (1946- 1984); Brazilian Foundation for Conser- vation of Nature (1966-1969, 1979-1984); National Research Council (1962-1964); Brazilian Institute for Development of For- estry (1961-1964); Federal Council of Cul- ture (1974-1985); Zoological Garden (1946- VOLUME 97, NUMBER 2 1972); Permanent Commission for the In- ternational Congresses of Entomology (1951-1980), President (1980-1984); As- sociation of Tropical Biology (1965-1967); International Union of Conservation of Na- ture (1963-1970, 1976-1981); and Group for Studies on Environment of the Vale do Rio Doce Company (1980-1991). He was made an honorable member of the Zoolog- ical Society of London, the Royal Ento- mological Society, and the International Congresses of Entomology, as well as many other societies. In addition, more than 45 animals—either species or genera—were named after him. After such a demanding career and many trips, not to mention some aging, his heart began to give him problems. In September of 1982, we flew to Cleveland, Ohio, for an operation on the mitral valve of his heart. For José, this was only a minor setback and, 233 upon recovery, he started to work again as hard as ever. In Brazil, 70 is the mandatory retirement age for government employees, so in 1984 he retired. José was able to maintain his research office at the museum where he con- tinued working every day. In 1989, he de- veloped a blood clot in the right leg, but he recovered quite well after surgery and con- tinued working at the Museum. In 1992, he broke his femur, which left him in a wheel chair. Even so, he continued working at home every day with his bugs, microscope, and correspondence to scientists around the world until October 21, 1994. I can say that it was very special to have shared my life with such an intelligent and capable husband. His willingness to help students and people, and his determined way of working and studying greatly enriched my life. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 236-239 THE MAKING OF AN ENTOMOLOGIST Marcio ANTONIO DE CARVALHO Belo Horizonte, Brazil The following is an outline of the early years and family life of José Candido de Melo Carvalho, as remembered by me, with the aid of relatives (stepmother: Candida Vilela Carvalho (1892-); half brothers: Bo- livar (1923-) farmer, Cassio (1925-) farmer, and Marcio (1936-) commercial pilot; half sisters: Mariana (1924-) farmer’s wife, Gil- da (1927-) farmer’s widow, Lucia (1929-) retired teacher) and his own notes. It is with great pleasure that we present this infor- mation on such an accomplished man and good brother. We believe the report that follows is an accurate description of what we know about José. Some topics may be hard for conser- vationists to appreciate, but these activities occurred in the 1920s in an aggressive en- vironment where such things were different than they are today. José, best known by family and friends as Zezinho (Joseph or Joe), was born on June 11, 1914. He was the first of two sons from the first marriage of our father. His mother, Ano, died prematurely in 1919, leaving five-year-old José and three-year- old Mauricio (1916-1973). Father married again in 1922, giving José six step brothers and sisters. The family was large, with un- cles from both sides numbering 26. A description of the environment near José’s home is necessary to understand his interest in nature. The area where our fam- ily of European descent settled was in the farmlands of Carmo do Rio Claro in Minas Gerais, approximately 230 miles NW of Rio de Janeiro, Brazil. This area, with mild ir- regular topography at an altitude of about 2300 ft, was developed during the 1800s and consists of partially cut semitropical forests. People in this part of the world were wise enough to preserve some untouched parts of forests as reserves. Unfortunately, devastation came after 1945. Our grandfa- thers used the land for mixed cultivation and grazing cattle. The climate in this region is very mild with temperatures varying mostly between 60° and 80°F. The dry win- ters during May, June, and July have oc- casional lows around 30°F, with some frost, but they are absolutely snow free. The rainy season occurs from October to March with a maximum temperature of 90°F. Because of this climate and location, the vegetation is rich and the wildlife plentiful. Before 1945, you occasionally heard of farmers losing cattle to attacks by jaguars. Snakes were common, including rattlesnakes, urutu, jar- araca, and corals. As a consequence, snake- bites were common. We hunted deer, paca, (coelogenys paca), water-hogs, and other an- imals. There are also many different birds, and the rivers, streams, and marshes were plentiful with fish and caimans. Nearly all of the goods we needed were produced locally on our farms. As a result, it was necessary for us to import only a few things such as salt, sugar, basic tools, some clothing, and medicine. Our main trans- portation was by horse and mule. For trans- porting cargo, we had two large big-wheeled wooden carts, each pulled by six to twelve oxen. Our farm had an orchard and garden that also included medicinal plants and flowers. We raised cows for milk and sheep for wool; we also had oxen, horses, mules, and donkeys to sell and for work around the farm. Our father had an ass that had only a single eye and, thus, gave him the name VOLUME 97, NUMBER 2 Carvalho family in Rio de Janeiro, 1985. José Car- valho (back), Milza (second row, left), daughter Iara Maria (second row, right), granddaughter Lilian (front row, left), grandson Leonardo (front row, right). Camoes (after the famous one-eyed Portu- guese Poet). Our primary crop was coffee, followed by maize, rice, beans, cotton, sug- arcane, and others. The houses were ma- sonry and quite large to accommodate our sizable families. Hard work was the norm for everyone, including young children. Riding horses was a natural part of country life. Chores in- cluded gathering eggs, helping handle the bullocks and shepherding, caring for live- stock, and helping raise the crops. Our grandfathers were a great influence in the education of our family. In our city we had a good nun’s college that taught the ladies to become teachers. But the boys had to travel quite a distance to the best high school. Our father and uncles provided con- siderable knowledge of the sciences, lan- guage, music, and religion. In our home we had a bookshelf containing books on varied subjects, including “‘Natural History” by Buffon that José began reading at an early age. He received a good perspective from his uncles too, who we visited regularly. On Christmas, New Years, Easter, and “City Dedication Day” everyone went to the city for religious activities. This was a good op- portunity to visit with cousins and other relatives. Of course the farms had many em- 237 ployees, and we usually had one of them as a special friend. One of José’s first teachers of botany and zoology was such a man, Se- bastiao Pereira, our father’s “‘ox cart man.” He knew a great deal about nature, partic- ularly the forest, and taught him much about the life there. Nearly every Sunday, they went hunting, fishing, looking for honey, and observing vegetation, birds, and animal signs. This knowledge, besides provoking José’s interests, proved particularly impor- tant for him later as a researcher, when he went on to explore by himself large areas of the Amazonian forest, as well as other parts of the world. At this time, however, José was being raised to become a farmer. No doubt José was born a man of great intellect and strong body (6 ft). This was apparent later by his success at college and accomplishments as an athlete. Somehow he remembered having received supple- mental feeding on mare and donkey milk as a child. José was quite precocious, and he learned to walk unassisted at seven months. At age three, our father had con- fidence enough to allow him to go alone on horseback from our farm to the city, a dis- tance of more than 15 miles. A cousin of the same age remembers him finding a bird’s nest on a visit to an uncle’s farm. He was so young that his coordination wasn’t well developed and he inadvertently stepped on the nest. After a few tears, he went on to examine the broken eggs and offspring. Af- ter their mother’s death, José and Mauricio were cared for between 1919 and 1922 by their aunt Mariana, who became extremely devoted to them. That aunt and another one lived on farms in another city 100 miles away. There José became better acquainted with 16 of his cousins. After their father’s marriage to our mother, they returned to the farm to get to know the new family. Our mother, still alive today at 102 years, and the two stepsons became great friends. Our mother’s im- mediate response to what José was like as a young boy, was “Very good boy, never teased (Nunca Amolou), very good.” At that 238 time, the region received mail by railroad that was about 30 miles away. From the train station a postman delivered the mail by donkey to our city and farms. One of José’s first jobs was going to an uncle’s farm every other day to bring correspondence and newspapers. As we described earlier, the area where we lived was rich in wildlife. Needless to say, some parts of it were dangerous because of the high concentrations of venomous snakes. Snakebites were a serious problem, so these parts of the farm were forbidden to the children. One of the forbidden places was a stream (Ribeirao), about a half mile from the farmhouse where the banks were covered with high vegetation and where snakes were particularly abundant. Well, José was a good boy, but one day when our father was away, José went fishing at the brook despite all warnings. He caught a good string of fish and returned to the farmhouse. Unfortunately for José, he and his father arrived together and, no doubt, the fish went flying in all directions. But as an indication of his determination, after the consequences of the encounter subsided, José picked up the fish one by one, went to the kitchen, and asked his stepmother to “please fry these fish for me!” Until he went to high school, José helped with farm work, learning well his probable future profession as a farmer. He remem- bers the first Ford car that arrived in the region in 1923. In the late 1920s, dad bought a car, named Oakland. The real political power in small cities of our country was the priest (vigario), a position ranked even above the mayor. After José finished elementary school, dad, convinced that his son should become a priest, sent 8-year-old José to seminary (1923-1924). The life there was of course devoted to religious studies and hard discipline from early morning to bed- time. José, however, didn’t like the new life. During his second year at the seminary, he developed an intestinal infection and his health was affected to such a degree that dad went to see him. Looking at his son’s situ- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ation, the old man’s heart softened to the extent that he changed his plans and sent him to public high school in a city near an uncle’s farm. The time in the seminary did instill discipline and patience in José, and resulted in a good basic high school edu- cation. In 1930, our state and the state where José and Mauricio attended high school, were adversaries in national politics and the sit- uation progressed to an armed revolution. Uncle Antonio sent his best employee Al- fredo on horse back to rescue Joe and broth- er Mauricio and bring them back to the farm’s safety. They fled well before day- break and remained at the farm until the armed conflict ceased. When José was in high school, the eco- nomic horizon on our father’s farm was bright and great investments were made in coffee culture and mechanical processing. Then came the effects of the world economic crisis in the late 1920s that severely affected dad’s plans. Farmers were obligated by the government to burn part of their coffee pro- duction to keep prices level. Mother and grandmother knew José needed a better ed- ucation, and convinced dad to send him to the Agriculture and Veterinary school in the city of Vicosa. José was only 15 but mini- mum age was 16, so father had to lie about his age to get him admitted. [This Vicosa School, now a facility of Minas Gerais State University, was created in 1927 under a grant from the U.S. Government under the supervision of Professor Peter Henry Rolfs, a U.S. citizen]. There José studied, learned, worked, and practiced sports with great en- ergy and enthusiasm and graduated at the head of his class. After graduation, he re- turned to help his father and work on the farm he inherited from his mother. He and brother Mauricio often competed to see which one could carry the biggest load of coffee. Both were able to handle around 200 Ibs. José was always interested in wildlife, and he frequently recruited his sisters to help hold different animals—from rattlesnakes to bats. He often helped exterminate ant VOLUME 97, NUMBER 2 nests for the neighbors. Mother remembers she didn’t like caterpillars and the time she asked José to remove a tobacco plant from a grassy area she used for drying clothes. Moments later José came back with several caterpillars in the palm of his hand and said “Look, they won’t harm you. See, I can even put one on my tongue.” We also remember a collection of 21 rattlesnakes in a room of his farm; they were eventually donated to a laboratory to make vaccines. The effects of the depression persisted, so in 1934 José decided to go back to school at Vicosa and with great effort passed the examinations to get into veterinary school. A new stint of hard work began, coupled with success in athletics to the point of being selected to represent Brazil in the 1936 Olympic Games in Berlin. However, while training, José pulled a muscle that elimi- nated him from competition. His “‘globe- trotter’’ career began in 1937, when he par- ticipated in an international university in- vitational in Paris, where he won fourth place in pentathlon. Sports aside, he still managed to find time to visit the museums in Paris. His student and athletic career at Vicosa ended in 1938 with graduation. While at school, he became interested in parasitology and zoology. In January 1939 he went to Rio de Janeiro to pursue research on ticks and the protozoans they transmitted. In February, he returned to Vicosa as a teacher and researcher. His first scientific work, published in 1939, was initiated as a student in 1937 and dealt with protozoans of Rana catesbiana Shaw. The diversity of people José knew as a boy and teenager gave him a great facility for making friends with everyone from In- dians in Amazon jungle to world VIPs. One of the best things that happened to José was the marriage to Milza Freire, the daughter of a farmer from our city in October 1939. This well-educated, charming, and petite woman proved to be a great companion for José. Following their marriage, our father died prematurely in November at age 57. 239 Remarkable to us was his travel to the United States in 1941 just before World War II, the result of a scholarship he received from Vicosa for his teaching and dedication to research. He went to the University of Nebraska in Lincoln for his M.S. Because of his many new friends and his own re- search efforts, he was admitted to the Ph.D. program at then Iowa State College in Ames. For his dissertation, he studied parasites of rabbits. His athletic training served him well, and rumors began to circulate that he could run down and catch cottontails and jack- rabbits. This story was published in the uni- versity newspaper, and soon the football and other athletic coaches were after him to join various university teams. However, he de- clined (he later claimed he could catch only young rabbits), preferring to concentrate on his studies instead. In 1943 he received his Ph.D. (thesis #689). Impressed on his mind was the quote at the University’s library: “You shall know the truth and the truth shall make you free.” In May 1943, José and Milza returned to Brazil after a complicated trip through Pan- ama and along the west coast of South America (because of German subs in the Atlantic) to Chile. From there, they traveled by train to Buenos Aires and home. From then on, José’s professional career soared and he went on to hold numerous important positions, including the director- ship at the National Museum of Natural History in Rio de Janeiro and the presi- dency of the XVI International Congress of Entomology, held in Kyoto, Japan. But these are the years best conveyed by his wife Mil- za. Even at 80 years, José, in company with Milza, his daughter Iara, a granddaughter Lilian, and a grandson Leonardo (the latter two of which are law students) continued to work with his insects on a daily basis until his death on October 21, 1994. Sometime before, he said to me “A hundred years, the maximum one can expect to live, is too short a time to accomplish everything.” PROC. ENTOMOL. SOC. WASH. 97(2), 1995, p. 240 JOSE C. M. CARVALHO: A STUDENT’S REMEMBRANCE PAULO SERGIO FIUZA FERREIRA Departamento de Biologia Animal—CCB, Universidade Federal de Vicosa, Vicosa, MG, Brazil. It was in 1966, the beginning of my sci- entific career, that I had the opportunity to meet Dr. José C. M. Carvalho at the Na- tional Museum of Natural History, Rio de Janeiro, Brazil. I vividly remember the first day we met. Dr. Carvalho was working in his office when I arrived to express my interest in studying Heteroptera. I presented to him my modest notes on the reduviid Ari/us carinatus (For- ster) that I had raised from eggs to adults in a small shoe box. He apparently appre- ciated the efforts of a sixteen-year-old high school boy trying to understand the amazing world of insects because he opened his lab- oratory and invited me to study Heteroptera under his supervision. It was not easy for Dr. Carvalho to get approval for me to work there—only college students were allowed in the Museum. After considerable effort, he convinced the administration to make an exception. Even so, some of the more conservative people still did not approve. Financial support was another problem we encountered. I was more interested in learning about bugs than worried about money, but Dr. Carvalho was concerned, for he knew that I was from a modest family that could not pay my way. His solution was to pay me out of his own resources until I entered the University. After that I was able to obtain assistantships from the gov- ernment by working on our cooperative projects. From 1966 until October 1973, I worked almost every day with Dr. Carvalho, be- coming more and more entomologically mature and learning much about his sci- entific philosophy. Under the tutilage of this hard-working, honest, and patient man, I transformed from a modest high school stu- dent to a young man with the opportunity to pursue a career in entomology. From our close relationship over the years, I grew con- siderably, and Dr. Carvalho became more like a second father to me, rather than just an advisor. I worked with him on nearly 30 papers, and we were still cooperating on various projects concerning Neotropical Miridae before his death. These papers and the other activities we shared, as well as his intro- ductions to national and international sci- entists, were very important in promoting my professional life. These experiences opened numerous opportunities for coop- erative work with other researchers, and eventually led to my M.S. (Universidade Federal do Parana) and Ph.D. (Texas A&M University) degrees, and a staff position at the Universidade Federal de Vicosa. I was very fortunate to have known Dr. Carvalho, who became my best friend and colleague. I will miss him greatly. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 241-249 LOULUCORIS, A NEW GENUS, AND TWO NEW SPECIES OF ENDEMIC HAWAIIAN PLANT BUG (HETEROPTERA: MIRIDAE: ORTHOTYLINAE) ADAM ASQUITH U.S. Fish and Wildlife Service, Pacific Islands Office, Three Waterfront Plaza, 500 Ala Moana Blvd, Suite 580, Honolulu, Hawaii 96813. Abstract. —The new endemic Hawaiian orthotyline genus Loulucoris, is diagnosed and described to accommodate two new species: Loulucoris kidoi, the type species of the genus, associated with the fan palm, Pritchardia (Arecaceae), on the island of Hawai’i, and Loulucoris cinygmiscus from the island of O’ahu. Key Words: The Hawaiian plant bug fauna (Heter- optera: Miridae) is remarkable in the un- usual autapomorphies displayed by the en- demic genera, and in its overall species rich- ness. Although a handful of new species have been added in recent years (Carvalho 1952, Carvalho and Usinger 1960, Gagne 1968, Asquith 1993), the fauna is still largely un- described and the Miridae certainly repre- sent the most speciose group of Heteroptera in Hawai’i (Howarth 1990). The tribal and generic placement of Hawaiian taxa, how- ever, is still largely based on the dated works of Kirkaldy (1902, 1904) and Zimmerman (1948b), with minor reassessments by Car- valho (1957-1960) and Schuh (1974). The subfamily Orthotylinae in Hawa1’1 is presently represented by the endemic genera Sarona Kirkaldy, Kalania Kirkaldy, Su- lamita Kirkaldy, Pseudoclerada Kirkaldy, Nesidiorchestes Kirkaldy, and endemic spe- cies in the widespread genus Orthotylus Fie- ber. In this paper I describe a new endemic genus and two new species in the subfamily Orthotylinae. Terminology of the genitalia follows that of Slater (1950) and Asquith (1991). Types and paratypes are deposited in the Bishop Museum, Honolulu (BPBM), Insecta, Miridae, plant bug, Hawaii and the American Museum of Natural His- tory, New York (AMNH). This paper is dedicated to José Carvalho in honor of his unequalled knowledge and productivity in the taxonomy of the Miri- dae. Loulucoris Asquith, NEw GENUS Diagnosis.— Recognized by its delicate form (Fig. 1); vertical, triangular head with subpedunculate eyes (Figs. 2, 3); shallow, median longitudinal sulcus and short trans- verse carina on vertex; weakly flattened pos- terolateral margins of pronotum; reddish spot on the cuneus. Description. — Male: Macropterous; gen- eral coloration greenish yellow to pale brown; dorsal surface smooth to faintly sha- greened, shining; dorsal vestiture with mod- erately long, inclined, dark setae; abdominal venter with long, pale setae. Head: Trian- gular in frontal view (Fig. 2), vertical (Fig. 3), width greater than height; vertex slightly concave, dorsal texture shagreened, with shallow, wide, median longitudinal sulcus and short indistinct transverse carina; ca- rina separating anterior shagreened surface from posterior polished area; three or four 242 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Loulucoris kidoi, male dorsal habitus male. VOLUME 97, NUMBER 2 erect, black setae along inner margin of eyes; frons prominent, extending only slightly an- terior of antennal fossae, anterior surface highly polished; tylus flat, vertical, slightly depressed at junction with frons; juga ver- tical, flat, triangular; lora vertical, flat, rect- angular; eyes subpedunculate, extending above vertex and occupying half of head height in lateral view, produced posteriad well past occiput, weakly emarginate along posteroventral margins; antennae inserted at lower third of eyes, fossa separated from eyes by one third its diameter; antennal seg- ment I weakly expanded basally, length slightly longer than width of vertex, with three or four erect, bristlelike setae on me- dial surface, all surfaces sparsely covered with short, inclined, dark setae; segment II linear, with short, inclined setae; segments III and IV linear, with short, inclined setae interspersed among longer, suberect setae. Pronotum: Trapezoidal in dorsal view, wid- er than long; flat to weakly sloping trans- versely; anterior margin straight; lateral margins straight to weakly sinuous, poste- rior halves weakly but thickly carinate; an- terior angles broadly rounded; posterior an- gles sharp; posterior margin broadly and shallowly concave; calli weakly convex, reaching lateral and anterior margins, sep- arated medially by shallow sulcus; meso- scutum broadly exposed, length one third of scutellum length; scutellum flat to weakly convex; metathoracic scent gland as in Fig. 4. Hemelytra: Subhyaline, subparallel sid- ed, widest at middle; claval vein elevated for entire length; radial vein elevated along basal half; cuneus twice as long as broad, with a large reddish spot occupying most of surface; membrane lightly suffused with fus- cous, more heavily on distal half; inner cell of membrane longer than length of cuneus but not extending past distal margin of cu- neus. Legs: Color greenish yellow; femora slightly flattened, tapered distally and ba- sally, with short, inclined, dark setae; fore femora with row of longer, erect, bristlelike setae on ventral surface; tibiae with short, 243 Fig. 2. Loulucoris kidoi, anterior view of head. inclined, simple setae; meso-/ and metati- biae with additional longer, semierect setae and several rows of minute spinulae; each tarsal segment slightly longer than segment immediately basad; claws strongly curved, thickened basally, pulvilli small, distal mar- gins angulate, parempodia convergent (Fig. >) Genitalia (Fig. 6): Genital capsule mod- erately large, broadly conical in ventral view, only slightly wider than long; and two an- teromedially directed flanges supporting the parameres; aperture vertical, large, subcir- cular. Left paramere short, L-shaped; with dorsomedially curved ventral process. Right paramere elongate, narrow, strongly curved medially. Vesica with one or two long, thick sclerotized spiculae. Female: Macropterous. Similar to male in color, vestiture, and structure except slightly broader. Genitalia (Fig. 7): Sclero- tized rings small, widely separated; lateral sclerotized area folded dorsomesally. Pos- terior wall consisting of a single saddle- shaped structure (J structure of Slater 1950) with lateral margins strongly folded ventro- mesally (K structure of Slater 1950). Etymology.—From the Hawaiian word, loulu (fan palm) and the Greek, coris (bug); referring to the host plant of the type spe- cies; gender masculine. Type species. — Loulucoris kidoi, new spe- cies. 244 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3-5. Loulucoris kidoi. 3, Lateral view of head; 4, Ostiole and evaporative area of metathoracic scent gland; 5, Pretarsus, left claw missing. Distribution. — Hawaiian Islands. thotylini (Zimmerman 1948b, Schuh 1974). Discussion. —Loulucoris is not known to _ In general habitus it is somewhat similar to occur outside Hawai’i and it is not similar Cyrtorhinus Fieber, that is represented in to any of the other endemic Hawaiian Or- Hawai’i by the introduced C. fu/vus Knight VOLUME 97, NUMBER 2 Fig. 6. = Va a-e. Male genitalia of Loulucoris kidoi. a, Genital capsule, dorsal view; b, left paramere, lateral view; c, left paramere, medial view; d, right paramere, posterior view; e, theca and spicula, dorsal view. and C. lividipennis Reuter. Loulucoris is easily distinguished from Cyrtorhinus by its triangular and vertical head. The strongly bent and thickened tarsal claws in Loulu- coris, compared to the narrow, weakly curved claws of Cyrtorhinus (Carvalho and Southwood 1955), also suggest that the two taxa are not closely related. Loulucoris appears to be related to species from the Indo-Pacific presently placed in the genus Zanchius Distant. These species also display the delicate body form; vertical head; narrow, apically furcate right para- mere, and the medially folded, apicodorsal angle of the left paramere found in Loulu- coris. The Pacific Zanchius species, includ- ing Zanchius carolinensis Carvalho, Z. fra- gilis Usinger, Z. piperi Usinger, and several undescribed species, also have thickened spiculae on the vesicae. Members of the ge- nus Zanchius; however, and the “Zanchius group” of genera, lack sclerotized spiculae (Schuh 1974). The presence of vesical spic- ulae in Loulucoris and the Indo-Pacific “Zanchius” place them in the “Orthotylus group” of genera as defined by Schuh (1974). A more extensive analysis of characters will be required to define strict synapomorphies that unite Loulucoris and the Pacific “*Zan- chius” species. Loulucoris differs from the Indo-Pacific **Zanchius”’ by its subpedunculate and pos- teriorly produced eyes, and the median lon- gitudinal sulcus. The flattened lateral pron- otal margins and posterior carina on the vertex are found in some but not all species of Pacific “‘Zanchius.” In addition, both species of Loulucoris have a distinct reddish spot in the middle of the cuneus, which I have not seen in any Pacific ““Zanchius” species. The relationships and biogeographic or- igins of several of the endemic Hawaiian mirid genera like Sulamita Kirkaldy, Pseu- doclerada Kirkaldy and Kalania Kirkaldy may remain obscure because of highly au- tapomorphic fascies. Loulucoris, however, if shown to be clearly united with the Pacific *“Zanchius” species, would be another con- firmed Indo-Pacific derivative in the Ha- waiian mirid fauna, along with Opuna shar- pianus (Kirkaldy) (Schuh 1984). 246 fi PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 7. a—b. Female genitalia of Loulucoris kidoi. a, posterior wall, anterior view; b, posterior wall, posterior view. Loulucoris Kidoi Asquith, NEw SPECIES Diagnosis. — Distinguished from the oth- er known species by its overall darker col- oration; short, acuminate tergal process on the right dorsolateral margin of the male genital capsule; the erect, basal process of the right paramere; found only on the island of Hawai’l. Description.— Male (n= 6): Length 3.18- 3.31 mm; pale greenish yellow coloration, dorsal vestiture with inclined, dark setae. Head: Width 0.73-0.74 mm; vertex 0.39- 0.41 mm; head brown, vertex and area bor- dering eyes paler, three or four erect, black setae along medial border of eyes; dorsal surface shagreened, anterior area of frons polished; tylus, and juga pale yellow, sur- faces polished, lora yellow distally, red on basal half; antennal segment I 0.46-0.54 mm, greenish yellow; segment IT 1.47-1.52 mm, greenish yellow basally, yellowish brown distally; segment III 1.25-1.35 mm, yellowish brown; segment IV 0.60-0.64 mm, yellowish brown; rostrum reaching or just surpassing mesocoxae, color yellow, apex black. Pronotum: Length 0.32-0.35 mm, posterior width 0.83—0.88 mm; uniformally brown, occasionally darker at middle of posterior disk; mesoscutum brown; scutel- lum brown, apex pale; dorsal half of tho- racic pleura dark reddish brown; venter pale greenish yellow. Hemelytra: General col- oration greenish yellow, suffused with large areas of brown on clavus, and basal and distal areas of corium; cuneus greenish yel- low with large, reddish spot; membrane slightly suffused with fuscous, less so ba- sally. Legs: Coxae and trochanters yellow; femora, tibiae and tarsi uniformally green- ish yellow. Genitalia: Dorsal margin of gen- ital capsule with short, acuminate tergal process on right side; posteroventral margin with deep socket for left paramere; sup- porting flange for right paramere broadly spatulate (Fig. 6a). Left paramere with broad area of apicodorsal edge serrate and folded medially (Fig. 6c). Right paramere narrow, elongate, evenly curved medially, apex nar- rowed, acuminate and curved dorsally (Fig. 6d), with large, erect, distally recurved pro- cess near base; smaller, basally recurved, dorsal process near apex. A single, thick spicula present, strongly recurved basally, spatulate distally (Fig. 6e). Female (n = 4): Macropterous. Color and size similar to male, except usually darker and slightly longer and wider; length 3.40- 3.46 mm; width of head across eyes 0.75- VOLUME 97, NUMBER 2 0.76 mm; width of vertex 0.43-0.44 mm; antennal segment I 0.47-0.48 mm; segment II 1.40-1.48 mm; segment III 1.38 mm; seg- ment IV 0.45-0.47 mm; length of pronotum 0.35—0.37 mm; posterior width of prono- tum 0.87-0.90 mm. Etymology.— Named for my colleague and good friend Michael Kido, University of Hawaii, Kauai Research Center, who first collected specimens of this species. Holotype 8, HAWAIIAN ISLANDS:HA- WADI: Stainback Hwy.,2.VIII.1991, ex Pritchardia sp., (A. Asquith & M. Kido). (BPBM). Paratypes. 8 6,22 HAWAIIAN IS- LANDS: HAWAII: Stainback Hwy., 2.VIII.1991, ex Pritchardia sp., (A. Asquith & M. Kido). (BPBM, AMNH). Additional specimens: Same data as type series, | 6, 2 2, all dissected, 2 nymphs (BPBM). The type series was collected from at- tached but dead leaves of Pritchardia bec- cariana Rock. Adults and nymphs were found deep in the leaf folds in association with psocids, although predation was not observed. Loulucoris cinygmiscus Asquith, NEw SPECIES Diagnosis.— Distinguished from Loulu- coris kidoi by its paler coloration; a red, dorsomedial, longitudinal stripe on the head of males; a large tergal process on the left dorsolateral margin of the male genital cap- sule, and the absence of the erect basal pro- cess on the right paramere; found only on the island of O’ahu. Description (n = 1).—Male: Length 3.35 mm; pale, straw yellow coloration, dorsal vestiture with inclined, pale setae. Head: Width 0.70 mm; vertex 0.38 mm; head yel- low, median sulcus reddish brown, erect se- tae bordering eyes gold to brown; dorsal sur- face shagreened, anterior surfaces polished; tylus, lora, juga pale yellow, surfaces highly polished; antennal segment I 0.48 mm, pale yellow; segment II 1.33 mm, pale yellow, slightly darker on distal half (antennal seg- ments III and IV missing from specimen); 247 rostrum reaching to mesocoxae, yellow, apex fuscous. Pronotum: Length 0.36 mm, pos- terior width 0.85 mm; pale yellow, anterior angles reddish brown; mesoscutum and scu- tellum straw yellow, apex of scutellum cream; ventral margins of thoracic pleura reddish brown; venter yellow. Hemelytra: General coloration pale, brownish yellow, weakly suffused with pale brown basally on clavus and corium; cuneus pale yellow with large reddish spot; membrane translucent. Legs: All segments uniformly yellow. Gen- italia: Dorsal margin of genital capsule with long, stout tergal process on left side; pos- teroventral margin entire, socket for left paramere absent; supporting flange for right paramere erect, narrowed, and recurved apically (Fig. 8a). Right paramere narrow, elongate, apex bifurcate, with two, nar- rowed, acuminate arms; basal process ab- sent (Fig. 8c). Left paramere with folded area of apicodorsal angle small, serrate (Fig. 8b). Vesica with two elongate spiculae, ven- tral spicula weakly sinuate and flattened dis- tally (Fig. 8d). Female (n= 1): Macropterous. Size sim- ilar to male. General coloration yellow, tinged with green; reddish areas on thoracic pleura indistinct; lacking longitudinal red- dish stripe on vertex. Length 3.61 mm; width of head 0.73 mm; vertex 0.39 mm; antennal segment I 3.61 mm; segment II 1.26 mm; segment III 1.09 mm; segment IV 0.70 mm. Etymology.—From the Greek, cinygm (phantom), referring to the few occasions this species has revealed itself to collectors. Holotype 6, HAWAIIAN ISLANDS: O’AHU: Castle Trail, 2000 ft., 28.XI.1937, Beating, (E. C. Zimmerman). (BPBM). Paratype 2, HAWAIIAN ISLANDS: O’A- HU: Waikane, 22.X.1947, (E. C. Zimmer- man). Additional specimens; HAWAIIAN ISLANDS: O’AHU: Halawa Valley, 11.1.1994, ex. Pritchardia, (A. Asquith) 1 é, 2 nymphs, (BPBM). An endemic genus with species found only on O’ahu and Hawai’i would be an unusual pattern within the Hawaiian insect fauna 248 Fig. 8. view; Cc, right paramere, posterior view; d, vesical spiculae. (DS = dorsal spicula, VS = ventral spicula.) (Zimmerman 1948a); therefore, it is highly likely that there are additional species of Loulucoris on other islands. With the both of the known species collected exclusively from Pritchardia, the fidelity of Loulucoris to fan palms seems likely. The genus Prit- chardia (Arecaceae) is restricted to tropical Pacific islands, with 19 endemic Hawaiian species (Wagner et al. 1990). The Pacific **Zanchius”’ species have not been recorded from palms, but are known from Marattia (Marattiaceae), Piper (Piperaceae), Cyrtan- dra (Gesneriaceae) and Ficus (Moraceae). Pritchardia hosts several other endemic Ha- walian insects with various degrees of as- sociation. The psyllid genus Megatrioza Crawford, for example, is restricted to fan palms in Hawai’i (Uchida and Beardsley 1988), while Nesodryas swezeyi Zimmer- man (Homoptera: Delphacidae) occurs on Pritchardia on Hawai'i and Nesodryas frey- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON DS VS a—d. Loulucoris cinygmiscus, male genitalia. a, genital capsule, dorsal view; b, left paramere, medial cinetiae Kirkaldy (Pandanaceae) is associ- ated with Freycinetia arborea Gaud. on O’a- hu (Zimmerman 1948c). ACKNOWLEDGMENTS I thank Michael Kido for his companion- ship and field assistance. Randall T. Schuh, American Museum of Natural History, and Dan Polhemus, Bishop Museum, kindly re- viewed the manuscript. I am particularly grateful to my wife Anna Asquith for pro- viding the fine illustrations. Tina Weath- erby patiently instructed me in the use of the electron microscope. This research was in part supported by a Research Centers in Minority Institutions Award RR-03061 to the Biological EM Facility of the University of Hawaii at Manoa. This is paper no. 3770 of the Hawaii Institute of Tropical Agri- culture and Human Resources Journal Se- ries. VOLUME 97, NUMBER 2 LITERATURE CITED Asquith, A. 1991. Revision of the Genus Lopidea in America North of Mexico (Heteroptera: Miridae). Vol. 16. Theses Zoologicae. Koeltz Scientific Books, Koenigstein. 280 pp. 1993. A new species of Cyrtopeltis from the Hawaiian Islands (Heteroptera: Miridae: Dicy- phinae). Pacific Science 47:17—20. Carvalho, J.C. M. 1952. A new species of Trigon- otylus Fieber, 1858, from Hawaii (Hemiptera: Miridae). Boletim do Museu Nacional, (Nova Ser- ie)(Zoologia) 111: 1-3. 1957-1960. Catalogue of the Miridae of the World. Arquivos do Museu Nacional, Rio de Ja- neiro. Part I. Cylapinae, Deraeocorinae, Bryocor- inae, 44: 1-158 (1957); Part II. Phyline, 45: 1-216 (1958); Part III. Orthotylinae, 47: 1-161 (1958); Part IV. Mirinae, 48: 1-384 (1959); Part V. Bib- liography & Index, 51: 1-194 (1960). Carvalho, J. C. M. and T. R. E. Southwood. 1955. Revisao do complexa Cyrtorhinus Fieber—Me- comma Fieber (Hemiptera—Heteroptera—Miri- dae). Boletim do Museu Paraense Emilio Goeldi, (Nova Serie) (Zoologia) II: 1-72. Carvalho, J. C. M. and R. L. Usinger. 1960. New species of Cyrtopeltis from the Hawaiian Islands with a revised key (Heteroptera: Miridae). Pro- ceedings of the Hawaiian Entomological Society 17: 249-254. Gagne, W. C. 1968. New species and a revised key to the Hawaiian Cyrtopeltis Fieb. with notes on Cyrtopeltis (Engytatus) hawaiiensis Kirkaldy (Heteroptera: Miridae). Proceedings of the Ha- waiian Entomological Society 20: 35-44. Howarth, F. G. 1990. Hawaiian terrestrial arthro- 249 pods: An overview. Bishop Museum Occasional Papers 30: 4-26. Kirkaldy, G. W. 1902. Hemiptera. /n Sharp, D., ed., Fauna Hawaiiensis, 3(2): 93-174. Cambridge Uni- versity Press. 1904. Some new Oahuan (Hawaiian) He- miptera. The Entomologist 37: 174-179. Schuh, R. T. 1974. The Orthotylinae and Phylinae (Hemiptera: Miridae) of South Africa with a phy- logenetic analysis of the ant mimetic tribes of the two subfamilies for the world. Entomologica Americana 47: 1-332. . 1984. Revision of the Phylinae (Hemiptera, Miridae) of the Indo-Pacific. Bulletin of the Amer- ican Museum of Natural History 177: 1462. Slater, J. A. 1950. An investigation of the female genitalia as taxonomic characters in the Miridae (Hemiptera). Iowa State Journal of Science 25: 1- 81. Uchida, G. K. and J. W. Beardsley, Jr. 1988. Tax- onomy and biology of Megatrioza palmicola group (Homoptera: Psyllidae). Proceedings of the Ha- walian Entomological Society 28: 57-100. Wagner, W.L., D. R. Herbst, and S.H.Sohmer. 1990. Manual of the Flowering Plants of Hawai’i. Bishop Museum Special Publication 83. Zimmerman, E. C. 1948a. Insects of Hawaii. Vol. I. Introduction. University of Hawaii Press, Hono- lulu. 206 pp. 1948b. Insects of Hawaii. Vol. IV. Heter- optera. University of Hawaii Press, Honolulu. 255 pp. 1948c. Insects of Hawaii. Vol. lV. Homop- tera: Auchenorhyncha. University of Hawaii Press, Honolulu. 268 pp. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 250-257 NEW GENUS AND NEW SPECIES OF COLPURINI (HETEROPTERA: COREIDAE) FROM THE PHILIPPINE REPUBLIC HARRY BRAILOVSKY Departamento de Zoologia, Instituto de Biologia UNAM, Apdo Postal No. 701-53 Mexico, 04510 D.F. Abstract. —One new genus (Carvalhygia) and three new species (C. carvalhoi, C. milzae, and C. nigra) collected in the Philippine Republic are described in the tribe Colpurini (Coreidae). Habitus view illustrations and drawings of the male and female genitalia are provided to help distinguish these taxa. Key Words: ippine Republic Only three genera and ten species of Col- purini have been described from the Phil- ippine Republic. The genus Hygia Uhler contains three subgenera: Co/pura Bergroth with two species (obscuricornis (Stal) and pallidicornis (Stal)), Microcolpura Breddin with one species (denticollis (Bergroth)), and Sphinctocolpura Breddin with five species (dentifer (Stal), maculipes (Stal), obscuripes (Stal), pictipes (Stal) and punctipes (Stal)). The genus Homalocolpura Breddin includes one species (sorbax Bergroth) and the genus Typhlocolpura Breddin has one species (vul- canalis Bergroth). The present paper adds one new genus and three new species. Three features of this new genus are the reduced to absent post- ocular tubercle, the reduced hemelytra that exhibit a coleopteroid condition, and the absence of ocelli. The following abbreviations are used for the institutions cited in this paper: BMNH (The Natural History Museum, London, England); UNAM (Instituto de Biologia, Universidad Nacional Autonoma de Méx- ico, México D.F.); and USNM (U.S. Na- Insecta, Heteroptera, Coreidae, Colpurini, new genus, new species, Phil- tional Museum of Natural History, Wash- ington, D.C.). All measurements are given in millime- ters. This paper is written in honor of the late Dr. José Candido Melo Carvalho, in rec- ognition of his voluminous and seminal contributions on the Miridae of the world, as well as on other groups of animals and key papers on ethnical South American groups. His distinguished and productive career has long served as an example of ex- cellence. Carvalhygia Brailovsky, NEw GENUS Diagnosis. — Carvalhygia Brailovsky, new genus is similar and closely related to Ly- gaeopharus Stal and Typhlocolpura Stal. These three genera share several characters: ocelli reduced or absent, tylus apically glo- bose and truncated, antenniferous tubercle unarmed, hemelytral membrane reduced, and the female abdominal sternite VII with plica and fissura. Carvalhygia is easily distinguished be- cause it is the only known genus in the tribe Colpurini with the postocular tubercle ex- VOLUME 97, NUMBER 2 tremely reduced to absent, eyes globose, slightly protuberant and the paratergite IX thick and folded downward. Carvalhygia exhibits the frontal angles of pronotum rounded, not exposed or pro- duced forward as short or long conical teeth, buccula with a sharp middle projection, and femora armed. These are characters that may or may not be present in other genera, mak- ing the combination unique to this new ge- nus. In Lygaeopharus the frontal angles are produced forward as long conical teeth, the femora are unarmed and the buccula en- tirely rounded, without sharp middle pro- jection; in Typhlocolpura the frontal angles are produced forward as long conical teeth, the femora could be armed or unarmed, and the buccula always with a sharp middle pro- jection. Generic description. — Head: Longer than wide, pentagonal and dorsally flat or slightly depressed; tylus unarmed, apically globose, extending anteriorly to and laterally higher than juga; jugum unarmed, thickened and shorter than tylus; antenniferous tubercle unarmed, quadrate, robust, apex truncated; side of head in front of eye unarmed and obliquely straight; antennal segment I ro- bust, thickest, slightly curved outward, shorter than head; segments II and III cy- lindrical and slender; segment IV fusiform; segment II the longest, segment IV the shortest and III longer than I; ocelli absent; posterior pit between eyes deep and diag- onally excavated; eyes globose, slightly pro- tuberant, and based on an hypothetical line the upper margin is raised above the vertex and frontal area; postocular tubercle ex- tremely reduced to absent; buccula round- ed, elevated, short, not projecting beyond antenniferous tubercle, with a sharp middle projection; rostrum reaching posterior third of abdominal sterna III or IV; rostral seg- ment I reaching posterior margin of the gula. Thorax.— Pronotum wider than long, tra- peziform and bilobed; anterior lobe longer than posterior lobe, each margin convexly rounded, emarginated and moderately ele- 251 vated; collar wide; frontal angles rounded, not exposed or produced forward as short conical teeth; humeral angles rounded, not exposed; posterolateral border obliquely straight and posterior border straight or slightly concave; calli slightly convex, never separated along midline by longitudinal fur- row; posterior margin without a transverse ridge. Anterior lobe of metathoracic peri- treme elevated and reniform, posterior lobe sharp, small. Legs.— Ventral surface of femora armed with two subdistal short spines and few more scattered along ventral surface; tibiae cylin- drical and sulcate. Scutellum.— Triangular, flat, wider than long or as long as wide; apex acute. Hemelytra.—Coleopteroid; clavus and corium fused; wings do not overlap but meet along the midline; membrane reduced to a small flap, leaving the posterior abdominal terga exposed. Abdomen.—Connexival segments strong- ly elevated; posterior angle of each connex- ival segment complete, not extending on a short spine; abdominal sterna with medial furrow, projecting to anterior third of ster- nite Von V: Integument.—Body surface rather dull. Head, pronotum, scutellum, clavus, cori- um, thorax, abdomen, and exposed parts of genital segments of both sexes with circular grayish-white farinose punctures and each punctuation with short, decumbent golden or silvery bristlelike setae; antennae, legs, and abdominal sterna with few long, erect setae; antennae and legs minutely granulate. Male genitalia.—Genital capsule simple and globose; posteroventral edge entire, transversely straight, with lateral angles rounded (Fig. 3). Female genitalia.—Abdominal sternite VII with plica and fissura; plica triangular, reaching middle third of sternite VII; gon- ocoxae I with dorsal third closed and ventral third opened, slightly triangular and with the external apical angle globose; paratergite VIII short, square, with visible spiracle; par- 252 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Dorsal view of Carvalhygia carvalhoi, 8. atergite IX longer than VIII, rectangular, Type species.—Carvalhygia carvalhoi entirely thick and folded downward (Figs. Brailovsky, new species. 4-6). Etymology.—I am pleased to name this new genus for Dr. José Candido Melo Car- NEW SPECIES valho, distinguished Brazilian hemipterist, Figs. 1, 3-5 in recognition of his splendid contributions Description.—Measurements: Male: to the sciences. Gender feminine. Head length 2.12; interocular space 1.20; Carvalhygia carvalhoi Brailovsky, VOLUME 97, NUMBER 2 253 Fig. 2. Dorsal view of Carvalhygia milzae, °. width across eyes 2.08; preocular distance 1.24; length antennal segments: I, 1.64; II, 2.40; III, 1.96; IV, 1.52. Pronotum: Total length of anterior lobe 1.26; total length of posterior lobe 0.88; total width of anterior lobe 2.28; total width of posterior lobe 2.84. Scutellar length 1.04; width 1.12. Total body length 10.95. Female.—Head length 2.28; interocular space 1.24; width across eyes 2.24; pre- ocular distance 1.36; length antennal seg- ments: I, 1.76; II, 2.76; II, 2.04; IV, 1.64. 254 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bs) Figs. 3-6. Caudal view. 5. Lateral view. 6. Female genital plates in lateral view of Carvalhygia milzae. Pronotum: Total length of anterior lobe 1.16; total length of posterior lobe 1.00; total width of anterior lobe 2.56; total width of poste- rior lobe 3.20. Scutellar length 1.36; width 1.44. Total body length 12.10. Male.— Dorsal coloration black with red brown reflections on posterior third of pro- notum, clavus, and corium; antennal seg- ments I to IV pale to dull orange; antero- lateral margin of pronotum and costal mar- gin of corium dull orange to chestnut brown; hemelytral membrane dark brown with in- ner portion creamy yellow; connexival seg- ments III to VII red brown with posterior margin yellow; abdominal terga black with 6 3-5. Carvalhygia carvalhoi. 3. Male genital capsule in caudal view. 4—5. Female genital plates. 4. lateral margin of VII yellow. Ventral col- oration black with following areas orange, with or without chestnut reflections: rostral segments I to IV, anterior and posterior lobe of metathoracic peritreme and pleural mar- gins of the abdominal sternites IV to VII; coxae black, trochanters yellow with chest- nut reflections, femora dull orange with bas- al join yellow, tibiae dull orange with two diffuse yellow rings, one subbasal, other one near middle, and tarsi dull orange with chestnut reflections. Male.— Rostrum reaching middle or pos- terior third of abdominal sternite IV; scu- tellum wider than long; hemelytral mem- VOLUME 97, NUMBER 2 brane reduced and reaching middle third of abdominal tergite IV. Female.—Coloration: Similar to male. Connexival segments VIII and IX red brown with posterior margin yellow; abdominal terga VIII and IX black. Gonocoxae I black with internal apical angle yellow; paratergite VIII and IX red brown with anterior third yellow. Genital plates. Paratergite [IX con- spicuously folded downward (Figs. 3-5). Variation.—The type material exhibits some color variation on most specimens: 1, antennal segment I red brown with basal third yellow; 2, antennal segment IV red brown; 3, femora red brown with dull or- ange reflections; 4, pleural margins of ab- dominal sterna III to VII red brown with dull orange reflections. Type material.— Holotype: 6, Philippine Republic, Sibuyan Island, Col. Baker (with- out additional data) (USNM). Paratypes: 3 66, 5 99, Philippine Republic, Sibuyan Is- land, Col. Baker (without additional data) (USNM, UNAM and BMNH). Etymology.—I am pleased to name this new species after the late Dr. José Candido Melo Carvalho. Carvalhygia milzae Brailovsky, NEw SPECIES Figs. 2, 6 Description.— Measurements: Female: Head length 2.20; interocular space 1.22; width across eyes 2.16; preocular distance 1.30; length antennal segments: I, 1.96; II, SF ih 2°32: 1V. 1.64. Pronotum: Total length of anterior lobe 1.12; total length of posterior lobe 0.96; total width of anterior lobe 2.16; total width of posterior lobe 2.64. Scutellar length 1.22; width 1.22. Total body length 12.00. Female.— Dorsal coloration: Head and anterior lobe of pronotum dark red brown; posterior lobe of pronotum bright orange red; scutellum, clavus and corium dull or- ange red, with apical margin of corium mostly dark brown; connexival segments III to IX pale orange brown with posterior mar- 255 gin yellow; abdominal terga red brown; an- tennal segment I bright orange; segment II dull orange with basal third pale yellow; seg- ment III dull orange and IV dark brown with subbasal yellow ring; collar, anterolat- eral margin of pronotum, apex of scutellum, and costal margin of corium bright to pale orange with yellow reflections; hemelytral membrane pale yellow. Ventral coloration: Head and prothorax dark red brown; me- sothorax, metathorax, abdominal sternite and genital plates pale red brown; rostral segment I yellow and segments II to IV bright chestnut orange; coxae and femora bright orange red; trochanters yellow; protibiae pale orange with two yellow rings one subbasal, other one near middle; mesotibiae pale or- ange with yellow reflections; tarsi pale or- ange with yellow reflections (posterior leg absent); anterior and posterior lobe of me- tathoracic peritreme bright yellow with or- ange reflections; pleural margin of abdom- inal sternite III to VII, posterior edge of abdominal sterna IV to VII, and internal apical angle of gonocoxae I yellow. Rostrum reaching anterior third of ab- dominal sternite IV; scutellum as long as wide; hemelytral membrane reduced and reaching posterior third of abdominal ter- gite IV. Genital plates. Paratergite IX slight- ly folded downward (Fig. 6). Type material.— Holotype: 2, Philippine Republic, Panay Mt. (Madaas), Col. W. Schultze (without additional data) (BMNH). Etymology.—Named for Milza, Carval- ho’s wife and indefatigable partner. Carvalhygia nigra Brailovsky, NEw SPECIES ig: 7 Description.— Measurements: Female: Head length 2.04; interocular space 1.22; width across eyes 1.86; preocular distance 1.16; length antennal segments: I, 1.72; II, 2.52; III, 2.00; IV, absent. Pronotum: Total length of anterior lobe 1.12; total length of posterior lobe 0.72; total width of anterior lobe 1.88; total width of posterior lobe 2.40. 256 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘a *\ ‘WA +{* =A iy ‘ Fig. 7. Dorsal view of Carvalhygia nigra, 9. Scutellar length 1.00; width 1.08. Total body length 11.00. Female.—Dorsal coloration black with following areas pale dirty yellow: anterolat- eral margin of pronotum and costal margin of corium; antennal segment I dark red brown, segments II and III dull orange with basal third yellow; segment IV absent: hemelytral membrane dirty yellow; connex- See Sees SS kre OS oh ival segments III to IX red brown with pos- terior margin yellow; abdominal terga black with intersegmental scars IV-V and V-VI pale orange yellow. Ventral coloration black with following areas yellow, with or without chestnut reflections: rostral segments I to IV, anterior and posterior lobe of metatho- racic peritreme, and posterior margin of pleural margins of abdominal sternite IV to VOLUME 97, NUMBER 2 VII; gonocoxae I black with internal apical angle yellow; paratergite VIII and IX black with upper margin yellow; coxae black; tro- chanters yellow; femora yellow, densely covered with dark red brown discoidal spots and with posterior third or posterior half and spines dark red brown; tibiae yellow with basal join and apical third dark red brown; tarsi yellow with chestnut reflec- tions. Rostrum reaching posterior third of ab- dominal sternite III; scutellum wider than long; hemelytral membrane extremely re- duced and reaching middle third of abdom- inal tergite III. Genital plates. Paratergite IX slightly folded downward. Type material.— Holotype: 2, Philippine Republic, Mindoro, Baco River, January— February 1910, J.J. Mounsey (BMNH). Etymology.—From the Latin nigra, meaning black. KEY TO THE KNOWN SPECIES OF CARVALHYGIA 1. Hemelytral membrane dark brown with inner portion creamy yellow (Fig. 1) ............. OE, ee carvalhoi Brailovsky, new species — Hemelytral membrane entirely yellow (Figs. 2, 1) wale tec te Re =a ie a he sot ea rant a 2 2. Hemelytral membrane extremely reduced, reaching middle third of abdominal tergite III; rostrum reaching posterior third of abdominal sternite III; profemora and mesofemora yel- low, with posterior third or posterior half, as well as several discoidal spots, dark red brown; scutellum wider than long ................. Sis Ocha tetas nks ena rere nigra Brailovsky, new species 257 — Hemelytral membrane never extremely reduced, reaching posterior third of abdominal tergite IV; rostrum reaching anterior third of abdominal sternite IV; profemora and mesofemora uni- formly bright orange red; scutellum as long as WIG arena aso) milzae Brailovsky, new species ACKNOWLEDGMENTS I thank the following colleagues and in- stitutions for the loan of specimens and oth- er assistance relevant to this study: Janet Margerison-Knight (BMNH); Richard C. Froeschner and T. J. Henry (USNM). Spe- cial thanks to Ernesto Barrera (UNAM) and Cristina Urbina for the preparation of the illustrations and to the Consejo Nacional de Ciencia y Tecnologia México (CONACYyT) for financial assistance to visit The Natural History Museum in London. LITERATURE CITED Bergroth, E. 1916. New and little known heteropter- ous Hemiptera in the United States National Mu- seum. Proceedings of the United States National Museum 51(2150): 215-239. 1918. Studies in Philippine Heteroptera I. Philippine Journal of Science 13(2): 43-126. Breddin, G. 1900. Materiae ad cognitionem subfam- ilae Pachycephalini (Lybantini olim). Ex Hemip- teris-Heteropteris, Fam. Coreidae. Revue d’ En- tomologie Caen 19: 194-217. Stal, C. 1870. Hemiptera Insularum Philippinarum. Ofversigt af Kongliga Vetenskaps-Akademiens Foérhandlinger 7: 607-776. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 258-330 A RECLASSIFICATION AND PHYLOGENY OF THE TERMATOPHYLINI (HETEROPTERA: MIRIDAE: DERAEOCORINAE), WITH A TAXONOMIC REVISION OF THE AUSTRALIAN SPECIES, AND A REVIEW OF THE TRIBAL CLASSIFICATION OF THE DERAEOCORINAE GERASIMOS CASSIS Australian Museum, 6-8 College Street, P.O. Box A285, Sydney South, N.S.W. 2000, Australia. Abstract.—The genera of Termatophylini are described, including two new genera, Democoris and Kundakimuka, from Australia. A key 1s given to the termatophyline genera. The type species of all the genera are described, and six new species of Australian ter- matophylines, Kundakimuka carvalhoi, K. queenslandica, Democoris lugens, D. leptocytus, Termatophylum melaleucae and T. weiri, are described. A key is given to the Australian termatophyline species. The tribe is re-defined on the basis of the short first labial segment, the presence of antero-lateral pronotal setae, and the exposed metathoracic spiracle. The genera Conocephalocoris, Hesperophylum and Termatomiris are removed from the Ter- matophylini and placed in the Deraeocorini. A cladistic analysis of the Termatophylini is provided on the basis of 29 morphological characters. The sister-group relationships are discussed and the characters are reviewed. Characters of the pterothoracic pleura, particularly the metathoracic spiracle and the scent efferent system of the metathoracic glands are extensively utilized in this study. These characters are also discussed for other suprageneric groups of Miridae. The tribal classification of the Deraeocorinae is reviewed, and it is concluded that none of the other tribes are adequately defined. It is also firmly established that the termatophylines are mirids, and that any similarity with the Antho- coridae is due to convergence. Key Words: Heteroptera, Miridae, Deraeocorinae, Termatophylini, revision, phylogeny, new genera, new species, Australia The Termatophylini have had a labile po- sition within the classification of the Cim- icomorpha, varying from family status (Reuter 1910, Wagner 1970), as a subfamily within the Anthocoridae (Reuter 1884a, b) and Miridae (China and Myers 1929), and more recently as a tribe of the mirid sub- family Deraeocorinae (Carvalho 1952, 1955a, 1957). Mention has been made of their salient similarity with some members of the Anthocoridae (Schuh 1976). Curi- ously, after Carvalho (1952) placed them in the Deraeocorinae, there has been little dis- pute as to their monophyly nor their fa- milial or subfamilial position. This view is supported in this work despite the present conclusion that the previous tribal defini- tion, based primarily on a porrect head, ne1- ther holds for the tribe, nor is exclusive for other Deraeocorinae. This work was initiated by the receipt of a box of mirid specimens from the Museum and Art Galleries of Northern Territories, amongst which, one specimen of a terma- VOLUME 97, NUMBER 2 tophyline from Smith Point, near Darwin, brought forth the question, “‘why is this mir- id so like an anthocorid?” This exaggerated specimen has gigantic hind femora, some- what like those found in members of the Halticini (Miridae: Orthotylinae). Aside from this apomorphy, the similarity of its overall facies with anthocorids is striking, and the absence of ocelli and presence of a hemelytral membrane cell, suggested an al- ternative taxonomic arrangement. Charac- ters of the male genitalia and the presence of femoral trichobothria strongly support the placement of the termatophylines with- in the Miridae. The anthocorid-like fea- tures, however, provided the impetus for making an analysis of the termatophylines, mindful of present definitions of the Miri- dae, Anthocoridae and other cimicomor- phan families (Schuh and Stys 1991). In this work I also support the placement of the termatophylines in the Deraeocori- nae, but in redefining them as a monophy- letic tribe, I have discovered that the other deraeocorine tribes are presently inade- quately defined. I have given a commentary on the characters presently used in these classifications and suggest alternative char- acters, particularly of the pterothoracic pleura. In particular I have introduced a morphological comparison of the metatho- racic spiracular opening and the scent ef- ferent system of the metathoracic glands in the Miridae, to reappraise previous char- acters, such as the basally cleft claws, pri- marily used to place the termatophylines within the Deraeocorinae. An invitation to provide a contribution to a Festschrift for Dr. José Carvalho, ren- dered this idea into its present form. The Termatophylini have not been previously recorded from Australia. In accumulating all the Australian termatophyline material it was clear that not all termatophylines were anthocorid-like, which led me to study the Australian fauna, redescribe the genera worldwide, including a checklist of species, phylogenetic definition and position of the 259 termatophylines, and a cladistic analysis of the genera. Jose Carvalho’s contributions on the Miridae, including the Australian fauna, remains unparalleled. Cassis and Gross (in press) report that Carvalho de- scribed more than half of the Australian mirid fauna. He had also commenced nu- merous projects on the Australian Miridae, and much of the termatophyline material in this work has passed through his hands, including Democoris lugens n. gen. and n. sp. I have provided a dedication to him by naming the unique Smith Point species in his honour, under the new generic name, Kundakimuka. In this work I have recognized eight gen- era and 30 species of Termatophylini. The genus Democoris is restricted to south tem- perate Australia. The genera Kundakimuka and Termatophylum Reuter are broadly dis- tributed in the Eastern Hemisphere, includ- ing Australia. Termatophylella Carvalho, Termatophylidea Reuter and Poppius, Ter- matophyloides Carvalho are restricted to the Western Hemisphere. Termatophylina Car- valho is restricted to the Oriental region. Arygrotelaenus Reuter and Poppius is re- stricted to the Middle East. I have provided a key to the world genera, which is based entirely on external characters, and a key to all known Australian species. In redefining the Termatophylini I have transferred Con- ocephalocoris Knight, Hesperophylum Reu- ter and Poppius and Termatomiris Ghauri to the Deraeocorini. MATERIALS AND METHODS About 110 specimens of termatophylines were examined in this study. This material was borrowed from the Natural History Museum, London and the National Muse- um of Natural History, Smithsonian Insti- tute, Washington, and the major collections found within Australia. The specimens of the type specimens were examined. No males of Arygrotelaenus elegans Reuter and Poppius were available for examination. The generic definitions incorporate observa- 260 tions of most termatophyline species. Not all species of Termatophylum and Terma- tophylidea were examined, but the descrip- tions in the literature were sufficient to con- fidently retain within them all previously assigned species. The male genitalic characters were ex- amined on temporary mounts in glycerin, using both a Zeiss SR Stereomicroscope and a Leitz Laborlux S Compound microscope. The line drawings were prepared using a camera lucida. Scanning electron micro- graphs were made from dried museum spec- imens, using a Cambridge S120 micro- scope. TAXONOMIC CHARACTERS The morphological terminology used in this work is derived from various sources. Many of the terms used for somatic char- acters are now conventional, following the modern works of authors such as Schuh (e.g. 1984). Some original interpretation was in- volved in my treatment of the forewing, the pterothoracic pleura and the male genitalia. In this work I introduce the use of com- parative morphology of the pterothoracic pleura, particularly the condition of the me- tathoracic spiracle and the scent efferent system of the metathoracic scent glands. The homologies are partly derived from Cara- yon (1971) and Staddon (1979) for the Het- eroptera, and Larsen (1945) and Southwood (1953) for the Miridae. The junction of the mesepimeron and metepisternum contains in some Miridae, a visible opening of the metathoracic spiracle (Figs. 57, 65). Its po- sition and associated structures are both phylogenetically and taxonomically signif- icant (Cassis 1984, Cassis in preparation). The external cuticular component of the metathoracic scent glands yields a wealth of systematic characters. The terminology of this system remains greatly confused, and requires standardization across the Heter- optera. Staddon (1979) has referred to this system as the scent efferent system, which I have adopted in this work. His nomen- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON clature of the scent efferent system com- ponents has not proven as useful, and I have preferred to use a translation of some of the terminology of Carayon (1971). I refer to the gutterlike channel which emerges from the ostiole as the peritreme. This structure is usually tonguelike and extends to the lat- eral margins of the metepisternum. Under light microscopy it often appears shiny, whereas under scanning electron micros- copy it is usually covered with microsetae. The peritreme 1s surrounded by highly dis- sected cuticle, which has a granulate ap- pearance and is referred to as the evapo- rative areas. The evaporative areas are com- posed of numerous components, and in this work I refer chiefly to the evaporative caps, which are raised and flattened. The char- acters of the scent efferent system are la- beled in Figs. 57, 119 and 123. I have in- cluded in the Phylogenetic analysis section below an evaluation of these characters in other Miridae. The forewing venation of the termato- phylines has been used extensively in this work, both as diagnostic and phylogenetic characters. The terminology and homology of the heteropteran forewing has been partly resolved by Tanaka (1926), Wootton and Betts (1986), and Betts (1986). Carver et al. (1991) refer to the posterior margin of the embolium in the Miridae as the R+M vein, whereas Knight (1941) considered this to be the costal vein. I accept the former inter- pretation and concur with Betts (1986) that the vein anteriad to the R+M vein is the subcostal vein, which is evident in some Miridae. In this work I refer to the groove posteriad to R+M vein as the median flex- ion line. Knight (1941) referred to this groove as the radial vein, but its appearance suggests a flexion line, rather than a tra- cheate vein. Its position, posteriad to the posterior margin of the embolium, elimi- nates its interpretation as a radial (or R+ M) vein. The median flexion line cannot be in- terpreted as the CuA vein, which according to Betts (1986), is an uninterrupted vein, VOLUME 97, NUMBER 2 parallel and adjacent to the claval furrow. The vein of the clavus is referred to as the first anal vein by Betts (1986), which I have adopted in this work, and is synonymous with the claval vein of Knight (1941). Other workers, such as Davis (1961), have pro- duced venational terminologies for the Het- eroptera, but it is beyond the scope of this paper to contrast in detail the various sys- tems. The characters of the forewing are la- beled in Fig. 47. The morphology of the male genitalia of the Deraeocorinae has received little atten- tion. Kelton (1959) gave a comparative morphological account, and reported con- siderable diversity in form throughout the subfamily. He stated that the most distinc- tive features included the flexible ductus seminis, and the bulbous spiculate process- es of the membranous vesica. Schwartz (1987) in a review of Mirinae and Deraeo- corinae male genitalia suggested that the secondary gonopore was of critical impor- tance, and that it differed in the two sub- families. He reported that the secondary gonopore of the Deraeocorini terminated adjacent and usually basad of the mem- brane, and that the aperture is large and opens into a depressed trough, often with associated serrate sclerotized plates. He also stated that the secondary gonopores in the other Deraeocorinae tribes were smaller than those found in the Deraeocorini. Schwartz (1987) reviewed the terminology of the ves- ical processes used in previous works (Kel- ton 1959, Clayton 1982, Stonedahl 1983), suggesting that they were too simplified. Stonedahl (1988) differentiated two types of vesical processes in the mirine genus Phy- tocoris Fallén, as follows: (1) sclerotized processes (equivalent to the ribbonlike strap of Schwartz (1987)) which originate at or near the level of the gonopore, which is sometimes continuous or associated with the basal sclerite (originating basad to the secondary gonopore and not present in the Deraeocorinae examined), and variously at- tached to the membranous sac; (2) lobal 261 sclerites (Schwartz 1987) which originate at or distally on the membranous lobes, and as simple outgrowths of the membrane cu- ticle (equivalent to spicules or spiculae of other authors). Stonedahl and Cassis (1991) in a revision of the deraeocorine genus Fingulus Distant described the male genitalia, and reported homologies and terminologies that have ap- parent general value in the Deraeocorinae, including the Termatophylini. They de- scribed the male genitalia of Fingulus as possessing a basal tubular, sclerotized skirt surrounding the distal region of the ductus seminis, a simple secondary gonopore, and a distal multilobed membranous sac with lobal sclerites and/or fields of spines. They found no evidence of sclerotized processes or basal processes. In the Termatophylini the ductus seminis terminates at the base of the membranous component of the vesica (Figs. 111-118), as reported for the Deraeocorini by Schwartz (1987). The secondary gonopore is small, simple and undifferentiated, as is found in Fingulus. Aside from the genus 7ermato- phylidea, the gonopore is associated with a differentiated basal portion of the membra- nous sac, which can be recognized as a V or U-shaped cavity marked by compact fields of spines (Figs. 111-113, 115-118), and is here referred to as the gonoporal cavity. It is difficult to ascertain in the Termatophy- lini whether the sclerites which are associ- ated with the gonoporal cavity are lobal sclerites (Figs. 111-113, 115-118), or rep- resent a unique type of vesical process. They are clearly outgrowths of the membranous cuticle, but are never found on the periphery of the membranous lobe. This could not be clarified because of the limited material available for examination. In the Terma- tophylini, the vesica is small and appears to be unilobed, and does not show any ob- vious inflation as is evident in other Der- aeocorinae and Mirinae. I have tentatively referred to these processes as lobal sclerites pending further study. The distal regions of 262 the membranous sac often have fields of spines (Figs. 111, 116-118). The vesica of Termatophylidea is unlike any other termatophyline, in possessing a sclerotized process connected to the base of the ductus seminis and extending to near the apex of the membranous lobe. It also lacks any obvious gonoporal cavity and there are no lobal sclerites or fields of spines. This is reminiscent of the condition found in the dicyphine genus Singhalesia China and Carvalho and in some species of Hyaliodes Reuter. The latter genus and Termatophy- lidea are superficially similar, but are clearly differentiated on characters of the head and the pterothoracic spiracle, and the presence of a sclerotized process in both genera is considered to be independently derived. Stonedahl (personal communication) has indicated that the sclerotized process 1s also present in the clivinemines, Ambracius Stal and Bothynotus Fieber. PHYLOGENETIC METHODS The phylogenetic relationships of the ter- matophyline genera were examined using cladistic methods (Wiley 1981). The clado- gram was produced, using in tandem, the computer software HENNIG86 and CLA- DOS. These programs are designed for pro- ducing cladograms of minimal length, and the manipulation of the character data. The character data matrix was built in a text editor and transferred to HENNIG86, where tree building algorithms were applied to the input file. The final result was produced by using the Branch and Bound option, where the data were treated in a stepwise manner. Choice between multiple trees of equal length was made by reweighting characters according to the rescaled consistency index. The characters were coded as either bi- nary or multistate characters. Most of the multistate characters were ordered, except for characters 4, 23 and 27 (see Table 2), which were coded as non-additive, because there was no evidence of transformation se- ries. Character state polarization was PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON achieved through outgroup comparison. The selection of outgroups was made difficult by current deficiencies in the tribal classifica- tion of the Deraeocorinae (Akingbohungbe 1974, 1978, Stonedahl and Cassis 1991). At present six tribes are recognized; Clivine- mini, Deraeocorini, Saturniomirini, Hy- aliodini, Surinamellini and Termatophyli- ni. A selection of genera from each of the tribes, aside from the ant-mimetic Surina- mellini, were initially included in the anal- ysis, to examine the distribution of char- acters throughout the Deraeocorinae. This produced cladograms with low consistency indices, and the outgroups were then re- stricted to two members of the Clivinemini (Ambracius and Bothynotus), and the genera Conocephalocoris and Hesperophylum, which previously had been included in the Termatophylini. The last two genera show many similarities with the Termatophylini, as presently defined, particularly in the head and hemelytral structure, and are consid- ered as sister-groups. The Clivinemini gen- era were also included in adherence to mul- tiple outgroup principles (Maddison et al. 1984), and were selected above other der- aeocorines because of some structural sim- ilarities with termatophylines, particularly in the form of the callosite region of the pronotum. Genera of the Hyaliodini (An- nona Distant and Hyaliodes Reuter), ini- tially included in the analysis, are similar to some Termatophylini, particularly the ge- nus Termatophylidea, but were excluded because they provided no additional clari- fication to any of the character state trees. The male genitalic characters were not ex- amined in any of the outgroups because of a lack of specimens. Stonedahl (personal communication) provided information on the male genitalia of some of the outgroup taxa. ACKNOWLEDGMENTS I thank the following individuals and in- stitutions for providing the specimens that were used in this study. Gary M. Stonedahl, VOLUME 97, NUMBER 2 International Institute of Entomology, Lon- don, Michael Webb, Natural History Mu- seum, London (NHM), Thomas J. Henry, Systematic Entomology Laboratory, ARS, USDA, % National Museum of Natural History, Washington, D.C. (USNM), John F. Donaldson, Queensland Department of Primary Industry, Brisbane (QDPI), Gra- ham Brown, Museum and Art Galleries of the Northern Territories (NITIMAG), Jan Forrest, South Australian Museum, Ade- laide (SAMA), and Tom A. Weir, Austra- lian National Insect Collection, Canberra (ANIC). Carmen Zurl provided the habitus illustrations of Kundakimuka queenslandi- ca and Democoris lugens, and assisted in the production of the photographic plates. Sue Lindsay and Geoff Avern prepared the scanning electron micrographs. Mary Carv- er of the Australian National Insect Collec- tion provided useful advice on forewing ve- nation. Gary Stonedahl assisted in many ways in this work and his prompt delivery of critical specimens, particularly of Con- ocephalocoris nasicus and Hesperophylum heidemanni, was vital. Both Gary Stone- dahl and Toby Schuh provided useful re- views of this paper. The Australian Biolog- ical Resources Study provided a grant to produce this work, and their continued sup- port of mirid taxonomy is duly acknowl- edged. Lastly, I would like to thank Tom Henry and Al Wheeler for the opportunity to contribute to the Festschrift. HISTORICAL REVIEW The first termatophyline, Termatophy- lum insigne, was described by Reuter (1884a), who recognized this species as unique, and erected a new subfamily, the Termatophylina, of Anthocoridae for it. He diagnosed the group on the basis of the four- segmented labium, including the short first segment, the absence of ocelli, hemelytral structure, lack of a hamus in the hindwing, and the three-segmented tarsi. He distin- guished it from another anthocorid subfam- ily, the Microphysina, on hemelytral and 263 tarsal characters. Reuter (1884b) in his monograph of the anthocorids, recognized three subfamilies, Anthocorina, Microphy- sina and Termatophylina, and provided a key to the subfamilies, differentiating the two latter subfamilies from the Anthocorina by the four-segmented labium. Subsequent- ly, Reuter (1910) in his review of the sys- tematics of the Miridae, recognized the ter- matophylines as a family, separating them from the mirids by the presence of an em- bolium. Poppius (1910) supported the fam- ily status, in describing a new Termatophy- lum species from the Ethiopian Region. Reuter and Poppius (1912) in the only previous comprehensive paper on the ter- matophylines retained the suprageneric group as a family, giving a detailed descrip- tion, and distinguishing them from the An- thocoridae. They made mention of the por- rect head and the short first labial segment, but erroneously concluded that the pretarsi lacked arolia (= parempodia). Schuh (1976) in a review of the mirid pretarsus has shown that setiform and fleshy parempodia are ho- mologous. Early European workers, such as Reuter and Poppius (1912), regarded fleshy parempodia as the only true “‘arolia,” and their conclusion that termatophylines lack “‘arolia” is a misinterpretation of setiform parempodia. Reuter and Poppius (1912) de- scribed three new genera, Arygrotelaenus, Hesperophylum and Termatophylidea, and seven new species. They also included a key to the four known termatophyline genera. Parshley (1923) described the family, large- ly repeating Reuter and Poppius’ (1912) di- agnosis, but curiously suggested that ter- matophyline genitalia were ““much as in the Anthocoridae.” China and Myers’ (1929) seminal paper on the classification of the cimicoid fami- lies, reappraised the relationships of the ter- matophylines, and placed them within the Miridae. They concluded that apart from the anthocorid-type head and the short first labial segment, the termatophylines were conclusively mirids, and treated them as a subfamily. They indicated that the porrect 264 head condition occurs in the Cylapinae, and noted its similarity to termatophylines. Car- valho (1951, 1952) analyzed the position of the termatophylines, and concluded that they were a tribe of Deraeocorinae on the basis of the basally cleft claws and setiform parempodia. He differentiated them from other deraeocorines by the porrect head, large eyes, short antennae, and small body size. Carvalho (1955a) provided a key to the termatophyline genera, recognizing sev- en genera, Hesperophylum, Termatophyli- dea, Arygrotelaenus, Conocephalocoris, Termatophylella, Termatophyloides and Termatophylum. He distinguished the gen- era mostly on head and pronotal characters, and the short antennae. Aside from Wagner (1970), who retained the termatophylines as a family, no subse- quent author has deviated from the tribal placement of this group within the Deraeo- corinae. Other works mostly include de- scriptions of new genera and species. Car- valho (1955b) described new taxa from the Neotropical region, recognizing two new genera, Termatophyloides and Termatophy- lella, from Central America, and two new species of Termatophylidea. Miyamoto (1965) gave detailed descriptions of two new termatophylines from Japan, providing ad- equate genitalic and habitus illustrations. Maldonado (1970) reviewed the distinctive genus Termatophylidea, describing three new species, with a key to all known species. Carvalho (1988) described a new genus, Termatophylina, from India, and gave il- lustrations of the male genitalia. Linnavuori (1974) described a new species of Terma- tophylum from Nigeria, and gave a key to the African species. Poppius (1910, 1915), China (1929), Usinger (1935), Carvalho (1983) and Ren (1983) described the other species. BIOLOGY The little that is known of the biology of Termatophylini suggests that the species are predators (Callan 1975). In Table | the bi- ological information that is known is listed, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON including prey and plant associations. This information was taken from the literature and label data. Myers (1935) was the first to report termatophylines as predators, de- scribing Termatophylidea pilosa as an im- portant predator of the cacao thrips, Selen- othrips rubrocinctus (Giard), a serious pest of cacao in the tropical parts of the Neo- tropical region. Callan (1943, 1975) and van Doesburg (1964) recorded other Termato- phylidea species as predators of thrips. Both T. maculata and T. opaca are also predators of the cacao thrips, and Callan (1975) re- ported that the former species also feeds on the larvae of the grass thrips, Calliothrips insularis (Hood). There are indications that other termatophylines feed on moth larvae. Kundakimuka queenslandica feeds on the xyloryctine moth, Xylorycta luteotactella (Walker), which feeds on a paperbark spe- cies, Melaleuca integrifolia. | have exam- ined specimens of Termatophylina indiana from the Natural History Museum, which has the label data: “‘associated with larval galleries of Lamida moncusalis,” which suggests that termatophylines may com- monly feed on moth larvae. The termatophylines are found in a va- riety of habitats, including flowers, foliage and moth larval galleries. Five termatophy- line species, Kundakimuka queenslandica, Termatophyloides pallipes, T. pilosulus, Termatophylidea maculata and T. opaca, have been found on leaves, with the last two species restricted to the underside of leaves (van Doesburg 1964, Callan 1975). Two species of Termatophylum, T. hikosanum and 7. melaleucae, have been collected from flowers. Van Doesburg (1964) reported that T. opaca and its prey were found under fine spider webs. The association of Termato- phylina indiana with moth larval galleries, suggests that termatophylines may be com- monly encountered in sheltered microhab- itats. The prey of Kundakimuka queenslan- dica, Xylorycta luteotactella, is also known to live in small tunnels, which the moth bores in the branches of their food plant (Common 1990). Also of interest is the fre- VOLUME 97, NUMBER 2 quency with which termatophylines are found at light. Five species of the genera Kundakimuka, Termatophylidea and Ter- matophylum have been taken at light or in light traps. This suggests that the termato- phylines are nocturnal predators. This hy- pothesis is morphologically supported by the presence of very large eyes, with enlarged ommatidia. There is no evidence of TJer- matophylina indiana being collected at light, which is the only termatophyline taxon that does not have enlarged ommatidia, and the eyes are of moderate size. Termatophylines are known from a wide variety of plants. They have been recorded from three monocot families, the Gramin- eae, Musaceae and Orchidaceae, and seven dicot families, the Anacardiaceae, Betula- ceae, Bombacaceae, Compositae, Legumi- nosae, Myrtaceae and Sterculiaceae (Table 1). The association of termatophylines with monocots is restricted to the Neotropical genera Termatophylella, Termatophylidea and Termatophyloides. There is no apparent overall pattern of plant association in the termatophylines, with some species having multiple plant associations, and 7ermato- phylidea opaca known from both a monocot and dicot. Some sympatric termatophyline taxa are found on the same plant genera: Arygrotelaneus simoni and Termatophylum insigne are both known from Acacia species, and Kundakimuka queenslandica and Ter- matophylum melaleucae are known only from Melaleuca species. SYSTEMATICS Tribe Termatophylini Termatophylina: Reuter 1884b: 167; Reu- ter 1884a: 218 (subfamily of Anthocori- dae). Termatophylidae: Reuter 1910: 70 (family status); Poppius 1910: 56; Reuter and Poppius 1912: 2 (diagnosis); Reuter 1912: 52 (key; diagnosis); Parshley 1923: 665 (description); Wagner, 1970: *** (foot- note). 265 Termatophylinae: China and Myers 1929: 97 (subfamily of Miridae). Termatophylini: Carvalho 1951: 133 (tribe of Deraeocorini); Carvalho 1952: 32, 42, 43, 50 (checklist); Carvalho 1955a: 22 (generic key); Carvalho 1957: 34 (cata- logue). Diagnosis.— The Termatophylini are best diagnosed by the short first labial segment (Figs. 19-20) which does not extend beyond the posterior margin of the bucculae. The head is moderately (Figs. 11, 14, 16, 17) to strongly porrect (Figs. 12, 13, 15, 18), with the apex of the clypeus in lateral view ter- minating above or at level of the antennifers (Figs. 11-18). Unlike any other Deraeocor- inae the metathoracic spiracle is external (= visible) as an oval or lanceolate opening, usually bounded by evaporative bodies on the dorso-caudal arm of the dorsal margin of the mesepimeron (Figs. 65, 67, 68). The pronotum has antero-lateral projecting se- tae on the anterior angles of the callosite region (Figs. 32-34). Description.—Macropterous, either smooth or coriaceous; ovoid (Fig. 1), elon- gate-ovoid (Fig. 2), or parallel-sided; mostly with rows of punctures on pronotum (Figs. 31, 33, 34, 38-42) and hemelytra (Figs. 47, 51-56); sparse to dense cover of setiform vestiture, rarely intermixed with adpressed, flattened scalelike setae. Head: moderately (Figs. 11, 14, 16, 17) to strongly (Figs. 12, 13, 15, 18) porrect; elongate (Figs. 4, 5, 7, 10) or transverse (Figs. 3, 6, 8, 9); vertex rounded, flattened, or narrowly bicom- pressed, posterior margin sometimes weak- ly carinate; frons moderately to strongly narrowed in front, barely to weakly decli- vent; clypeus short, weakly to moderately declivent, terminating above (Figs. 12, 13, 15) or at level of antennifers (Figs. 11, 14, 16, 17, 18) in lateral view; bucculae usually short and terminating caudally at or just beyond antennifers, most often very nar- row, with lateral margins either subparallel (Figs. 20, 22, 23), weakly to moderately ar- cuate (Figs. 19, 21, 26) sometimes strongly PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 266 EE a aE SE ay ee B1EP [2QP| B1ep [2qP| 6861 Wonaeuury S961 CLOWRAIN Qss6l ouTRAIeD eB1ep [9qe| cL6l uRTeD ‘Sol SIOAW CL6I UPl -[&D ‘p96 Binqss0q uPA qcsol oueAre) CL6I URTIRD Se6l J9duIS/) SL6I “€P6l UBIRD OL61 OpeuopleW OL61 OpeuopleW OL61 OpeuepleW Qcs6l OU4feAIe) B1ep [9qe| 696] OIOWPATIPY B1ep [9qe] 6861 ‘SL6] Wonaeuury JOUIIIJIY [aeaoeLIAJ] DiAdauanbuinb ‘W ‘pyofisasajul vona[vjaw [aesourmmnsaq] ‘ds piovop [aeoo -esny] ‘ds psnpy se20eplyoiO (9v1 -IsOdwio)] DIDIYIDAG DIUOUIAA deOUIWIRID [avaoelnjog] DIDUIWINID Snu] [avaoerpseoeuy] 2/0) -uap1990 wnipépopUuYy “[aeaoe -I[ND191$] OVIDI DUWOAGOIY [ [avaovoequiog] S7usisul DAIYIDd aed0ePIYIO [ovo -BLIAYA]] D/Ofiusajul DINaVDjAW [aesouruinsaq] ‘ds piIovop uONeIOOSsy 1URId JSOIOJUILI UT IYBI] 12 1SOIOJ -UIBI Ul 14ST] 1B SoWIZ0BI UO WIOSSO]q 991] UO sqioy uo [e1onidopidoq] SIJDSNIUOU DPIWUVT JO Sat -I9]]e3 [PAIL] YIM poyeroosse SnoLiesos3y, SdARI] JO OpIsiopun uo SIAR JO BPIsIopuN uo 1st] 1e 1yst] 18 ade IO} UO sqioy 3urdsoms 1ysty 1e siqey pure sieugey [eroidoueshy [] SNJIUIIOAGNA SA14YIOUIIS [ersidoueshy [] SNJIUIIOAGNA SA14YJOUIIS [e13}dou -esky [] Siupjnsul sdiysjor]D) [erojdoueshy]] SNJOUIIOAGNA SA1dYJOUIJIS [e1njdop -1daq] vjjajavjoamn] DIIAAOjAX Aaig turAydoveula | iam wnjdAydojouLsa | aponapjau unjAydojouidd ausisul WnjAydoousa [ uinuvsoyly unjAydojoussa | snynsojid sapiojAydojoussa [ Dupipul DuljAydojvusa | psojid vapyjAydojousa | vovdo vapyAydojousd [ pjojja20 vapyAydojwutsa I DIDNIDU vapIjAYydojDULsa | pulpaAy vapiAydojoussa [ DII14JSUOI DapyAYydojoUsa [ pauunag vapijAydojousa [ saploajnf DjjajAydojoudsa [ poIpupjsuaanb pynwuiyopuny sadyjod pynuiyopuny 10Y]DAADI DYNWUIYDPUNY IUOUIS SNUIADIJAJOABAAP ie GO eo ee ee EE EE a eel satsads rurAydoyeu3 | ‘satoads rurAydoieulay Jo suoneroosse ued pue ‘siiqey pue syeriqey ‘Kaid Jo ist] “T FGeL VOLUME 97, NUMBER 2 Figs: arcuate (Fig. 24), and in one case extremely broad, laterally almost contiguous with an- tennifers, with margins also explanate (Fig. 25); gula elongate, narrow (Figs. 22, 23) to moderately broad (Figs. 19, 20, 21, 24), con- cave (Figs. 19, 20, 22-26) to weakly convex (Fig. 21), often with transverse ridges, mar- gins rarely carinate, sometimes with rows of setae on lateral margins, rarely with a Democoris lugens, dorsal habitus of female. - ’ fs > “ Tow rweiv bers uaseverad . LAA ri ~ AS 4009 a a dense matt of whitish, microsetae (Figs. 22, 23). Eyes: moderately sized (Figs. 12, 13, 15, 17, 18) to very large (Figs. 11, 14, 16), most often contiguous, or nearly so, with pronotal collar, rarely removed; ommatidia usually very large, rarely small (Fig. 13), with short to elongate setiform interocular setae (Figs. 11-14). Antennae: inserted be- low midheight of eyes; usually short; mostly 268 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Kundakimuka carvathoi, dorsal habitus of male. with AIII and AIV thinner than AI and AIT]; with setiform vestiture, usually longer than width of segments, rarely intermixed with minute, flattened, adpressed setae on AII; AI usually short, barely surpassing apex of clypeus, rarely moderately surpassing tip of clypeus in females, moderately to greatly thickened; AII usually longest segment, usu- VOLUME 97, NUMBER 2 "XOWIOA = A ‘snadAjo = o ‘ayew ‘anonajpjau ‘DoIpud]Suaanb DYNWIYDOPUNY ‘*¢ “Peasy JO MIIA [BSIOG “9-¢E “SBLY 270 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON == Figs. 7-10. Dorsal view of head. 7, Arygrotelaenus elegans, female. 8, Democoris lugens, female. 9, Ter- matophylella fulvoides, male. 10, Termatophyloides pilosulus, male. ally cylindrical and weakly incrassate api- cally (Figs. 27-29), sometimes fusiform and swollen in males (Fig. 30), and if so, only incrassate in females; AIII cylindrical, ei- ther subequal in length to AII to much shorter; AIV weakly fusiform, subequal in length to shorter than AIII. Labium: in- serted terminally on head, above or at the antennifers in lateral view (Figs. 11-14), slender, sometimes fragile, usually reaching apices of forecoxae, at most posterior mar- gin of mesosternum; LI very short and nar- row, never surpassing posterior margin of bucculae (Figs. 19, 20); LII usually longest segment. Pronotum: tripartite, subtrapezoi- dal (Figs. 31-34, 39-42); lateral margins lin- ear, weakly to strongly divergent posteri- orly; flattened (Figs. 31, 43), evenly tumid (Fig. 40), or biconvex with callosite and disc regions separated by deep incision (Figs. 32, 46); pronotal collar rounded, moderately elongate (Figs. 31-34, 39-42), width vari- able reaching midwidth of eyes to maxi- mum eye width, posterior margin marked by a shallow to deep groove, which is punc- tate (Figs. 31, 33, 34, 35, 40, 42) or non- 7a f)| VOLUME 97, NUMBER 2 -4a[ “py ‘ayeulay ‘vuvipul vuijdAydojoussa ‘ €l ‘sigjtuuajue = B ‘snodAjo = 9 ‘aejndonq = q ‘a[BU ‘avonajDjau UinjAYdojoU ‘gyeulay “ds vapydydojpuisa yl ‘ZT “oyew DIIpuvjsuaanb vynNwu1yDpUNY ‘| ] “Pedy JO MIIA [INV] “py I] ‘s3ty 272 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 15-18. Lateral view of head. 15, Arygrotelaenus elegans, female. 16, Democoris lugens, female. 17, Termatophylella fulvoides, male. 18, Termatophyloides pilosulus, male. punctate (Figs. 32, 39, 41); callosite region separated by a shallow to deep transverse groove, submarginally directed anteriorly, usually meeting pronotal collar (Figs. 31- 34, 39-42), groove usually punctate (Figs. 31, 33-35, 40, 41, 42) rarely with conical, tumid processes (Figs. 32, 36), sometimes callosite region demarcated along midline by a short row of punctures (Figs. 37, 42), with calli clearly separated; antero-lateral pronotal setae, placed dorsad to tergopleur- al suture, and either anteriad or posteriad to propleural suture; disc usually longer than callosite region, sometimes subequal in length, rarely shorter, most often obliquely raised above callosite region, posterior mar- gin truncate, or weakly bisinuate. Scutel- lum: triangular, flattened to moderately rounded, sometimes with lateral margins punctate; anterior margin rarely with small tubercles (Fig. 38). Hemelytra: margins par- allel-sided to strongly arcuate (Fig. 1), usu- ally not greatly extending laterally beyond posterior angles of pronotum, rarely much broader; clavus nearly always with a punc- tate anal vein (Figs. 47, 51-56), sublaterally placed, rarely anal vein as a sinuate groove (Figs. 49, 50); embolium either narrow (Figs. 48, 51, 54, 55, 56), usually less wide than length of pronotal collar, to very broad, sub- equal to or greater than length of pronotal collar (Figs. 47, 52, 53); corium mostly with a punctate R+M vein (Figs. 47, 51-56), variable in length, from midlength of cori- um to just anteriad to cuneal fracture, either shorter (Figs. 51, 56) or longer than medial flexion line (Figs. 47, 48, 52-55), sometimes median flexion line very short (Fig. 49); me- ~ N VOLUME 97, NUMBER 2 -d2[. ‘TZ “apeulay ‘puvipul vurdydojoussa ‘ ‘JUIUIBasS [eIQR] IsIy = | ‘e[Nd = 3 ‘oeynoonq = q ‘ajewW ‘avonajnjaw WinjAydojDUL "‘peoy JO MIA [BI]UDA IZ ‘qyeway “ds vapydydojouiay “OZ ‘seul ‘vIIpuDjsuaanb DyNWU1yYDPUNY ‘6| ‘77-61 “S84 274 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 24 Figs. 23-26. Ventral view of head. 23, Arygrotelaenus elegans, female. 24, Democoris lugens, female. 25, Termatophylella fulvoides, male. 26, Termatophyloides pilosulus, male. dial flexion line mostly subparallel, and weakly divergent posteriorly to R+M vein, or strongly divergent from origin; cuneus mostly as long as wide; membrane either hyaline, translucent or opaque, concolorous or with bicolored pattern; usually glabrous, rarely with dense cover of microsetae; membrane with one cell, very narrow (Figs. 52, 53) to broad (Figs. 48, 54), subelliptical to subquadrate; membrane vein sometimes obscure (Fig. 51), often with an indistinct swelling subcaudally adjacent to cuneus. Thoracic pleura: tergopleural suture usually distinct as a groove anteriad to propleural suture (Figs. 43-46), posteriorly evident as a weak depression, sometimes as a pit above propleural suture; propleural suture most often distinct, sometimes obscure; proepi- sternum short, subtriangular or subquad- rate, explanate; proepimeron large, sub- triangular, strongly depressed mesally, sometimes with one or two pits (Figs. 44, 46); mesepimeron usually polished, rarely rugose, variable, either short and subquad- rate (Figs. 57, 60, 61-64), or large and elon- gate (Figs. 58, 59), with dorsal margin an- VOLUME 97, NUMBER 2 gulate, and dorso-caudal arm bearing an oval to lanceolate spiracular opening (Figs. 65, 66, 67), or recessed and not visible in lateral view (barely visible in latero-caudal view) (Fig. 68), spiracle nearly always bounded by evaporative areas (Figs. 65, 67, 68), in one case absent (Fig. 66), ventral margin of me- sepimeron often obscure, at most as a weak depression, sometimes with vestiture; me- tepisternum bipartite, with a well developed metathoracic gland scent efferent system (Figs. 57-64, 69-72), tumid, occupying about half of the segment, often dorsally contiguous with the mesepimeric evapora- tive areas (Figs. 57, 59, 63, 64), sometimes separated (Figs. 60-62), peritreme usually placed on mid-length of scent efferent sys- tem (Figs. 58, 70, 60, 72, 61) sometimes just caudad of mid-line (Figs. 57, 69, 62, 63, 64), rarely along posterior margin of me- tepisternum (Figs. 59, 71), peritreme tonguelike (Figs. 69, 70) or fingerlike (Fig. 72), mostly polished, with microsetae, mostly straight, rarely weakly arcuate dor- sally (Figs. 62, 64), often with posterior margin raised above plane of evaporative areas, rarely with apex raised; evaporative bodies either elongate (Figs. 73, 74, 76) or subquadrate (Fig. 75), caps mostly widely separated; remainder of metepisternum de- pressed relative to scent efferent system, margins rounded. Legs: usually of moderate length, rarely elongate; metatrochanters rarely with minute spine (Fig. 83); femora mostly fusiform, rarely elongate and cylin- drical, with metafemora rarely greatly en- larged (Fig. 83), and armed with spinelets (Figs. 81-82) or large spine distally on the ventral surface (Figs. 83-84); three or four ventral mesofemoral and four to six ventro- subdistal metafemoral trichobothria (Figs. 77-80), intermixed with moderate to sparse cover of semierect setiform vestiture; tibiae mostly cylindrical, sometimes weakly subquadrate, sometimes with rows of spi- nelets (Figs. 86-88), rarely without rows of spines (Fig. 85), always with moderate to sparse cover of semiadpressed setiform ves- 275 titure, foretibiae always with a well-devel- oped tibial comb; tarsi cylindrical, with TI and TII subequal in length, TIII longest seg- ment, always with semiadpressed setiform vestiture; pretarsus with basally toothed claws, mostly with setiform parempodia (Figs. 90, 96, 91, 97, 92, 98), sometimes parempodia flattened and ribbonlike (Figs. 89, 95, 93, 99, 94, 100), rarely weakly flat- tened with apices expanded, without pul- villi or pseudopulvilli. Abdomen: depressed, very broad, shiny, anthocoridlike, usually with regular distribution of semiadpressed semierect setae, often intermixed with elon- gate trichobothrialike setae laterally on ter- minal segments, abdomen rarely narrow and cylindrical. Male genitalia: genital capsule small, conical (Fig. 101), genital opening dorso-caudal (Fig. 102), small, narrow, ven- tral margin carinate, obliquely raised from left to right; tergite X large, mostly free, rarely partially fused to left dorsal margin of genital opening; left paramere (Figs. 103- 110) placed in a weak excavation of ventro- sinistral corner of genital opening (Fig. 101), sensory lobe very small, with a few sensory hairs on outer margin, shaft arcuate, in rest- ing position at 45° to ventral margin of gen- ital opening (Fig. 102), directed dextrally, apex of shaft either spatulate (Figs. 106, 109) or evenly tapered (Figs. 103-105, 107, 108, 110); right paramere very small, apically ta- pered, placed in ventro-dextral corner of genital opening; aedeagus directed sinis- trally in resting position, crossing shaft of left paramere; phallotheca straplike, apical- ly dissected, sclerotized, broadly connate with vesica and connected basally to phal- lobase by a thin membrane; vesica (Figs. 111-118) with one membranous lobe; duc- tus seminis terminating at base of lobe, without liplike sclerotized secondary gon- opore, usually with gonopore opening into a gonoporal cavity, marked by a U or V-shaped field of spines; usually without sclerotized process (Figs. 111-113, 115- 118), present in one genus (Fig. 114); lobal sclerites usually mesally placed, and linear PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 276 ‘BWI ‘avInajDja WinjAydojDULAa J ‘O€ “aeUlay ‘DUDIpUul DUYAYdowUlia] “67 ‘o[eWay “ds vapydydowusay “BZ ‘a[eW ‘YIIpuDjsuaanb vynu1yHpuny ‘LZ ‘WwauIdas [euUDUR PuoddIg “(—¢-/7 ‘S3IY 277 VOLUME 97, NUMBER 2 ‘pe ‘opeulay ‘vuvipul vuljAydojoussa f, ‘ "BJ9S [PJOUOI [e19}e]-019]Ue = S ‘UOIZOI ISIP = P ‘UOTOI dJISOT[eD = ‘ BO S1e]]OO 2 ‘o[BU ‘avINaDjaut WunjAYdoJDULsa €E ‘qeurgy “ds vapiydydojouisal “TE ‘d[eUL ‘YIIpuDjsuaanb pynu1yopuny ‘[¢E “UINJOUOId JO MIIA [BSIOG ‘PpE-[E ‘SBL4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 278 JOLI9]Ue UO sass9901d . “gjoJoqn} = 1 WIN][IINIs JO UTBIVW JOLIO\UR ‘oyeUay ‘ds NapIjAydojMUida] ‘QE “UOTZII IISOT][ed JO auUTTPru ‘ayeUT ‘apmnajojaut winjAydojpusia I *L¢ “wnjouold Jo sassaooi1d ‘ayeutay ‘ds vapijdydojpussa] “QE ‘ajeul ‘YIIpuHjsuaanb vynulyopuNny “CE (RE BL) WINT[aINds Jo uTsIeW ‘ (LE CE ‘SBI4) UOTBaI d]ISO][eO pu Ie]]OO [e1OUOId Jo UTBIeW JOLIO\sCd UO sassad01d puke suOT}EIOUNd Jo UONROYyTUseU IOYZIY “gE—CE “SBLy VOLUME 97, NUMBER 2 39 Figs. 39-42. Dorsal view of pronotum. 39, Arygrotelaenus elegans, female. 40, Democoris lugens, female. 41, Termatophylella fulvoides, male. 42, Termatophyloides pilosulus, male. to sublinear (Figs. 112, 113, 115) or sickle- shaped (Figs. 111, 116-118), and basally connected to apex of gonoporal cavity; apex of membranous lobe sometimes with fields of spines (Figs. 111, 116-118). CHECKLIST OF THE TERMATOPHYLINI OF THE WORLD Arygrotelaenus Reuter and Poppius, 1912 elegans Reuter and Poppius, 1912 Egypt simoni Reuter and Poppius, 1912 Middle East, Sudan Democoris Cassis n. gen. lugens Cassis n. sp. Australia leptocytus Cassis n. sp. Australia Kundakimuka Cassis n. gen. carvalhoi Cassis n. sp. Australia pallipes (Miyamoto, 1965) nov. comb. Japan queenslandica Cassis n. sp. Australia Termatophylella Carvalho, 1955 fulvoides Carvalho, 1955 Mexico Termatophylidea Reuter and Poppius, 1912 brunnea Maldonado, 1970 Puerto Rico PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 280 ‘ginjns [eins{dos19} = d} ‘uo1swiidso0id = d ‘ainjns [einajdoid = sd . ‘umnusajsidaoid = ad ‘ayew ‘aponajvjaw wnjAydojoutsa I ‘Op ‘q[eulay ‘DuDIpul DUuljAYydojoUsa] ‘Cp ‘ajeulay “ds vapydydojpuial ‘pp ‘seul ‘DIIpunjsuaanb pynuiyopuny “Ep “WINOUOId JO MOIA [BINL] “OP-Cp “SBI4 VOLUME 97, NUMBER 2 281 ETON A Figs. 47-50. Hemelytral characters. 47, Kundakimuka queenslandica, male, hemelytra. 48-50, Termato- phylidea sp., female. 48, hemelytron. 49, anterior aspect of corium and clavus. 50, serrate anal vein of clavus. a = anal vein; m = median flexion line; r = R+M vein. 282 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 51-56. Hemelytra. 51, Arygrotelaenus elegans, female. 52, Democoris lugens, female. 53, Termato- phylella fulvoides, male. 54, Termatophylina indiana, male. 55, Termatophyloides pilosulus, male. 56, Terma- tophylum insigne, male. 283 VOLUME 97, NUMBER 2 ‘gpoesids o1oesioyiejow = s ‘owesied = d ‘wnusajsidajau = sw ‘UOJOWIdasoW = OUI ‘sole JATIRIOAdBAD = Bd ‘a[BUI ‘aVINajDjaWU WiNnjAYydo]MU lA], ‘QQ ‘aeulay ‘vUDIpUI DUYAYdoIDULdAT, “6S ‘apewWay “ds vapljAydojwUsa] “Bg “aye ‘poipunjsuaanb pynwuiyopuny ‘L¢ “eina[d 1deIOYIOIN dG ‘“OQ9-L¢ ‘s3l4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 284 sapiojAydojwusa y, ‘pg “ayewu ‘sapioayn{ vjjajAydoywutsa | . ‘geul ‘snynsojid €9 ‘aTeULEJ ‘sUaSN] SOJOWAG ‘79 ‘a{BULD} ‘sSUDSa]a SnuanjajossAUp “1g “eINa|d s19eIOYIOINg “p9-19 “SBI4 €9 LQ VOLUME 97, NUMBER 2 Mesepimeron and metathoracic spiracle. 65, Kundakimuka queenslandica, male. 66, Termatophylidea sp., female. 67, Termatophylina indiana, female. 68, Termatophylum melaleucae, male. s = metathoracic spiracle. Figs. 65-68. 285 286 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON peritreme. 72. Scent efferent system of metathoracic glands. 69, Kundakimuka queenslandica, male. 70, Termatophylidea sp., female. 71, Termatophylina indiana, , Termatophylum melaleucae, male. p Figs. 69— female. 72 VOLUME 97, NUMBER 2 ‘sdvo dANeIOdRAD = 99 “a]RU ‘aDInaDjaU WNjAYdoJMULdA] “QL ‘afeuUlay ‘DU_IPUl DUI;AYdOJDULAA | ‘cy copeuray “ds vapydydomwuuay ‘pl ‘ajew ‘voIpunjsusanb pynuiyopuny “¢/ “Warscs JUoIaYo 1U9dS puek sWoAed JO UONBoyTUseU JOYsIH “9/-€L “S84 A: ye! PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 288 ‘qjeuray “ds vapydydojmuusay ‘OX ‘6L ‘afew ‘ausisu wunjdydojwusay ‘gL ‘a{ew ‘vI1punjsusanb vynwiyopuny . “BLIYJOQOYOL} JO UOTIISOd 0} JayoI si9quINu LL “BLIYIOgOYIN} SIBIOYIBIIIY “O8-LL “S3l4 289 VOLUME 97, NUMBER 2 ‘soulds JO SMOJ JO UOBOYTUseW JOYsIY ‘DIIpUD/susanb DynUIYDpUNY “py ‘ayeUl ‘vI2Ipuvjsuaanb Dynuiyopuny ‘¢g ‘auids JO uoneoytuseW JoYsIY “few ‘10Y/DA4DI DYNWUIYDpUNY ‘ZR ‘aU ‘1OYIDAIDI DyNWIYDpUNY ‘1g ‘saideds DynWIYDpuNy Jo BIOWIAJe\IW UO soUIdg “pE-[g “S314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 290 DYNULYOPUNY ‘Bg ‘afeul ‘v2Ipunjsuaanb vynwi1yopuny ‘1g ‘apewlay “ds vapyjdydojnuisa J ‘ 98 “9[BUloy ‘ouids [eiqn) = s} ‘oyew ‘70YjDAUDI SUDBAJA SNUADJAIOABAAP “CQ “IeIQH JO sINjeUuIdS “gg—Cg ‘sBLy VOLUME 97, NUMBER 2 constricta Maldonado, 1970 Venezuela hyalina Maldonado, 1970 Venezuela maculata Usinger, 1935 Caribbean, Central America ocellata Carvalho, 1955 Costa Rica opaca Carvalho, 1955 Guyana, Surinam pilosa Reuter and Poppius, 1912 Caribbean, Central America Termatophylina Carvalho, 1988 indiana Carvalho, 1988 India Termatophyloides Carvalho, 1955 pilosulus Carvalho, 1955 Guatemala Termatophylum Reuter, 1884 grande Reuter and Poppius, 1912 India hikosanum Miyamoto, 1965 Japan insigne Reuter, 1884 Middle East, Sudan melaleucae Cassis n. sp. Australia montanum Carvalho, 1983 Brazil nigrum Poppius, 1910 Kenya obscurum Reuter and Poppius, 1912 Sri Lanka, Indonesia ochraceum Reuter and Poppius, 1912 India orientale Poppius, 1915 Taiwan rhea Linnavuori, 1974 Nigeria turneri China South Africa weiri Cassis n. sp. Australia yunnanum Ren, 1983 China 291 KEY TO GENERA . Dorsum without rows of punctures on pro- notum (Fig. 32) and hemelytra (Fig. 48) .... Pesaro eee Termatophylidea (Neotropical region) Dorsum with punctures on pronotum (Fig. 31), and R+M and anal vein (Fig. 47) .......... 2 . Body with rows of flattened scalelike setae on pronotum\and hemelytrany seen ee Le Ae ne Argyrotelaenus (Middle East, Sudan) Body without rows of flattened scalelike setae on pronotum and hemelytra, with setiform hairs only, rarely intermixed with scattered, flat- tenedsiscalelikeisetae’ =... eee eee: 3 . Eyes removed from collar by more than width of second antennal segment ..............- 5 et es ann ae Termatophylina (India) Eyes contiguous with collar or removed by less than width of second antennal segment ..... 4 . Punctate R+M vein shorter than medial flex- ONLINE (ISOS) ea 45 siocvexcestnsn eee ee 6 Punctate R+M vein longer than medial flexion Hine 1G 4) et re nc pad Ce 7 . Posterior margin of pronotal collar punctate (Figs. 31, 35); males with swollen, fusiform second antennal segment (Fig. 30) ......... .... Termatophylum (Eastern Hemisphere, Brazil) Posterior margin of pronotal collar not punc- tate (Figs. 32, 36); males with cylindrical sec- ond antennal segment, weakly incrassate api- cally (Fig. 27) .... Termatophyloides (Guatemala) . Bucculae extremely broad and explanate, lat- erally almost reaching antennifers (Fig. 25) .. a end ieee tay Termatophylella (Mexico) Bucculae moderately broad with margins weakly arcuate;(Fig! 19): 0.2. een eee 8 . Body ovoid, extremely broad (Fig. 1), with em- bolium wider than interocular distance; pro- notum extremely tumid; metafemora not swol- ler re ene et eters Democoris (Australia) Body elongate to elongate-ovoid (Fig. 2), em- bolium not wider than interocular distance; pronotal disc obliquely raised above callosite region in lateral view (Fig. 43); metafemora greatly swolleni(Fig:183) sense ae soe eae. ... Kundakimuka (Australia, Papua New Guinea, Japan, India, Hawaii) KEY TO AUSTRALIAN SPECIES OF TERMATOPHYLINI . Body ovoid, extremely broad (Fig. 1); prono- CUMstumid Mer rer eer en een as 2 Body elongate to elongate-ovoid (Fig. 2); pro- NOtUMIMNOtUMIGVe a eee oe 3 292 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 89-94. Ventral view of pretarsus. 89, Arygrotelaenus simoni, male. 90, Kundakimuka queenslandica, male. 91, Termatophylidea sp., female. 92, Termatophylina indiana, female. 93, Termatophylum insigne, male. 94, Termatophylum melaleucae, male. p = parempodia. NO Clypeus reddish; dorsum with elongate semi- erect: Setaen haem eset ake. Democoris lugens Clypeus dark brown; dorsum with short sem- ladpressed seta «. 45. ..1542- Democoris leptocytus R+M vein longer than medial flexion line (Fig. 47) R+M vein shorter than medial flexion line (Fig. St) hs ee a ee che oe 5 . Metafemora with a large ventro-subdistal spine (E1gsc83184))) gee cote Kundakimuka carvalhoi Metafemora with rows of short, black spines (Figs. 81.82) 52.2. Kundakimuka queenslandica VOLUME 97, NUMBER 2 293 3: Figs. 95-100. Lateral view of pretarsus. 95, Arygrotelaenus simoni, male. 96, Kundakimuka queenslandica, male. 97, Termatophylidea sp., female. 98, Termatophylina indiana, female. 99, Termatophylum insigne, male. 100, Termatophylum melaleucae, male. Hemelytral membrane yellowish, concolorous; GENERIC AND SPECIES DESCRIPTIONS apex of left paramere pointed (Fig. 110) .... DEES ts Rh Sn Fe Termatophylum weiri Arygrotelaenus Reuter and Poppius Hemelytral membrane patterned, mostly dark : brown; apex of left paramere spatulate (Fig. Arygrotelaenus Reuter and Poppius 1912: 109) eRe he awe Termatophylum melaleucae 14; Carvalho 1952: 50 (list), Carvalho 294 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 101-102. Pygophore of Kundakimuka queenslandica, male. 101, lateral view. 102, dorso-caudal view. Ic = left clasper. 1955a: 22 (key), Carvalho 1957: 35 (cat- alogue), Stichel 1956: 176 (description). Figures.—7, 15, 23, 39, 51, 61, 85, 89, 95, 1032 111: Type species.—Arygrotelaenus elegans Reuter and Poppius, 1912, by original des- ignation. Diagnosis. — Arygrotelaenus is recognized by the flattened scalelike setae on the head and pronotum. It is most similar to Ter- matophylum, with the male second anten- nal segment swollen and fusiform in both genera. The vestiture of the male second antennal segment is also similar, with se- tiform, semierect setae, intermixed with flattened, scalelike setae. The antennae of the females differs in the two genera, with the first antennal segment greatly thickened in Arygrotelaenus species. Unlike Terma- tophylum (Fig. 31), the posterior margin of the pronotal collar and callosite region are not punctate in Arygrotelaenus (Fig. 39). Description. — Parallel-sided; coriaceous; rows of punctures on anal and R+M veins (Fig. 51); longitudinal rows of silvery, flat- tened scalelike setae on head, pronotum, he- melytra and thoracic pleura, intermixed with very sparse distribution of setiform vesti- ture. Head: strongly porrect (Fig. 15), elon- gate (Fig. 7); vertex and frons narrowly bi- laterally compressed, with an indistinct row of flattened scalelike setae, lateral margins strongly declivent, almost vertical; poste- rior margin of head carinate; frons barely declivent; clypeus weakly declivent, termi- nating above antennifers (Fig. 15); lateral margins of bucculae weakly arcuate (Fig. 23); gula elongate, narrow, weakly concave, margins not carinate, with a dense matt of whitish microsetae (Fig. 23). Eyes: moder- ately sized, almost contiguous with pronotal collar, separated by a distance less than the width of the second antennal segment; om- matidia large, interocular setae absent. An- tennae: short; AI greatly swollen in females, thicker than AIT; AII weakly incrassate dis- tally in females, subequal in width to AIII; AII fusiform and swollen in males, thicker than AI; AIII and AIV thin, short, subequal in length, combined length subequal to AII length. Labium: barely reaching apices of forecoxae; LII longest segment. Pronotum: (Fig. 39) lateral margins moderately diver- gent; with four longitudinal rows of flat- tened scalelike setae, on lateral margins, and either side of the midline; collar broad, ex- tending to lateral margins of eyes, posterior margin delimited by a shallow, nonpunctate groove; callosite region marked by a trans- verse and sublateral nonpunctate groove, with midline weakly to moderately, trans- versely rugose, calli not distinctly separated, pronotal antero-lateral setae absent; disc subequal in length to callosite region, strongly rounded, obliquely raised above VOLUME 97, NUMBER 2 103 104 107 295 105 10 1 4 J) Figs. 103-110. Left clasper of male, lateral view. 103, Arygrotelaenus simoni. 104, Democoris leptocytus. 105, Kundakimuka queenslandica. 106, Termatophylidea pilosa. 107, Termatophylina indiana. 108, Termato- phylum insigne. 109, Termatophylum melaleucae. 110, Termatophylum weiri. callosite region, posterior margin truncate. Scutellum: weakly declivent, lateral mar- gins not punctate. Hemelytra: (Fig. 51) nar- row, barely extending beyond posterior an- gles of pronotum; lateral margins subpar- allel; anal vein punctate; embolium narrow, width less than length of pronotal collar; weakly punctate R+M vein short, barely surpassing mid-length of corlum, medial flexion line longer than R+M vein, about 7/’s length of corium, subparallel to R+—M vein, and barely divergent posteriorly, re- gion between veins marked by a row of flat- tened setae; cuneus longer than wide, lateral margins almost linear; membrane opaque, with color pattern, mesally brown and lat- SN? Se, ——= > RAL ¥ S Figs. 111-118. muka queenslandica. 114, Termatophylidea pilosa. 115, Termatophylina indiana. 116, Termatophylum insigne. 117, Termatophylum melaleucae. 118, Termatophylum weiri. erally pale; without microsetae; membrane cell subquadrate, broad; membrane vein in- distinct, without subcaudal swelling. Tho- racic pleura: (Fig. 61) tergopleural suture distinct above propleural suture, sometimes marked by a row of flattened scalelike setae; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON = S = ss SS — 7 a 4 2 ——- Se 118 — Vesica, dorsal view. 111, Arygrotelaenus simoni. 112, Democoris leptocytus. 113, Kundaki- proepisternum subtriangular; proepimeron without pits; mesepimeron short, subquad- rate, dorsal margin strongly arcuate, dorso- caudal arm with an oval spiracular opening, with evaporative area, ventral margin marked by a depression and a row of flat- VOLUME 97, NUMBER 2 tened scalelike setae, submesally with a row of flattened scalelike setae; metepisternum scent efferent system occupying half of seg- ment; peritreme mesally placed, tonguelike, parallel-sided, not raised above plane of evaporative areas, evaporative bodies mostly subquadrate, remainder of metepi- sternum with numerous flattened scalelike setae. Legs: moderately long; coxae of mod- erate length; femora fusiform; femoral tri- chobothria not distinct in available speci- mens; tibiae cylindrical, without rows of spinelets, with moderate distribution of stiff, semi-adpressed setae; parempodia flat- tened, ribbonlike (Figs. 89, 95); Abdomen: broad, anthocoridlike, venter semipolished. Male genitalia: genital capsule minute, con- ical; left paramere (Fig. 103) very small, sen- sory lobe minute, with a few sensory hairs, shaft evenly arcuate, tapered apically; ve- sica (Fig. 111) with membranous lobe, gon- oporal cavity present, basally with a pair of weakly sclerotized lobal sclerites, apex of membranous lobe with fields of spines. Distribution.— Egypt, Yemen, Sudan, Is- rael, Saudia Arabia, Iraq. Remarks. —Arygrotelaenus contains two very similar species confined to the Middle East and the Sudan. The genus was de- scribed by Reuter and Poppius (1912) for the Egyptian species, A. e/egans. They er- roneously indicated that the membrane cell is obsolete, and although it is greatly re- duced in both Arygrotelaenus species, the membrane vein is visible. Arygrotelaenus species are the most colorful of the terma- tophylines, and are mostly dark with highly contrasting pale markings on the scutellum and hemelytra. Aside from Stichel’s (1956) redescription of the genus and species, little else has been published on these taxa. Lin- navuori (1975, 1980, 1989) noted their presence throughout the Middle East and indicated that A. simoni is found in asso- ciation with Acacia species. Arygrotelaenus is the sister taxon of Ter- matophylum, and is separated from it by the presence of rows of flattened scalelike 297 setae on the head and pronotum. Some spe- cies of Termatophylum, such as T. insigne, have setae of this type, but they are scat- tered, and never produced into rows. The head structure is also different, with the head of Arygrotelaenus species elongate and strongly porrect, whereas the head in Ter- matophylum species is mostly transverse and less porrect. The vertex and frons in Ary- grotelaenus are narrowly bilaterally com- pressed, and raised dorsally above the eyes, which occurs to a lesser extent in 7erma- tophylum turneri. Arygrotelaenus is also dif- ferentiated from Termatophylum, by the smaller eyes, the lack of interocular setae, and the absence of punctations on the pos- terior margins of the collar and callosite re- gion, and along the midline of the callosite region. Arygrotelaenus elegans Reuter and Poppius Arygrotelaenus elegans Reuter and Poppius 1912: 14, Linnavuori 1951: 104 (descrip- tion), Carvalho 1952: 50 (type species), Carvalho 1957: 35 (catalogue), Wagner 1963: 484 (description), Linnavuori 1975: 6 (Sudan), Linnavuori 1980: 15 (Sudan; biogeography). Figures. —7, 15, 23, 39, 51, 61, 85. Syntypes.—UZMH. Type locality: Des- cehena, Egypt. Not examined. Specimens examined.—Egypt, 1912-24, H12 (NHM,; 2 females). Diagnosis.—This species is very similar to Arygrotelaenus simoni, but can be differ- entiated from it by minor color differences in the hemelytra, and the relative length of the second antennal segment, which is a lit- tle longer than the first antennal segment in A. elegans, and shorter in A. simoni. Description.—Female. Small, length 2.17-—2.23, width 1.13-1.24, mostly dark brown, with yellow markings on hemelytra, and scutellum mostly yellow. Head: length 0.45-0.48, width 0.33-0.34, interocular dis- tance 0.14-0.15, dark brown. Eyes: black tinged with red. Labium: length 0.80, reach- 298 ing apex of forecoxae, yellow, reaching api- ces of forecoxae; LII longest segment, reach- ing posterior margin of head; LIII reaching middle of forecoxae. Antennae: length 0.24, dark brown; AI length 0.25-0.27, basally dark brown, remainder yellow; AIII length 0.14-0.18, yellow; ATV length 0.14—-0.16, yellow. Pronotum: length 0.47, posterior width 0.66-0.70. Scutellum: anterior half brown, remainder yellow. Hemelytra: cla- vus dark brown; corium with anterior half yellow, remainder dark brown; length of R+M vein half of corial length; medial flex- ion line almost reaching cuneal fracture; cu- neus dark brown; membrane opaque, mesal brown band, laterally whitish. Legs: uni- formly yellow. Males not available. Remarks.— Reuter and Poppius’ (1912) original description adequately separated this species from A. simoni, primarily on the basis of color differences. Linnavuori (1951) redescribed the species without fig- uring the male genitalia. Stichel (1956) and Wagner (1963) gave detailed descriptions of the external characters of this species. Lin- navuori (1975) listed the species from Egypt. Linnavuori (1980) listed 4. e/egans from the desert zone of the Saharan region. The coloration, size and shape of the two Arygrotelaenus species are similar and fur- ther study is needed to clearly differentiate the two Arygrotelaenus species. The minor differences in the antennal segment lengths needs to be confirmed with additional ma- terial. No males of A. elegans were available for this study and a comparison of the male genitalia of the two species is necessary to confirm the separation of these taxa. Democoris Cassis n. gen. Figures.—1, 8, 16, 24, 40, 52, 62, 104, Ie. Eytomology.—demos (Greek)—fat; koris (Greek)— bug. Type species.—Democoris lugens Cassis n. sp. Diagnosis.— This genus is recognized by the broad, ovoid body and the extremely PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tumid pronotum. Both species have a fin- ger-like peritreme, with its apex weakly ar- cuate and raised above the plane of the evaporative areas. Description.—Ovoid, extremely broad (Fig. 1); coriaceous, weakly rugo-punctate; with rows of punctures on pronotum (Fig. 40), scutellum, R+M and anal veins (Fig. 53); dorsum with dense distribution of moderate sized to elongate, semiadpressed or semierect, pale setae. Head: transverse (Fig. 8), moderately declivent (Fig. 16), pos- terior margin carinate; vertex weakly to strongly rounded; frons strongly narrowed in front; clypeus moderately declivent, barely reaching antennifers in lateral view; lateral margins of bucculae moderately ar- cuate (Fig. 24); gula moderately broad, weakly concave, with transverse ridges, margins not carinate, with a few elongate, semierect, pale setae. Eyes: very large; con- tiguous with pronotal collar (Fig. 16), cov- ering most of the lateral aspect of the head; ommatidia large, with short or elongate, in- terocular setae. Antennae: short; with mod- erate cover of semiadpressed setae; AI not thickened; AII weakly incrassate distally; AIII cylindrical; AIV not present on avail- able specimens. Labium: reaching apices of forecoxae; LII longest segment. Pronotum: (Fig. 40) lateral margins moderately to weakly divergent posteriorly; moderately to extremely tumid, not biconvex; collar nar- row, at most reaching mid-width of eyes, posterior margin marked by a deep groove with a few separated deep punctures; cal- losite region marked by an arcuate, trans- verse and sublateral, punctate groove, with- out any midline groove, callosite region tu- mid; pronotal antero-lateral setae moder- ately elongate, anteriad to propleural suture; disc subequal in length to callosite region; declivent posteriorly, posterior margin truncate. Mesonotum: small, barely visible from above, strongly declivent. Scutellum: flattened with lateral margins marked by row of punctures. Hemelytra: (Fig. 52) extreme- ly broad, extending beyond posterior angles of pronotum (Fig. 1); lateral margins strong- VOLUME 97, NUMBER 2 ly arcuate; anal vein punctate; embolium very broad, much wider than the length of the pronotal collar; R+M vein punctate, elongate, almost reaching cuneal fracture; medial flexion line shorter than R+M vein, about 7’s length of corium, weakly diver- gent from R+M vein; cuneus length sub- equal to width; membrane translucent, dusty, concolorous, without microsetae; membrane cell narrow to very narrow, vein indistinct, with a posterior, subcaudal, ob- scure thickening. Thoracic pleura: (Fig. 62) tergopleural suture indistinct, weakly de- pressed; propleural suture distinct; proepi- sternum subquadrate; proepimeron without pits; mesepimeron large, broad, subtrape- zoidal, anterior half flattened, posterior half tumid, dorsal margin strongly angulate, dorso-caudal arm with an oval spiracular opening, with evaporative areas, ventral margin marked by a weak, arcuate depres- sion; metepisternum scent efferent system occupying about half of segment, not con- tiguous with mesothoracic evaporative ar- eas; peritreme mesally placed, narrow, fin- gerlike, with apex tapered, weakly arcuate, and raised above the plane of the evapo- rative areas, remainder of metepisternum with a few short, adpressed whitish setae. Legs: moderately long; coxae large, elon- gate; femora elongate, subcylindrical; four mesofemoral and six metafemoral tricho- bothria; tibiae cylindrical, without rows of spinelets, with stiff, semiadpressed setae; pretarsus with setiform parempodia. Ab- domen: broad, anthocoridlike, venter pol- ished. Male genitalia: genital capsule min- ute, conical; left paramere (Fig. 103) mod- erately large, sensory lobe weakly expanded, with a few sensory hairs, shaft evenly ar- cuate, tapered apically; vesica (Fig. 112) with membranous lobe, gonoporal cavity pres- ent, with large, mesal, sublinear lobal scler- ite, tapered apically. Distribution. — Australia. Remarks. — Democoris is morphological- ly atypical in comparison with most ter- matophyline genera. It contains two species confined to the arid and dry mediterranean 299 environs of South Australia. As yet nothing is known of the biology of the two included species. This genus is similar to the Middle American genus Termatophylella, but can be separated from it by differences of the bucculae and the peritreme. The bucculae in Democoris are moderately broad, with the margins arcuate, whereas in Termato- phylella, the bucculae are greatly enlarged, with the margins, greatly enlarged, explan- ate, and laterally almost extending to the antennifers. The peritreme of Democoris is also distinct, being very narrow and finger- like, with the apex strongly tapered, pro- jected away from the body, and above the plane of the evaporative areas. In Terma- tophylella the peritreme is tonguelike, quite broad, with the posterior margin weakly raised along its length, as is typical ofa num- ber of other termatophyline genera. These differences support the present separation of the two genera. Dorsally these genera could be confused, both having an enlarged, somewhat tumid pronotum. All the species are broad, with the embolium particularly broad; however, the body length of Ter- matophylella fulvoides is significantly small- er than the two Democoris species. The rows of punctures on the pronotum, and anal and R+M veins are very similar in both genera. Generally in both genera the membrane cell is narrow, and this is particularly so for 7. fulvoides and D. leptocytus. Democoris is the sister taxon of the clade containing Arygrotelaenus and Termato- phylum (Fig. 131), and is related to these taxa on the basis of the recessed metatho- racic spiracle and the reduced scent efferent system. Democoris also has a close relation- ship with the genus Kundakimuka, although saliently the two genera are dissimilar. De- mocoris contains oval species, with the pro- notum greatly tumid, whereas in Kundaki- muka the body is elongate to elongate to elongate-ovoid. Democoris also differs from Kundakimuka by the following characters; the narrow membrane cell, the metathorac- ic spiracle recessed and not visible in lateral view, the evaporative areas of the mesepi- 300 meron and metepisternum not contiguous, the peritreme positioned on the midline of the scent efferent system, and the parem- podia setiform and not expanded apically. Democoris lugens Cassis n. sp. Figures.—1, 8, 16, 24, 40, 52, 62. Eytomology.—/ugens (latin)—dark. Holotype.—Female. Mitcham. S[outh] Aust[ralia], 17-i-1981, at light, R.V. South- cott (SAMA). Other specimens examined.— Paratype: Female. South Australia: Barossa (SAMA). Diagnosis.—D. /ugens is distinguished from D. leptocytus by the extremely broad body, and the dense distribution of elongate semierect, whitish setae. The clypeus is red- dish-orange, as opposed to the fuscous clyp- eus of D. leptocytus. Description. — Female. Large, length 4.10- 4.55, width 2.11-—2.30, extremely broad, uniformly fuscous, with dense cover of long semierect, whitish setae. Head: length 0.49— 0.58, width 0.72-0.73, interocular distance 0.28, vertex fuscous; frons strongly nar- rowed in front of antennifers, reddish-or- ange; clypeus reddish-orange; maxillary and mandibular plates reddish-orange. Eyes: black tinged with red, with elongate, erect, pale interocular setae. Antennae: AI short, length 0.22, yellow; AII weakly incrassate apically, length 0.53, mostly yellow, with bases and apices weakly embrowned; AIII shorter than AIV, length 0.28, yellow; AIV length 0.36, yellow. Labium: length 1.06, barely surpassing forecoxae; LII reaching midpoint of collar; LIII reaching beyond midpoint of forecoxae. Pronotum: strongly tumid, lateral margins moderately diver- gent; length 0.78-0.96, posterior width 1.67— 1.82; pronotal antero-lateral setae elongate. Mesonotum: barely visible, fuscous. Scutel- lum: fuscous. Hemelytra: embolium ex- tremely broad, almost as wide as interocular distance; membrane short, translucent, with obscure embrownment, with one small, narrow membrane cell (Fig. 52), width sub- equal to embolium width, vein with obscure PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fuscous markings. Legs: coxae brown with apices yellow; trochanters yellow; femora brown, with apices yellow, dorsal margins with short adpressed, pale setae, ventral margins with elongate, semierect, pale setae, with four mesofemoral trichobothria and six metafemoral trichobothria; tibiae yellow- ish, with dense distribution of strong, straight, semiadpressed, pale setae; tarsi yel- lowish. Male unknown. Remarks.—This species is the largest of all the known termatophylines, over 4 mm, and a quarter again the length of its con- gener. This spectacular species is only known from two female specimens from the Ade- laide area and the Barossa Valley of South Australia. Its broad and tumid body is di- agnostic for the genus, and the species is clearly distinguished by color characteris- tics and the vestiture of the dorsum. Democoris leptocytus Cassis n. sp. Figures. — 104, 112. Eytomology.—/epto (Greek)—narrow; kyto (Greek)—cell. Holotype.— Male: South Australia: Mt Davies, 11 Nov[ember] 1963, at light, col- lectors P. Aitken and N. B. Tindale (SAMA). Diagnosis. —D. /eptocytus is readily iden- tified by the very narrow membrane cell, and can be differentiated from its congener by other characters given in the diagnosis of D. lugens. Description.— Male. Small, length 2.95, width 1.60, broad, uniformly fuscous, with dense cover of medium sized, semiad- pressed, whitish setae. Head: length 0.42, width 0.64, interocular distance 0.25, ver- tex fuscous, with posterior margin yellow; frons fuscous; clypeus orange-brown; max- illary and mandibular plates orange-brown. Eyes: black with reddish tinge on inner mar- gins, ommatidia large, with interocular se- tae small, barely surpassing maximum height of ommatidia. Antennae: AI length 0.22, yellow, with a few semiadpressed se- tae; AII length 0.33, weakly incrassate api- VOLUME 97, NUMBER 2 cally, mostly yellow, with an indistinct em- brownment basally, with moderate cover of semiadpressed, pale setae; AIII length 0.21, yellowish-brown, with a moderate cover of semiadpressed, pale setae. Labium: mostly yellow, length 0.54, reaching apices of fore- coxae; LII reaching proxyphus; LIII reach- ing midlength of forecoxae. Pronotum: moderately tumid, callosite region more so than disc, lateral margins weakly divergent, setae thickened along lateral margins; uni- formly fuscous; length 0.61, posterior width 1.96; callosite region more so than disc; pronotal antero-lateral setae moderately sized; posterior margin strongly truncate. Hemelytra: clavus, embolium and cuneus mostly fuscous, apex of clavus with a small yellow marking, and an indistinct red mark- ing above cuneal fracture, corium orange brown; membrane opaque, yellowish- brown, membrane cell very narrow, about as wide as the length of the pronotal collar, vein dark brown and indistinct. Legs: short; coxae dark, shiny brown; femora mostly dark brown with apices yellow; trichoboth- ria not clear in type specimen; tibiae light yellowish-brown. Male genitalia: left para- mere (Fig. 104); vesica (Fig. 112). Female unknown. Remarks.—The male genitalia of this species is very simple, and unlike other ter- matophylines, does not have large basal spicules. This species which is only known from one male specimen is not considered to be conspecific with D. /ugens, known from two female specimens, because of its larger size, more tumid shape of the latter species, and the different vestiture. The two species are restricted to South Australia, with D. leptocytus confined to the arid northwest corner of the state, and D. /ugens found in the Mediterranean zone of South Australia. Kundakimuka Cassis n. gen. Figures.—2, 3, 11, 19, 27, 31, 35, 43, 47, S76 16931697 3507 75, 8182. 83.184,-87, 88; DOR G Ole 0Z 1054113: Eytomology.—kunda (Larrickia)—big, 301 kimuka (Larrickia)—leg. Derived from the Larrickia aboriginal language, which is the predominate language group of the tropical parts of the Northern Territory. Type species.—Kundakimuka carvalhoi Cassis n. sp. Diagnosis.— Kundakimuka is character- ized by the flattened pronotum (Fig. 31), the setiform parempodia which are weakly flat- tened apically (Figs. 90, 96), and the en- larged metafemora, which have either a large subapical spine (Figs. 81, 82), or small bris- tlelike spinelets (Figs. 83, 84) on the ventral surface. This genus is somewhat similar to Termatophylum, however, the punctate R-+M is shorter than the medial flexion line vein in the latter (Fig. 56), and longer in Kundakimuka (Fig. 47). Description. — Elongate to elongate-ovoid (Fig. 2), moderately dorso-ventrally flat- tened, with rows of punctures on pronotum, and anal and R+M veins, moderate cover of long semierect, pale setae. Head: trans- verse (Fig. 3), weakly porrect (Fig. 11); ver- tex rounded; frons moderately declivent; clypeus strongly narrowed in front, strongly declivent, barely reaching antennifers; buc- culae margins moderately arcuate (Fig. 19); gula moderately broad, weakly concave, weakly transversely rugose. Eyes: very large, covering most of lateral aspect of head, con- tiguous with pronotal collar (Fig. 11); om- matidia large, with elongate, interocular se- tae. Antennae: short; AI fusiform, not swol- len; AII weakly incrassate apically (Fig. 27); AIII and AIV slender, narrower than pre- ceding segments. Labium: reaching between the foretrochanters and midpoint of me- sosternum; LII longest segment. Pronotum: (Fig. 31) broad, strongly flattened; lateral margins moderately divergent posteriorly; collar narrow, extending to mid-width of eyes, posterior margin delimited by deep, nonpunctate groove; callosite region marked by a transverse and sublateral sinuate, punc- tate groove (Fig. 36), indistinct near pos- terior margin of pronotal collar, with a short punctate groove along midline from pos- 302 terior margin, pronotal antero-lateral setae elongate, anteriad to propleural suture; disc region flattened, subequal in length to cal- losite region, posterior margin weakly bi- sinuate. Mesosternum: short, visible from above, flattened. Scutellum: flattened, lat- eral margins with a short row of deep punc- tures. Hemelytra: (Fig. 47) moderately broad, just wider than posterior angles of the pronotum (Fig. 2); anal vein punctate; embolium moderately broad, wider than length of pronotal collar; R+M vein punc- tate, elongate, almost reaching cuneal frac- ture; medial flexion line elongate, just short of R+M vein, weakly divergent posteriorly; cuneus as long as wide; membrane trans- lucent, embrowned, concolorous; mem- brane cell subquadrate, broad; membrane vein with a subcaudal thickening. Thoracic pleura: (Fig. 57) tergopelural suture anteri- orly distinctly grooved, posteriorly weakly depressed (Fig. 43); proepisternum subquadrate; propleural suture distinct; proepimeron without pits; mesepimeron narrow, V-shaped, dorsal margin strongly angulate, dorso-caudal arm with a lanceo- late spiracular opening (Fig. 65), with evap- orative area, ventral margin marked by a weak depression, and a few elongate, semi- erect setae; metepisternum scent efferent system large (Fig. 69), occupying about half of the segment, contiguous with metatho- racic evaporative areas; peritreme (Fig. 73) placed just caudad of midline of scent ef- ferent system, weakly dorso-caudally pro- jected, broad, tonguelike, with either apex tumid and raised above plane of evapora- tive areas, or apex tapered and posterior margin weakly raised above evaporative ar- eas, either ending before or sometimes just dorsally beyond evaporative areas; remain- der of metepisternum short, depressed, with a few semierect, pale setae; metepimeron raised, narrow. Legs: moderately long; cox- ae broad, moderately long, weakly separat- ed; metatrochanters sometimes with short spine subdistally (Fig. 83); femora fusiform, with metafemora greatly enlarged each with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON large subdistal ventral spine (Figs. 83, 84), or moderately enlarged with rows of recum- bent, bristlelike spines (Figs. 81, 82), four mesofemoral and six metafemoral tricho- bothria (Fig. 77); tibiae cylindrical to weak- ly arcuate, with four rows of minute spine- lets (Figs. 87, 88), intermixed with semi- erect stiff setae; parempodia weakly flat- tened and expanded apically; pretarsus (Figs. 90, 96) with claws strongly cleft basally, par- empodia setiform, and apically flattened; Abdomen: broad, anthocoridlike, venter, semipolished. Male genitalia: genital cap- sule small, with tergite X partially fused to the dorsal margin of the genital opening; left paramere (Fig. 105) small with a very small sensory lobe, and a narrow straplike shaft, evenly arcuate, and tapered apically; vesica (Fig. 113) with membranous lobe, with weakly spiculate gonoporal cavity, with broad ill-defined lobal sclerite. Distribution.— Australia, Papua New Guinea, Japan, India, Hawaii. Remarks.— Kundakimuka is a distinct genus found in the eastern Indo-Pacific, where new species require description, and one species, closely allied to K. queenslan- dica, has broadly spread. The Japanese spe- cies K. pallipes has been transferred to Kun- dakimuka from Termatophyloides on the basis of the head structure (see Remarks section of Termatophyloides for discussion). The two Australian species described below are the only species critically evaluated in this work. Kundakimuka is externally sim- ilar to Termatophylum, but can be distin- guished from it by numerous characters, in- cluding differences in the length of the R+M vein, the shape of the second antennal seg- ment, the punctation of the pronotum, char- acters of the pterothoracic pleura, metafe- mora, pretarsi, and male vesica. The male second antennal segment in Kundakimuka is weakly incrassate apically, but is always fusiform and swollen in Termatophylum species. The metathoracic spiracle in Kun- dakimuka is exposed, and the associated evaporative areas are contiguous with those VOLUME 97, NUMBER 2 of the scent efferent system of the metepi- sternum. In Termatophylum species the sp1- racle is recessed, with the opening not vis- ible, and the evaporative areas are not con- tiguous with those of the metepisternum. The pretarsi of the two genera show some similarity, with the parempodia flattened in Termatophylum, and only apically flattened in Kundakimuka. The lobal sclerites of the two genera are markedly different; in Kun- dakimuka they are obscure, and in Ter- matophylum they are sickle-shaped. Kundakimuka carvalhoi Cassis n. sp. Figures.—2, 83, 84, 88. Eytomology.—This species is named af- ter José Carvalho, in honor of his contri- butions to world miridology. Holotype.— Male. Smith Point, N[orth- ern] T[erritory], 3-vili-1982, C. Wilson and S. Collins, ex light trap, registration no. 8592 (NTMAG). Diagnosis.—This species can be separated from all other termatophylines by the greatly expanded metafemora, each with a large, ven- tro-subdistal triangular spine (Figs. 83, 84) and the short subdistal spine on each of the me- tatrochanters (Fig. 83). Description.— Male. Small, length 2.40, width 1.01, strongly dorso-ventrally flattened; uniformly yellowish-brown, smooth, with moderate cover of long semierect, golden se- tae. Head: length 0.29, width 0.46, interocular distance 0.19. Eyes: black with reddish tinge. Antennae: uniformly yellowish; AI weakly fu- siform, length 0.13, less than interocular dis- tance, with a few semierect setae; AII, length 0.33, with semiadpressed golden setae; AIII, length 0.20, cylindrical, with adpressed, short setae, and semierect setae; AIV, length 0.13, weakly fusiform, setation as AIII. Labium: length 0.84, reaching midpoint of mesoster- num; LII and LIII subequal in length, LII reaching proxyphus, LII reaching proxyphus, LIII reaching apices of forecoxae. Pronotum: strongly flattened, length 0.43, posterior width 0.82, with uniform distribution of semierect 303 setae; antero-lateral pronotal setae elongate, in- serted dorsad of tergopleural suture and an- teriad of propleural sulcus. Thoracic pleura: peritreme dorsally tumid, raised above and extending just dorsally beyond evaporative ar- eas. Scutellum: yellowish-brown, lighter pos- teriorly, lateral margins without punctures. Hemelytra: uniformly yellowish brown, with R+M and anal veins fuscous, membrane faintly embrowned. Legs: metatrochanters each with a short, subdistal spine (Fig. 83); fore and mesofemora fusiform, not expanded; metafe- mora, greatly expanded, about 2.5 x wider than preceding femora, with a very large, ventral, triangular spine (Figs. 83, 84), apex of meta- femora expanded into a cuplike opening; with four mesofemoral and six metafemoral tricho- bothria; tibiae weakly arcuate, dorsal face with lateral row of closely pressed sclerotized teeth (Fig. 88), ventral face with row of scattered sclerotized teeth, foretibiae with well-devel- oped ventral tibial comb; pretarsal claws with strong basal teeth, and weakly flattened seti- form parempodia, each weakly expanded at apex. Male genitalia: genital capsule minute, conical; left paramere small, weakly arcuate. Female unknown. Distribution.—Northern Territory (known only from type locality). Remarks. — Kundakimuka carvalhoi differs from K. queenslandica in color, with the for- mer having a concolorous, pale yellowish- brown color, and the latter species being dark brown with lighter markings on the dorsum. The peritreme of the two species also differs with the type species having the apex weakly tumid and raised, and extending beyond the dorsal margin of the evaporative areas, where- as in K. queenslandica the peritreme is only weakly raised along its posterior margin, and terminates ventrad of the dorsal margin of the evaporative areas. The peritremes are similar in placement; just posteriad of the midwidth of the scent efferent system, and dorso-cau- dally directed. The metafemora in the two Australian Kundakimuka species are enlarged, but particularly so in K. carvalhoi, with each having an enlarged, subdistal spine. 304 Kundakimuka queenslandica Cassis n. sp. Figures.—3, 11, 19, 27, 31, 35, 43, 47, 57, 65, 69, 81, 82, 87, 90, 96, 101, 102, 105, 113. Eytomology.—The species name refers to the State of Queensland distribution. Holotype.—Male. Maryborough, Queens- land, 18-I[V-1973, D.A.I. N3700 13277, C.LE. collfection] A.6716 13277 (QDPI). Other specimens examined.—Paratypes: QLD: Maryborough, 18-IV-1973, D.A.L., N3700, suspected predator of Neodrepta lu- teodactella (Lepidoptera) larva on Melaleuca integrifolia (Myrtaceae) (NHM: 3 males, 2 fe- males; QDPI: 1 male, 1 female); QLD: Gym- pie, Lagoon Pkt, 4-VII-1968, D.A.I., numer- ous preying on larvae of Neodrepta luteodac- tella on Melaleuca integrifolia foliage, N3106 (NHM: 2 males, 2 females); QLD: Bauple, (CSR), 1-VII-1974, with Neodrepta luteodac- tella on Melaleuca integrifolia, N3852 (QDPI: 2 females, | male). Diagnosis. — Distinguished from other Kun- dakimuka species by the presence of small spines on the ventral surface of the meso- and metafemora. Description.—Male. Small, length 2.33- 2.82, width 0.98—1.36, moderately dorso-ven- trally flattened, mostly dark brown, with a few yellowish-orange markings on the dorsum, and a moderate cover of long, semi-erect, golden setae. Head: length 0.34—0.46, width 0.49-0.53, interocular distance 0.23-0.24, mostly brown with clypeus, maxillary and mandibular plates yellowish brown. Eyes: mostly reddish with fuscous tinge. Antennae: AI length 0.14—0.16, proximal half yellowish-brown, distal half yel- low, AII length 0.34—0.37, weakly incrassate apically, yellow; AIII length 0.22-0.24, yellow; AIV length 0.22-0.24, brown. Labium: length 0.88-1.20, reaching between apices of fore- coxae and foretrochanters. Pronotum: mod- erately flattened, length 0.48-0.54, posterior width 0.82-0.96, mostly brown, with a yel- lowish marking on midline of pronotal disc. Thoracic pleura: peritreme short of dorsal mar- gin of evaporative areas, polished orange dark brown, slightly raised above evaporative areas. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mesonotum: brown with distinct reddish markings on each side of midline. Scutellum: brown laterally, with an indistinct reddish-yel- low to yellowish marking along posterior half of midline. Hemelytra: mostly brown, lighter brown to yellowish-brown on anterior parts of corium, embolium, and cuneal fracture, mem- brane mostly dusty brown, sometimes mesally pale and translucent. Legs: forefemora mostly yellow, sometimes with a dorso-proximal in- fuscation; mesofemora and metafemora fus- cous with apices yellow, ventral margins with indistinct rows of very short, recurved, black spines (Figs. 81, 82), metafemora moderately incrassate, four mesofemoral and six metafe- moral trichobothria; tibiae cylindrical with four rows of minute, closely pressed sclerotized, fuscous teeth (Fig. 87); pretarsal claws (Figs. 90, 96) strongly cleft basally, with setiform par- empodia, apically flattened (Fig. 90). Genita- lia: tergite X as a U-shaped process, connate apically with dorsal margin of genital opening; left paramere (Fig. 105); vesica (Fig. 113). Female. Similar to male in structure and color. Length 2.79-3.04, width 1.30-1.39. Head: \ength 0.36-0.37, width 0.54—0.60, in- terocular distance 0.54-0.60. Antennae: AI length 0.14-0.16, AII length 0.36-0.38, ATI length 0.23-0.24, AIV length 0.22-0.23. La- bium: length 0.98-1.02. Pronotum: length 0.58- 0.60, posterior width 1.06. Remarks.—K. queenslandica is somewhat similar to Termatophylum species in color and texture, and in coastal Queensland where these species are sympatric, differentiating the two genera requires careful examination. In such cases the relative lengths of the R+ M vein and medial flexion line veins easily distinguishes the two genera, with Termatophylum species having a short R+M vein. Specimens of Kundakimuka from India, Pa- pua New Guinea and Hawaii have been re- ceived from the Natural History Museum which are very similar to K. queenslandica. They possess spiculate ventral surfaces of the meso- and metafemora. Adam Asquith (per- sonal communication) has indicated that the termatophyline found in Hawaii is a recent VOLUME 97, NUMBER 2 introduction. It is possible that the specimens from India and New Guinea could also be introductions, and that this species may be adventitious in other areas of the Indo-Pacific. At present these specimens are not considered to be conspecific with K. queenslandica. All specimens of Kundakimuka queenslan- dica have been recorded from Melaleuca in- tegrifolia (Myrtaceae), which also harbors Ter- matophylum weiri. The label data on these specimens indicates that K. queenslandica feeds on the lepidopterous larvae of Neodrepta lu- teotactella. Termatophylella Carvalho Termatophylella Carvalho 1955b: 643; Car- valho 1955a: 22 (key), Carvalho 1957: 35 (catalogue) Figures.—9, 17, 25, 41, 53, 63. Type species.— Termatophylella fulvoides Carvalho 1955, by original designation. Diagnosis.— Termatophylella is recognized by the greatly enlarged bucculae (Fig. 9), with the lateral margins explanate, and almost reaching the antennifers when viewed ven- trally. Description.—Ovoid, with rows of punc- tures on transverse groove of pronotum, anal and R+M veins; sparse cover of short, sem- iadpressed setae. Head: transverse (Fig. 9), strongly porrect (Fig. 17); vertex strongly con- vex; frons declivent, rounded, wide; clypeus short, weakly declivent, terminating above an- tennifers; bucculae greatly enlarged, and wide- ly separated, explanate, and almost contiguous with antennifers (Fig. 25), longer than first la- bial segment; gula elongate, narrow, weakly convex, lateral margins carinate, with trans- verse ridges, with a row of soft, moderately long, semierect setae. Eyes: large, almost con- tiguous with pronotal collar, distance less than the width of the second antennal segment; om- matidia large, with short, interocular setae. An- tennae: short; AI fusiform, not swollen; AII weakly incrassate distally; AIII subequal in length to AII, longer than AIV. Labium: reach- ing apices of forecoxae; LIV longest segment. 305 Pronotum: (Fig. 41) biconvex; lateral margins weakly carinate; collar narrow, reaching mid- width of eyes, posterior margin delimited by a deep nonpunctate groove; callosite region tumid, marked by a postero-transverse and sublateral punctate groove, with a faint, non- punctate impression along midline, pronotal antero-lateral setae not present in available specimens; disc shorter in length than callosite region, rounded, posterior margin truncate. Mesonotum: large, declivent, visible from above. Scutellum: weakly rounded, lateral margin not punctate. Hemelytra: (Fig. 53) broad, extending well beyond lateral margins of pronotum; embolium broad, width greater than length of pronotal collar; anal vein punc- tate, elongate, almost reaching cuneal fracture; medial flexion line long, almost *4’s of length of corium, shorter than R+M vein, strongly divergent to R+M vein; cuneus very large, longer than wide; membrane translucent, dusty, concolorous; without microsetae; membrane cell very narrow; membrane vein indistinct, without subcaudal thickening. Thoracic pleu- ra: tergopleural suture distinct anteriorly; proe- pisternum subquadrate; proepimeron without pits; mesepimeron (Fig. 63) moderately sized, rugose, dorsal margin moderately angulate, dorso-caudal arm with a short lanceolate spi- racular opening, with evaporative area, ventral margin marked by a weak depression; mete- pisternum scent efferent system (Fig. 63), oc- cupying about half of the segment, not contig- uous with metathoracic evaporative areas; per- itreme submarginal to posterior margin of seg- ment, dorsally dorso-caudally projected, tongue-shaped, weakly tumid at dorsal end, weakly raised above plane of evaporative ar- eas; evaporative bodies mostly with rounded appearance; remainder of metepisternum weakly depressed. Legs: moderately long; cox- ae elongate; femora fusiform; three mesofe- moral and four metafemoral trichobothria; tibiae cylindrical, with stiff, semiadpressed se- tae, without rows of spinelets; parempodia se- tiform. Abdomen: broad, anthocoridlike, ven- ter semipolished. Male genitalia: left paramere at right angles to ventral margin of genital 306 opening, straplike with a very small sensory lobe. Male aedeagus not examined. Remarks. — 7 ermatophylella is a monotypic genus restricted to Mexico. This genus is known from only two specimens. I did not examine the interna! male genitalia of T. fulvoides pend- ing the collection of more material of this spe- cies. Carvalho (1955a) distinguished 7erma- tophylella from other termatophylines by the transverse head, however, this character also occurs in Democoris and Kundakimuka. The genus shows numerous similarities with the Australian endemic genus Democoris, but can be distinguished by characters of the bucculae and the peritreme of the scent efferent system (see Democoris Remarks section). Termato- phylella is the sister genus of the clade con- taining Kundakimuka + Democoris + Ary- grotelaenus + Termatophylum, and is united with this group by the presence of the enlarged eyes, which are contiguous with the posterior margin of the pronotum. Termatophylella fulvoides Carvalho Termatophylella fulvoides Carvalho 1955b: 644; Carvalho 1957: 35 (catalogue). Figures.—9, 17, 25, 41, 53, 63. Holotype.— Male. Mexico Br. Tex. 69391, 7/IL/1950, 329, on orchid plants (USNM;; type seen). Other specimens examined. — Mexico, State of Vera Cruz, 3-20-1962, Jackson collfection], ex bromelaids (USNM; male). Diagnosis.—This species is clearly differ- entiated from other Western Hemisphere ter- matophylines by the very narrow membrane cell (Fig. 53). The enlarged bucculae distin- guishes it from all termatophylines. Description.—Male. Small, length 2.26- 2.51, width 1.24—1.30, mostly dark brown, with sparse cover of short semiadpressed, pale se- tae. Head: porrect, rounded, longer than wide, length 0.38, width 0.49-0.52, interocular dis- tance 0.24—0.28, mostly dark reddish-brown, posterior margin and mandibular plate or- ange-brown, apex of clypeus red. Eyes: red, with short interocular setae, barely surpassing PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON maximum height of ommatidia. Antennae: short, AI length 0.20 brown, fusiform; AII length 0.32, basally narrower than AI, weakly incrassate apically, yellow; AIII length 0.24, mostly yellow, weakly embrowned apically; AIV length 0.34, yellow, weakly fusiform. La- bium: length 0.96, reaching mesocoxae; LI barely reaching posterior margin of bucculae; LIT reaching proxyphus; LIII short of apices of forecoxae; LIV longest segment. Pronotum: uniformly fuscous; length 0.64, posterior width 0.94. Mesonotum: mostly fuscous, with red- dish tinge. Scutellum: fuscous. Hemelytra: em- bolium reddish-brown; clavus and corium dark brown; membrane translucent, yellowish, cell narrow, subequal in width to length of pronotal collar, vein brown. Legs: femora dark brown, four meso- and six metafemoral trichobothria; tibiae mostly yellowish-brown, basally dark brown; tarsi yellow. Abdomen: dark brown. Male genitalia: genital capsule small, conical, mostly dark brown, tergite X orange brown; left paramere small, orange-brown, sensory lobe minute, shaft weakly arcuate, spatulate. Female unknown. Remarks.—This species is known from or- chids and bromelaids (Table 1), however, this information probably represents sitting re- cords, and does not indicate true host plant records. More material is needed to fully de- scribe this species, and field studies are needed to establish its hosts and habits. Termatophylidea Reuter and Poppius Termatophylidea Reuter and Poppius 1912: 2, 5, Usinger 1935: 271 (note), Carvalho 1952: 50 (list), Carvalho 1955a: 22 (key), Carvalho 1957: 35 (catalogue), Maldonado 1970: 119 (taxonomic history; species key), Callan 1975: 389-391 (biology). Figures.—4, 12, 20, 28, 32, 36, 38, 44, 48, 49, 50, 58, 66, 70, 74, 79, 80, 86, 91, 97, 106, 114. Type species.— Termatophylidea pilosa Reuter and Poppius 1912 by monotypy. Diagnosis. — Termatophylidea is character- ized by glassy, broad hemelytra, with the em- VOLUME 97, NUMBER 2 bolium enlarged. Unlike any other termato- phyline, they do not have rows of punctations on the pronotum (Figs. 32, 36) and hemelytra (Fig. 48), and the anal vein of the clavus is present as a serrate groove (Figs. 49, 50). They also do not possess evaporative areas sur- rounding the metathoracic spiracle (Fig. 66), and they have a small pit along the anterior margin of the metepisternum (Fig. 58). Description. —Elongate-ovoid to ovoid, mostly smooth with pronotum granulate; without rows of punctures on dorsum (Figs. 32, 48), with moderate cover of elongate, semi- erect, pale setae, arising from pronounced bas- es, intermixed with microsetae on hemelytra. Head: elongate (Fig. 4) strongly porrect (Fig. 12); vertex weakly rounded, postoccular mar- gins either subparallel or moderately conver- gent posteriorly, posterior margin very weakly carinate; frons moderately narrowed; clypeus subhorizontal, terminating above antennifers; bucculae short, margins subparallel (Fig. 20); gula moderately broad, weakly concave, dis- tinctly transversely rugose, without any dis- tinctive vestiture. Eyes: small to moderately sized, separated from anterior margin of pron- otal collar by a distance equal to or greater than width of second antennal segment; om- matidia large, with elongate, interocular setae. Antennae: moderately sized, subcylindrical, with AI, AIII and ATV subequal in length, and All 3x to 4x longer than other segments; AI not thickened; AII thickest segment, incrassate apically (Fig. 28); ATI and AIV fusiform. La- bium: reaching between apices of forecoxae and midlength of mesosternum. Pronotum: (Figs. 32, 36, 44) biconvex; lateral margins strongly divergent posteriorly; pronotal collar narrow (Fig. 32), extending to mid-width of eyes, marked posteriorly by a deep nonpunc- tate groove; callosite region marked by a deep, nonpunctate, transverse and sublateral groove (Fig. 36), incomplete mesally, with or without tubercles either side of midline contiguous with the posterior margin of the callosite region, mostly smooth, with a few setae restricted to the midline and below the posterior margin of pronotal collar; antero-lateral pronotal setae 307 caudad of propleural suture; disc longer than callosite region, mostly with a granular ap- pearance, with elongate, semierect setae with distinct raised bases, posterior margin weakly bisinuate. Mesonotumi: visible from above, de- clivent, sometimes with a dense matt of whit- ish microsetae. Scutellum: lateral margins not punctate, sometimes with tubercles just cau- dad of anterior margin (Fig. 38). Hemelytra: (Figs. 48-50) hyaline; broad, widest caudad of midlength of corium; lateral margins strongly arcuate; anal vein sinuate (Figs. 48, 49), non- punctate, region laterad of vein with dense matt of microsetae; embolium narrow, thickened; R+M vein not punctate, almost reaching cu- neal fracture; corlum broad, mostly glassy, sometimes with darker markings, weakly to moderately setate, setae with tuberculate, sometimes embrowned bases; medial flexion line either obscure and short or obsolete; cu- neus either as long as wide, or usually longer than wide; membrane transparent to translu- cent, usually concolorous; with dense matt of microsetae; membrane cell large, subquadrate; membrane vein with subcaudal thickening. Thoracic pleura: tergopleural suture indistinct (Fig. 44), sometimes marked by a subanterior pit, remainder as a very weak depression; pro- pleural suture indistinct, usually marked by a pit; proepisternum subquadrate; proepimeron usually with a subanterior pit; mesepimeron elongate (Fig. 58), usually smooth and shiny, dorsal margin weakly angulate, with narrow, lanceolate spiracular opening on dorso-caudal arm of dorsal margin (Fig. 66), without evap- orative areas, lined with microsetae, ventral margin as a weak depression; metepisternum with reduced scent efferent system (Fig. 58), occupying about ' of the segment, not reach- ing metathoracic spiracle, dorsal margin usu- ally emarginate, peritreme (Fig. 70) tonguelike, broad, along midwidth of scent efferent sys- tem, dorsally directed, short of dorsal margin of scent efferent system, not raised dorsally or along posterior margin, above evaporative ar- eas; evaporative bodies widely separated, caps mostly elongate (Fig. 74), much longer than wide, remainder of segment weakly depressed, 308 anterior margin carinate, with a pit dorsad of dorsal margin of scent efferent system. Legs: elongate, cylindrical; coxae elongate; femora elongate and narrow, with four mesofemoral and six metafemoral trichobothria (Figs. 79, 80), with moderately dense distribution of elongate, semierect setae; tibiae elongate and narrow, cylindrical, with rows of spinelets (Fig. 86), intermixed with elongate, semiadpressed setae; pretarsus (Figs. 91, 97) with claws greatly thickened basally, apically strongly arcuate, parempodia setiform, not flattened apically. Female abdomen: broad. Male abdomen: sub- cylindrical. Male genitalia: genital capsule small, conical; left paramere minute (Fig. 106), with small sensory lobe, and shaft weakly ar- cuate, broad, and weakly enlarged apically; ve- sica (Fig. 114) with narrow membranous lobe, with a sclerotized process, directly connected to the termination of the ductus seminis, and almost reaching apex of vesica, no gonoporal cavity, lobal sclerites or fields of spines. Remarks. — Termatophylidea is restricted to the Caribbean, Central America, and northern South America. Termatophylidea is atypical of most termatophylines in not having punc- tate rows on the pronotum and hemelytra. It is superficially similar to Deraeocorinae: Hy- aliodini because of the glassy, broad hemelytra, however the head structure is markedly dif- ferent. The head of Termatophylidea is strong- ly porrect like all other termatophylines. This genus is confidently retained in the Termato- phylini because of the short first labial segment and the presence of antero-lateral pronotal se- tae. Aside from the nonpunctate dorsum it differs from other termatophylines most mark- edly in the form of the metathoracic spiracle and the male aedeagus. The spiracle lacks any surrounding evaporative areas, although the opening is exposed as in most termatophy- lines. The male genitalia are atypical for ter- matophylines in possessing a sclerotized pro- cess, which is connected to the apex of the ductus seminis. This is reminiscent of the male genitalia of some Hyaliodini (Kelton 1959), and suggests a possible relationship between the hyaliodines and the termatophylines. The present description of Termatophylidea is based PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON on observations of 7. maculata, T. opaca, and T. pilosa, and an undescribed species from Mexico, and the descriptions of other species by Carvalho (1955a) and Maldonado (1970). Callan (1975) has reviewed the biology of Ter- matophylidea species, and reports that T. pi- losa, T. maculata and T. opaca are noted pred- ators of thrip pest species. Termatophylidea pilosa Reuter and Poppius Termatophylidea pilosa Reuter and Poppius 1912: 5 (. sp.), Usinger 1935: 271 (mote); Myers 1935: 22 (biology), Carvalho 1952: 50 (list), Carvalho 1955b: 647 (description), Carvalho 1957: 36 (description), Maldon- ado 1970: 125 (key), Callan 1975: 389-391 (biology). Figures. — 106, 114. Syntypes.—Male and female specimens. Mandeville, Jamaica (as Jamaika), IV/1906, [E.P.] Van Duzee (UZMH,; not examined). Specimens examined.—Jamaica, J. G. My- ers, 1933 #3515, Brit. Mus. 1933-400 (NHM; male). Diagnosis. — Termatophylidea pilosa is rec- ognized by the embrowned markings on the clavus and corium. The male vesica is dis- tinctive (Fig. 114), possessing a sclerotized pro- cess, however the male genitalia of no other Termatophylidea species has been examined. Description.— Male. Small, length 3.10, width 1.20, uniformly yellow with hemelytra mostly transparent, with sparse cover of semi- erect, pale elongate setae. Head: extremely por- rect, rounded, length 0.52, width 0.46, inter- ocular distance 0.20; postoccular margins tu- mid; clypeus with lateral margins tinged with red; maxillary and mandibular plates yellow, tinged with red. Eyes: black, tinged with red, moderately sized. Antennae: AI length 0.24, with a few elongate, semiadpressed setae, yel- low, with slight reddish tinge, AII length 0.70, uniformly and weakly swollen, with dense dis- tribution of elongate, semiadpressed pale setae, uniformly yellow; ATI and AIV missing in available specimen. Labium: length 1.14, reaching just beyond the apices of the forecoxae; LII elongate, about twice length of VOLUME 97, NUMBER 2 LIII. Pronotum: length 0.54, posterior width 0.82; mostly yellowish, with grooves and lat- eral margins embrowned and polished, with moderate distribution of elongate, semierect setae. Scutellum: moderately rounded, yellow, with a few elongate, semierect setae, with two minute processes on either side of the midline, just caudad of anterior margin. Hemelytra: mostly transparent with clavus and meso-cau- dal aspect of corlum weakly embrowned; em- bolium thickened, with moderate distribution of semierect, elongate setae; corium with a few scattered semierect, elongate setae; membrane vein weakly tinged with red. Legs: uniformly yellow; femora very long, narrow, with mod- erate distribution of semiadpressed setae; tib- iae elongate, cylindrical, with four rows of min- ute spines, with dense distribution of semiad- pressed setae. Abdomen: not broad, subcy- lindrical, venter yellowish and brown laterally, with sparse distribution of elongate, semirect setae. Male genitalia: left paramere (Fig. 106); vesica (Fig. 114). Female not available. Remarks. — Termatophylidea pilosa has re- ceived some attention in the literature because it is a predator of the economic pest, the cacao thrips, Selenothrips rubrocinctus. Myers (1935) and Callan (1975) gave details of the biology of this species (see Biology section). Carvalho (1955b) redescribed this termatophyline spe- cies, and also recorded it from Central and South America. Usinger (1935) made a com- parison of this species with 7. maculata, and included a habitus illustration of the male of the species. Maldonado (1970) in a review of Termatophylidea, gave a key to species, and distinguished 7. pilosa mainly on color char- acteristics. Termatophylina Carvalho Termatophylina Carvalho 1988: 235. Figures.—5, 13, 21, 29, 33, 45, 54, 59, 67, TV TS9I2)/ 98107 1S. Type species.— Termatophylina indiana Carvalho, 1988: 235, by original designation. Diagnosis.— Termatophylina is diagnosed by the narrow body and elongate appendages. 309 It has numerous other apomorphies including the peritreme positioned on the posterior mar- gin of metepisternum (Fig. 59) and the eyes distantly removed from the pronotum (Figs. 5, 13). The pronotum is similar to Kundaki- muka, but the head is much longer in Ter- matophylina. Description. — Elongate, parallel-sided, dark brown, coriaceous, with rows of punctures on pronotum (Fig. 33), and anal and R+M veins (Fig. 54), with moderate cover of elongate, pale, semierect setae. Head: elongate (Fig. 5), porrect (Fig. 13); vertex rounded, posterior margin not carinate; frons short; clypeus large, weakly de- clivent, terminating above antennifers; buc- culae moderately large, wide, lateral margins arcuate (Fig. 21), posteriorly extending just be- yond antennifers; gula elongate, moderately wide, weakly convex, weakly transversely ru- gose. Eyes: removed from the pronotum by a distance subequal to length of pronotal collar, moderately large; ommatidia small, with a few elongate, interocular setae. Antennae: relative- ly long, longer than head and pronotum com- bined; all segments cylindrical and narrow, AII cylindrical (Fig. 29). Labium: elongate and slender, reaching posterior margin of mesos- ternum; LII elongate reaching posterior mar- gin of head; LIII longest segment, reaching midpoint of forecoxae. Pronotum: (Fig. 33) weakly flattened, subtrapezoidal; collar round- ed, moderate length, broad, almost extending to lateral margins of eyes, posterior margin marked by a moderately deep, impunctate groove; callosite region marked by a sinuate, transverse, weakly punctate groove; antero-lat- eral pronotal setae elongate, dorsad of tergo- pleural suture and anteriad to propleural sul- cus; disc region weakly rounded, subequal in length to callosite region, posterior margin weakly excavate on either side of midline. Mesonotum: relatively long and visible from above, weakly declivent. Scutellum: weakly flattened, not punctate. Hemelytra: (Fig. 54) narrow, barely surpassing posterior angles of pronotum; clavus broad with a punctate anal vein; embolium narrow, a little wider than width of first antennal segment; corium with an elongate, punctate R+M vein, about %4 310 length of corium, subequal in length to a weak- ly divergent medial flexion line; cuneus slightly longer than wide; membrane with a moder- ately wide membrane cell. Thoracic pleura: ter- gopleural suture mostly distinct (Fig. 45), mesally obscure placed at midheight of eyes; propleural suture distinct; proepimeron subquadrate, explanate; proepisternum subquadrate, depressed; mesepimeron elon- gate (Fig. 59), broad, dorsal margin angulate, dorso-caudal arm bearing a lanceolate spirac- ular opening (Fig. 67), with evaporative bod- ies, ventral margin of mesepimeron marked by a depression; metepisternum with a well- developed scent efferent system (Fig. 71), dor- sal margin contiguous with evaporative areas of mesepimeron; peritreme almost contiguous with posterior margin of metepisternum, tongue-shaped, posterior margin raised above evaporative areas; evaporative bodies subquadrate to elongate (Fig. 75); remainder of metepisternum weakly depressed, dorsally with elongate setae. Legs: all segments elon- gate, cylindrical; femora narrow, with four me- sofemoral and six metafemoral trichobothria; pretarsi (Figs. 92, 98) with narrow, strongly C-shaped claws, weakly cleft basally, parem- podia short and setiform. Abdomen: moder- ately broad, venter semipolished. Male geni- talia: genital capsule small, conical; left para- mere (Fig. 107) moderately large, sensory lobe weakly tumid, with a few sensory hairs, shaft, narrow, evenly arcuate, tapered apically; ve- sica (Fig. 115) with membranous lobe, gono- poral cavity present, with large, mesal, narrow, linear lobal sclerite. Distribution. —India and Malaysia. Remarks. —Carvalho (1988) described this genus from India, suggesting that it was closely allied to Termatophylum. From my investi- gations it is evident that Termatophylina dif- fers considerably from Termatophylum, par- ticularly in the head structure, with the former having an elongate head and the eyes greatly removed from the pronotal collar. Unlike any other termatophyline, Termatophylina has very small ommatidia. The scent efferent system 1s also distinctive, with the peritreme running PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON along the posterior margin of the metepister- num. The pronotal grooves are also reduced, with the degree of punctation minimal, and unlike in Termatophylum, the posterior mar- gin of the pronotal collar is not punctate. Ter- matophylina also contains an undescribed spe- cies from West Malaysia. This species is rep- resented by two specimens in the Natural His- tory Museum (London). This species is larger than 7. indiana, but its overall facies are very similar to the type species. Termatophylina indiana Carvalho Termatophylina indiana Carvalho 1988: 236 (n. sp.). Figures.—5, 13, 21, 29, 33, 45, 54, 59, 67, TARTS 292) 98: LOWailsS: Holotype.— Male. India: Calicut Universi- ty, Madras, Suchitra col[lection], Nov[ember] 1987—Jan[uary] 1988 (NHM; not seen). Specimens examined.—India: Kerala, Cal- icut Univer[sity] Campus, 22-II-1984, S. A. George, C.I.E.A. 15860 (NHM; male, 2 fe- males). Diagnosis.— 7ermatophylina indiana is dis- tinguished from an undescribed Termatophy- lina species by its smaller size, and the shape of the lobal sclerite of the male genitalia (Fig. 115); Description.— Male. Moderate size, length 2.73, width 0.74, mostly dark brown, with head lighter, and reddish and yellowish tinges on hemelytra, with moderate distribution of elon- gate, semiadpressed, golden setae. Head: elon- gate, length 0.53, width 0.42, interocular dis- tance 0.18, vertex mesally medium brown, lat- erally yellow, maxillary and mandibular plates yellow; clypeus yellowish to yellowish-brown. Eyes: reddish-black. Antennae: with a mod- erate cover of semierect setae; AI length 0.22, yellow, sometimes proximally embrowned; AI length 0.55, yellow; ATI length 0.38, proxi- mally more yellowish, distally reddish-brown; AIV length 0.24, reddish-brown. Labium: length 1.24. Pronotum: dark brown, with elon- gate, erect to semierect, golden setae, antero- lateral pronotal setae dorsad of tergopleural VOLUME 97, NUMBER 2 suture, and anteriad of propleural suture. He- melytra: mostly dark brown, embolium light brown, R+M vein reddish, cuneal fracture and antero-lateral margins of cuneus yellowish with reddish tinge; membrane dark brown, with dense cover of microsetae. Legs: mostly yel- low, with meso and metafemora with subapi- cal reddish tinge. Male genitalia: left paramere (Fig. 107); vesica (Fig. 115). Female. Similar to male in structure, color and vestiture. Length 3.07-3.10, width 0.84— 0.90. Head: length 0.52-0.69, width 0.43-0.46, interocular distance 0.17-0.21. Antennae: AI length 0.24; AII 0.50-0.54; AIII length 0.36- 0.37, AIV 0.24-0.31. Labium: length 1.29- 1.44. Distribution. — India. Host association. — Associated with the lar- val galleries of Lamida moncusalis Walker (Lepidoptera: Pyralidae). Remarks.—Carvalho (1988) described T. indiana, and included a habitus illustration, and detailed figures of the male genitalia. His illustration of the male vesica indicates the presence of a sclerotized process. In my dis- sections the sclerite was not connected to the apex of the ductus seminis, and appeared to be associated with a gonoporal cavity as in most other termatophylines, and is therefore considered to be a lobal sclerite. Termatophyloides Carvalho Termatophyloides Carvalho 1955b: 641 (n. gen.), Carvalho 1955a: 36 (key), Carvalho 1957: 36 (catalogue). Figures.— 10, 18, 26, 42, 55, 64. Type species. — Termatophyloides pilosulus Carvalho, 1955, by monotypy. Diagnosis. — Termatophyloides is separated from other termatophylines by a combination of characters, and does not contain a single autapomorphy. Like Termatophylum, the cal- losite region of this genus has a longitudinal row of punctures along the midline. It can be separated from Termatophylum by the elon- gate, punctate R+M vein, which is longer than the median flexion line in Termatophyloides. 311 Description.—Ovoid; coriaceous; rows of punctures on pronotum (Fig. 42), anal vein and R+M veins (Fig. 55); moderate distri- bution of semierect, elongate setae. Head: elongate (Fig. 10), porrect (Fig. 18); vertex rounded; frons declivent, rounded; clypeus short, weakly declivent, terminating above an- tennifers; bucculae short, margins weakly ar- cuate (Fig. 26), not surpassed by first labial segment; gula elongate, narrow, weakly con- cave, lateral margins carinate, with transverse ridges, and a row of soft, moderate-sized, semi- erect setae adjacent to eyes. Eyes: large, almost contiguous with pronotal collar, distance less than the width of the second antennal segment; ommatidia moderately large; elongate intero- cular setae. Antennae: short; AI subcylindrical; AII weakly incrassate distally, AI] subequal to AII length, longer than ATV; AIII and AIV weakly fusiform. Labium: reaching apices of forecoxae; LII longest segment. Pronotum: (Fig. 42) weakly biconvex; lateral margins moder- ately divergent; collar broad, almost reaching lateral margins of eyes, separated from callosite region by a deep, nonpunctate groove, callosite region marked by a postero-transverse and sublateral, punctate groove, and punctate groove along midline; pronotal antero-lateral setae caudad of propleural suture; disc weakly biconvex, longer than callosite region, poste- rior margin truncate. Mesonotum: barely vis- ible from above, strongly declivent. Scutellum: flattened, lateral margin not punctate. Hemel- ytra: broad, extending well beyond lateral mar- gins of pronotum; lateral margins arcuate; anal vein punctate; embolium broad, subequal to pronotal collar length; punctate R+M vein short, only slightly surpassing 7/3 length of co- rium; medial flexion line much longer, and strongly divergent from the R+ M vein; cuneus as long as wide; membrane translucent, dusty concolorous, without microsetae; membrane cell moderately broad, subelliptical, with a weak expansion subcaudally. Thoracic pleura: ter- gopleural suture distinct anteriad to propleural suture; proepisternum subquadrate, granulate; proepimeron weakly rugose, without pits; me- sepimeron (Fig. 64) moderately short, broad, 32 with dorsal margin strongly angulate, dorso- caudal arm of margin with a lanceolate, spi- racular opening with evaporative areas, ven- tral margin marked by a weak depression, and a row of elongate, anteriorly directed, semiad- pressed setae; metepisternum scent efferent system (Fig. 64) occupying about half of the segment, contiguous with metathoracic evap- orative area; peritreme submarginal to poste- rior margin of segment, dorso-caudally pro- jected, tongue-shaped, almost reaching dorsal margin of evaporative areas, not raised above plane of evaporative area; evaporative bodies mostly rounded, remainder of metepisternum depressed, with elongate, semiadpressed setae. Legs: moderately sized; coxae elongate; femora fusiform; three mesofemoral and four meta- femoral trichobothria; tibiae cylindrical, not slender, with four rows of spinelets, and stiff, semiadpressed setae; pretarsi with basally toothed claws and setiform parempodia. Ab- domen: broad, semipolished. Male genitalia: genital capsule small, genital opening small; left paramere at right angles to ventral margin of genital opening, shaft broad. Male aedeagus not examined. Remarks. —Carvalho (1955b) described Termatophyloides from a single female spec- imen from Mexico. Carvalho (1955a) distin- guished this genus from other termatophylines by the presence ofa transverse punctate groove along the posterior margin of the callosite re- gion, elongate pubescence, and the labium ex- tending to the apex of the forecoxae. None of these characters are exclusive to Termatophy- loides, and can no longer be considered diag- nostic for the genus. A lack of material does not allow an examination of the male genitalia, however the genital capsule and the left par- amere, indicate a close relationship with the other Central American genus Termatophy- lella. Miyamoto (1965) in describing a new species, 7. pallipes, from Japan, indicated nu- merous differences from the type species, par- ticularly in head structure. I have moved T. pallipes to Kundakimuka, on the basis of the similar head structure, although the presence of a punctate groove along the midline of the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON callosite region of this species is consistent with the present diagnosis of Termatophyloides. Termatophyloides pilosulus Carvalho Termatophyloides pilosulus Carvalho 1955b: 643 (n. sp.), Carvalho 1957: 36 (catalogue). Figures. — 10, 18, 26, 42, 55, 64. Holotype.— Female. Mexico: Tamazun- chale, San Luis Potosi, 30-XII-1948, coll[ector] Insp[ector] Leary, Laredo Tex{as], 49529, on orchids (USNM, 62946; type seen). Other specimens examined.—Mexico on orchids, 10-10-1953, Brownsville, Tex[as] (USNM, male). Diagnosis.—This species 1s recognized by the dark coriaceous body, the presence of a punctate groove along the midline of the cal- losite region, and the elongate R+M vein. Description.— Male. Small. Length 2.17, width 1.20, mostly black, with elongate, sem- iadpressed vestiture. Head: conical, small, a little longer than wide, length 0.41, width 0.37, dark brown, with reddish tinge, particularly on apex of clypeus, and maxillary and mandibular plates. Eyes: red, removed from pronotal col- lar by a distance equal to width of second antennal segment, ommatidia of moderate size. Antennae: very small, mostly cylindrical, with a moderate distribution of pale, semiadpressed setae; AI small, a little thickened, length 0.13, dark brown; AII weakly expanded distally, mostly yellow, apex with a weak reddish tinge, length 0.24; ATII mostly yellow, with an apical red annulus, length 0.14; ATV red, length 0.14. Labium: yellow, small, reaching midlength of forecoxae; LIT reaching midlength of eyes, lon- gest segment; LIII reaching proxyphus. Pro- notum: uniformly black, antero-lateral setae dorsad of tergopleural suture, and anteriad of propleural suture. Legs: coxae mostly dark brown, apices with reddish tinge; trochanters reddish; femora mostly dark brown, some- times with obscure reddish tinge, apices yel- low; tibiae yellow; tarsi yellow, THI with ob- scure embrownment, longest segment. Abdo- men: dark brown, broad, anthocoridlike, ven- VOLUME 97, NUMBER 2 ter with regular, moderately distributed, elongate, adpressed, pale setae, posterior seg- ments with a few elongate, erect setae, partic- ularly laterally. Male genitalia: see generic de- scription. Female. Similar to male in color, shape and vestiture. Length 2.17, width 1.10. Head: length 0.40, width 0.34, interocular distance 0.16. Pronotum: length 0.40, posterior width 0.84. Antennae: AI length 0.14; AII length 0.24; AIII length 0.16; AIV length 0.16. Pronotum: length 0.40, posterior width 0.84. Remarks.—This species, like Termatophy- lella fulvoides, has been taken on orchids, which probably represents a sitting record. It was in- tercepted in quarantine in Texas, USA. More material is needed for examination to firmly establish the relationships of this species. Termatophylum Reuter Termatophylum Reuter 1884a: 218 (n. gen.), Reuter 1884b: 167 (description), Reuter and Poppius 1912: 3, 5 (key; description), Car- valho 1952: 50 (list), Carvalho 1955a: 23 (key), Stichel 1956: 175 (key), Carvalho 1957: 36 (catalogue). Figures.—6, 14, 22, 30, 34, 37, 46, 56, 60, 68, 72, 76, 80, 93, 94, 99, 100, 108, 109, 110, MiG e ls: Type species.— Termatophylum insigne Reuter, 1884 by monotypy. Diagnosis.— 7ermatophylum is separated from other termatophylines by the abbrevi- ated punctate R+M vein (Fig. 56), which is shorter than the medial flexion line. The mid- line of the callosite region has a punctate groove (Figs. 34, 37) in Termatophylum, which is not the case in the saliently similar species of Kun- dakimuka (Figs. 31, 35). Also, the posterior margin of the callosite region of Termatophy- lum is punctate (Fig. 34), which is not the case in the phylogenetically related genus Arygro- telaenus (Fig. 39). Description.—Elongate to elongate-ovoid; semipolished to weakly coriaceous; with rows of punctures on anal and R+M veins (Fig. 56), pronotum (Fig. 34) and scutellum, with 313 sparse to moderate cover of semierect, pale setae. Head: transverse (Fig. 6) weakly porrect (Fig. 14); vertex weakly to moderately round- ed, posterior margin weakly carinate; frons weakly declivent; clypeus moderately decli- vent, terminating above antennifers; lateral margins of bucculae subparallel (Fig. 22), ter- minating caudally at midpoint of antennifers; gula elongate, narrow, concave, with trans- verse ridges, and matt of microsetae (Fig. 22). Eyes: very large, covering the lateral aspect of the head, almost touching pronotum (Fig. 14), separated by a distance less than the width of the second antennal segment, ommatidia large, with short to elongate, interocular setae. An- tennae: short to moderately long, with sem- iadpressed setae, in males intermixed with flat- tened, scalelike, adpressed setae on AII; AI short, weakly thickened; AII swollen and fu- siform in male (Fig. 30), cylindrical and weakly incrassate subapically in females, AIII cylin- drical, ATV weakly fusiform, not swollen. La- bium: reaching between apices of forecoxae and apex of mesosternum; LII longest seg- ment. Pronotum: (Figs. 34, 37, 46) lateral mar- gins strongly divergent, weakly to moderately biconvex; collar rounded, not wide, barely reaching midwidth of eyes, posterior margin marked by a deep punctate groove; callosite region (Fig. 34), marked by a transverse and sublateral, punctate groove, and a short, mid- line, punctate groove (Fig. 37); calli separated and moderately tumid antero-lateral pronotal setae anteriad of propleural suture; disc longer than callosite region, moderately raised above callosite region in lateral view, posterior mar- gin truncate. Mesonotum.: short to very short, visible from above. Scutellum: flattened to weakly rounded, lateral margins each with a row of punctations, rarely along entire length. Hemelytra: (Fig. 56) not broad, barely sur- passing posterior angles of pronotum, margins subparallel to weakly arcuate; anal vein punc- tate; embolium narrow, mostly less wide than width of second antennal segment of male; punctate R+M vein, usually surpassing mid- length of medial flexion line, rarely beyond, sometimes with punctations less deep and dis- 314 tinct than punctures of anal vein and prono- tum; medial flexion line elongate, almost reaching cunea! fracture, not strongly divergent from R+M vein, subparallel anteriorly; cu- neus as wide as long; membrane translucent to opaque with a bicolored pattern, usually pale anteriorly and brown posteriorly, mem- brane cell large, subquadrate, vein somewhat indistinct, sometimes thickened subcaudally. Thoracic pleura: tergopleural suture distinct anteriorly (Fig. 46); proepisternum, subtrian- gular; proepimeron with a pit ventro-mesally placed; mesepimeron (Fig. 60) broad, dorsal margin strongly angulate, dorso-caudal arm with a small band of evaporative areas, spi- racular opening (Fig. 68) not visible, recessed, ventral margin of mesepimeron marked by a depressed line; metepisternum scent efferent system (Fig. 60) occupying almost half of seg- ment, not contiguous with evaporative areas of mesepimeron, with peritreme at midlength or just posteriad of mid-length of evaporative areas, peritreme (Fig. 72) weakly depressed, lanceolate and narrow, to moderately broad and tonguelike, not raised dorsally or poste- riorly above plane of evaporative areas; evap- orative bodies variable, mostly elongate (Fig. 76), remainder of segment weakly depressed with a few semierect, pale setae. Legs: mod- erately long; femora fusiform, not greatly swol- len; four mesofemoral and six metafemoral trichobothria; tibiae cylindrical, without any rows of spinelets, with stiff, semiadpressed pale setae; pretarsi (Figs. 94, 100) with claws strong- ly cleft basally, and angulately C-shaped, with short, flattened, ribbonlike parempodia. Ab- domen: broad, polished, with moderate dis- tribution of soft, semiadpressed setae; Male genitalia: genital capsule conical (Fig. 101, 102); genital opening small; left paramere (Figs. 108— 110) with a very short sensory lobe, with a few short, erect, pale setae, shaft arcuate, some- times evenly tapered apically, sometimes broader and spatulate apically; vesica (Figs. 116-118) with a membranous lobe, with a gonoporal cavity, and one or two sickle-shaped, lobal sclerites, apex of membranous lobe with fields of spines. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Remarks. — Termatophylum is the most di- verse genus of the termatophylines, and pres- ently contains 13 species, of which, all except one, are confined to the Old World, particu- larly the tropical and subtropical regions. Ter- matophylum montanum was described from Brazil by Carvalho (1983) but has not been examined in this study. Reuter (1884a) first described the genus for a species from Egypt. Reuter and Poppius (1912) provided a detailed redescription of the genus and listed diagnostic characters for Termatophylum in a key to the known genera. They separated Termatophy- lum and Arygrotelaenus primarily on the pres- ence of hairy eyes in the former genus, which remains a valuable distinguishing character. Carvalho’s (1955a) generic key to the terma- tophylines did not contain any significant di- agnostic characters, and incorrectly stated that the pronotum did not have rows of punctures behind and between the calli. Termatophylum is now clearly recognized by the punctation of the pronotum and characters of the ptero- thoracic pleura and male genitalia. The biology of Termatophylum is poorly known with host plant records known for T. insigne and T. me- laleucae (see Table | and Biology section). Termatophylum insigne Reuter Termatophylum insigne Reuter 1884a: 218 (n. sp.), Reuter 1884b: 168 (description), Reu- ter and Poppius 1912: 7, 9 (key, description), Carvalho 1952: 50 (type species), Stichel 1956: 175 (description), Carvalho 1957: 36 (catalogue), Linnavuori 1974: 3, 4 (diag- nosis; species key), Linnavuori 1975: 6 (Egypt; host plant), Linnavuori 1980: 15 (Sudan), Linnavuori 1989: 29 (Yemen). Figures.—93, 99, 108, 116. Holotype.—Female. Egypt (as Aegyptus) (?UZMH,; not seen). Specimens examined.—Sudan: Kordofan, Tendelti-Umm Ruwaba, 25-1-1963, R. Lin- navuori (NHM; male); Sudan: N[orth- ern] Prov[ince] Ed Dammer-Shendi 1-XI]I- 1962, R. Linnavuori (NHM; male); Sudan: Khartoum, junction of Niles, 23-1-1966, P. VOLUME 97, NUMBER 2 Stys collector (AM; female); Sudan: Khar- toum, Sunt Forest, 19-11-1965, P. Stys col- lector (AM; 3 males; female); Sudan: Khar- toum, Burri, 10-1965, P. Stys collector (AM; female); Sudan: Khartoum, Burn, 10-12-1965, Williams collector (AM; male). Diagnosis. — Termatophylum insigne is dis- tinguished by the color pattern of the dorsum; mostly brown, with yellowish markings. The male left paramere (Fig. 108) and the vesica (Fig. 116) are also distinctive for the species. Description.— Male. Small, parallel-sided, length 2.33-2.64, width 0.84-0.93, mostly yel- lowish-brown, with brown and yellow mark- ings, hemelytra opaque, with sparse cover of semiadpressed, short, golden setae. Head: uni- formly brown, with uniform distribution of short, whitish, adpressed setae, moderately porrect, length 0.43-0.48, width 0.53-0.56, in- terocular distance 0.14-0.17; maxillary and mandibular plates yellowish-brown, some- times with reddish; gula yellow. Eyes: reddish- black, with moderately elongate, interocular setae. Antennae: AI short, length 0.14~0.17, barely passing apex of clypeus, yellow, with a few semiadpressed setae; AII length 0.39-0.44, fusiform, uniformly swollen, proximal half to two-thirds yellow, distal half to third brown, sometimes with reddish tinge, particularly at apex, with moderate cover of flattened, ad- pressed, scalelike, whitish setae, intermixed with elongate, semiadpressed setae; AIII cylin- drical, much narrower than AII, length 0.22- 0.24, yellow, with moderately sized semiad- pressed, pale setae AIV weakly fusiform, length 0.14-0.17, vestiture as AI. Labium: slender but not fragile, reaching apices of forecoxae, length 0.98-1.00, LIT longest segment but only slightly longer than LIT. Pronotum: with sparse cover of moderately long, semi-erect, pale se- tae; collar yellowish-brown, laterally more brown; callosite region yellowish-brown, with grooves and lateral regions darker brown; disc uniformly brown. Mesonotum: barely visible from above. Scutellum: anterior half brown, posterior half yellow to yellowish-brown, with sparse cover of semierect, moderately sized, pale setae. Hemelytra: with sparse cover of S15 semiadpressed, moderately sized, pale setae on clavus, embolium, corium and cuneus; clavus yellowish brown, with inner margins darker; embolium R+M vein short, about *4 length of medial flexion line, R+M vein only weakly punctate, mostly yellowish, posteriorly em- browned; corium with anterior 74’s laterad of medial flexion line vein yellow to yellowish- brown, remainder brown, except for indistinct yellow marking mesally above cuneal fracture; cuneus antero-laterally yellow, remainder brown; membrane with vein embrowned, membrane patterned, mostly brown, with two indistinct yellow markings posteriad of cuneus, without any apparent microsetae. Thoracic pleura: proepisternum proximally brown, dis- tally yellow, proepimeron proximally brown, distally yellow; proxyphus uniformly yellow. Legs: moderately sized, mostly yellow with distal half of metafemora shiny brown; me- tafemora almost 1.5x< longer than mesofe- mora; tibiae and tarsi yellow. Abdomen: broad, anthocoridlike, brown, shiny. Male genitalia: left paramere (Fig. 108); vesica (Fig. 116) with a sickle-shaped lobal sclerite, and two ill-de- fined, basal lobal sclerites. Female similar to male in color, shape and vestiture. Length 2.67-2.79, width 0.86-0.96. Head: \ength 0.45-0.49, width 0.52, intero- cular distance 0.16-0.18. Antennae: AI length 0.15-0.16, uniformly dark brown; AII cylin- drical, weakly expanded distally, yellow; ATII length 0.24, yellow; AIV length 0.16-0.18, yel- low. Labium: length 0.94-0.96. Pronotum: length 0.59-0.60, posterior width 0.82-0.88. Remarks.—This species is broadly distrib- uted in the Middle East, including the Sudan, and is known to inhabit Acacia (Linnavuori 1975). Reuter (1884a) briefly described the species from a single female specimen. Reuter and Poppius (1912) gave a detailed description of the species, including the male, and differ- entiated it from other Termatophylum species, primarily on color characteristics. Stichel (1956) briefly described the species, and Linnavuori (1974) gave a detailed diagnosis, including structural and color features. The majority of Termatophylum species are differentiated on 316 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON external color characters, and a study of the male genitalia is necessary to confirm the va- lidity of these species. Termatophylum melaleucae Cassis n. sp. Figures.—6, 14, 22, 30, 34, 37, 46, 56, 60, 68, 72, 76, 94, 100, 108, 109, 117. Eytomology.—This species is named after its host-plant genus association. Holotype.— Male. QLD: 19:18.2S 146: 49.1E, Townsville, Oonoonba Kennan St[reet] 19 May 1993 ike Brown, NQMgqn93028.PO04 1-042, ex Melaleuca quin- quenervia (AM). Specimens examined.—Paratypes: QLD: 19: 18.2S 146:49.1E, Townsville, Oonoonba Ken- nan St[reet], 19 May 1993, L. M. Brown, NQMgqn93028.PO4 1-042, ex Melaleuca quin- quenervia (AM; 1 male; 2 females); QLD: Heathlands, 11:45S 142:35E, 18 August 1992, J. Cardale, P. Zborowski, at light (ANIC; male); QED» 1'8:07.98 145:55E,, 2128: kmeS, Tully: Double Barrel C[ree]k Swamp, 11 May 1993, J.K. Balciunas, NQMMgqn93026.P099, ex Melaleuca quinquenervia (AM; female). Diagnosis.— 7ermatophylum melaleucae is distinguished by the bicolored hemelytral membrane. It is also differentiated from its close relative, 7. weiri, by the spatulate apex of the left paramere (Fig. 109), and the single lobal sclerite (Fig. 117). Description.—Male. Small, length 2.51- 2.73, width 0.96-1.02, parallel-sided, mostly dark brown, with moderate cover of pale, semierect setae. Head: length 0.38—0.48, width 0.51-0.53, interocular distance 0.12-0.15, ver- tex and frons mostly dark brown to black, clypeus, maxillary and mandibular plates dark brown, sometimes with reddish tinge. Eyes: reddish-dark brown. Antennae: with moderate cover of semierect setae, less so on AI; AI length 0.16-0.18, dark brown, with moderate to strong reddish tinge; AIT length 0.36-0.41, mostly dark brown with reddish tinge, some- times with base yellow, also with short flat- tened, scalelike setae; AIII length 0.23-0.24, dusty yellow; AIV length 0.21-0.24, dusty yel- low. Labium: length 0.96-1.08, yellow, LII longest segment, reaching base of forecoxae, LHI reaching apex of forecoxae, LIV reaching posterior margin of mesosternum. Pronotum: dark brown to black, grooves darker, posterior margin with a minute, pale yellowish band; length 0.49-0.58, width 0.84-0.86. Scutellum: flattened, mostly dark brown to black, apex with yellow marking. Hemelytra: color vari- able, mostly with clavus and cuneus dark brown to fuscous, embolium reddish-brown to brown, with corium brown, but darker mesally, but sometimes hemelytra lighter brown, with cla- vus and corium with reddish tinge, and lateral aspect of corium almost translucent, medial flexion line and R+M veins usually darker; R+M vein occupying ¥, to % of corium, medial flexion line more than “% of corium; membrane with moderate cover of pale microsetae, bi- colored, with anterior half pale to translucent, remainder pale to medium brown, anterior margin of darker areas V-shaped, cell mod- erate size, mostly pale, with vein weakly raised, and caudal half embrowned, and weakly thick- ened caudo-laterally. Legs: mostly yellow. Genitalia: left clasper (Fig. 109) with apex of shaft spatulate; vesica (Fig. 116), with one sick- le-shaped lobal sclerite. Female similar to male in shape but more elongate-ovoid; color similar, in some speci- mens hemelytra sometimes more uniformly dark brown, with lighter markings along cu- neal fracture; vestiture similar to males. Length 2.63-3.04, width 1.10—1.20. Head: length 0.48- 0.49, width 0.52-0.54, interocular distance 0.16-0.17. Antennae: AI length 0.19-0.24, dark brown to fuscous; AIT length 0.37-0.43, weak- ly incrassate apically, proximal *4 yellow, re- mainder dark brown to fuscous, sometimes with reddish tinge; ATI] length 0.24-0.26, dusty yellow; AIV length 0.21-0.22, dusty yellow. Labium: length 1.02-1.08. Pronotum: length 0.48-0.50, posterior width 0.90-0.94. Remarks.—This species is easily recognized by the patterned hemelytral membrane. The description of this species is based on speci- mens from northern Queensland, however, numerous specimens from southern Queens- land, acquired after the observational phase of VOLUME 97, NUMBER 2 this work, appear to be conspecific, which sug- gests that the species is broadly distributed in the coastal region of the State. This needs to be confirmed by examination of the male gen- italia. Termatophylum weiri Cassis n. sp. Figures.— 110, 118. Eytomology.—This species is named after the collector Tom A. Weir, of the Australian National Insect Collection, CSIRO, Canberra. Holotype.— Male. QLD: 15.47S 145.17E, Moses C[ree]k, 4 km N by E of Mt. Finnigan, 14-16 October 1980, T. Weir (ANIC). Other specimens examined. —Paratypes: QLD: 15.47S 145.17E, Moses C[ree]k, 4 km N by E of Mt. Finnigan, 14-16 October 1980, T. Weir (ANIC; | male; 2 females); QLD: Cairns district, F. P. Dodd (SAMA; female). Diagnosis.— Termatophylum weiri is rec- ognized by the unicolorous hemelytral mem- brane, the evenly tapered apex of the left clasp- er (Fig. 110), and the vesica with two lobal sclerites. Description.—Male. Small, length 2.29- 2.33, width 0.85-0.90, parallel-sided, mostly light brown, with sparse cover of elongate, semiadpressed pale setae. Head: length 0.36- 0.40, width 0.46—-0.52, interocular distance 0.12, vertex fuscous, clypeus, maxillary and mandibular plates reddish-brown. Eyes: fus- cous, with a reddish tinge. Antennae: Al length 0.14-0.15, not thickened, yellowish-brown, with a few erect, pale setae; AII length 0.38- 0.40, yellowish-brown, with moderate cover of erect, brown, elongate, bristlelike setae, and short, flattened, adpressed pale setae; AIII length 0.22-0.24, cylindrical, narrow, yellow, with moderate cover of elongate, semierect, pale setae; AIV length 0.16-0.17, narrow, weakly fusiform, with moderate cover of elon- gate, semierect, pale setae. Labium: length 0.88— 0.96, reaching beyond midlength of mesoster- num; LII longest segment, reaching midlength of forecoxae; LIII reaching beyond apices of forecoxae. Pronotum: subtrapezoidal, mostly yellowish-brown; collar light yellowish-brown; callosite region yellowish-brown, but darker 317 than pronotal disc and collar; pronotal disc yellowish-brown. Scutellum: brown, with an- terior half with fuscate tinge, lateral margins with weak punctations. Hemelytra: clavus yel- lowish-brown, with darker embrownment par- ticularly along anal vein; cortum yellowish brown, R+M vein punctations not deep, not embrowned, barely surpassing midlength of medial flexion line, setal bases fuscous; mem- brane yellowish, translucent, without any color pattern, membrane cell large, subquadrate, vein weakly embrowned. Genitalia: left clasper (Fig. 110) with apex of shaft evenly tapered; vesica (Fig. 117) with two sickle-shaped lobal scle- rites. Female. Similar to male in structure and color. Length 2.17-—2.29, width 0.93-0.96. Head: length 0.38, width 0.46, interocular dis- tance 0.12-0.14. Antennae: Al length 0.16, AI length 0.36, weakly incrassate apically: AIII length 0.22-0.24; ATV length 0.18-0.19. La- bium: length 0.89-0.96. Pronotum: length 0.50- 0.53, posterior width 0.78—-0.82. Remarks.—There is some doubt as to whether the female from Moses Creek, QLD, is conspecific with the holotype male from the same locality, as it is considerably darker. The female from the Cairns district is saliently very similar to the holotype male, particularly in the lighter coloration. GENERA REMOVED FROM THE TERMATOPHYLINI Conocephalocoris Knight, 1927 Conocephalocoris Knight 1927: 141 (n. sp.), Carvalho 1952: 53 (list), Carvalho 195Sa: 22 (key), Carvalho 1957: 35 (catalogue). Type species.—Conocephalocoris nasicus Knight, 1927, by monotypy. Specimen examined.—USA: Utah, Wash- ington Co., Leeds Canyon, 28 July 1965, W. J. Hanson collector (Utah State University; male). Remarks. — Conocephalocoris was originally described in the Deraeocorini by Knight (1927), which was followed by Carvalho (1952). Car- valho (1955a) transferred the genus to the Ter- 318 matophylini, presumably on the basis of the elongate, porrect head. His diagnosis of the Termatophylini refers to the short antennae, but in Conocephalacoris the antennae are elon- gate, with the second segment extremely long and thickened. Carvalho (1957) retained its position within the Termatophylini in his world catalogue. Conocephalocoris cannot be retained in the Termatophylini, as presently defined. There is little doubt that it is similar to termatophy- lines, in possessing rows of punctures on the pronotum, and R+M and anal veins. How- ever, these characters are present in numerous other Deraeocorinae, particularly the Hyaliod- ini. The head of Conocephalocoris is porrect, but this has evidently been independently de- rived in the termatophylines, and some Der- aeocorini genera, such as Conocephalocoris, Eurychilopterella Reuter, Hesperophylum and Termatomiris. Conocephalocoris has intero- cular setae, which has not been observed in any other non-termatophyline Deraeocorinae, and is present in all termatophylines, except Arygrotelaenus. The short first labial segment, not surpassing the posterior margin of the bucculae, is the most critically defining character for the Ter- matophylini. In C. nasicus the first labial seg- ment far surpasses the bucculae. The ptero- thoracic pleura of this species are different to the termatophylines, with the dorsal margin of the mesepimeron almost linear, carinate, and without an exposed spiracular opening, or associated evaporative areas. The peritreme of the scent efferent system of the metepisternum is spoutlike, and greatly raised above the evap- orative areas, and is unlike any Termatophy- lini. The pronotum C. nasicus has an indistinct callosite region, but is not bounded posteriorly and sublaterally by a deep groove. Numerous other characters of C. nasicus are not present in other termatophylines, and these include: 1) clavus with an incomplete punctate anal vein, and a punctate longitudinal groove, antero-mesally placed; 2) hemelytra much lon- ger than abdomen; 3) membrane with two cells; 4) labium extremely elongate, reaching last ab- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dominal segment; 4) absence of antero-lateral pronotal setae. Stonedahl (personal communication) is presently reviewing the North American gen- era Conocephalocoris, Eurychilopterella and Hesperophylum, and has indicated support for my conclusions. It is unknown at present whether these genera, and genera such as Ter- matomiris, form a monophyletic group, which are defined by having a porrect head. Hesperophylum Reuter and Poppius, 1912 Hesperophylum Reuter and Poppius 1912: 3, 16 (n. gen.), Knight 1941: 64, 74 (key), Car- valho 1952: 50 (list), Carvalho 1955a: 22 (key), Carvalho 1957: 35 (catalogue). Type species.— Hesperophylum heide- manni Reuter and Poppius, 1912, by mono- typy. Remarks.—Hesperophylum was de- scribed in the Termatophylini by Reuter and Poppius (1912), predominantly on the basis of the porrect head. This North American monotypic genus is removed from the Ter- matophylini on the same basis as Conoce- phalocoris; it has an elongate first labial seg- ment which extends far beyond the buc- culae. It also differs from termatophylines by the following characters: 1) flattened sec- ond antennal segment; 2) callosite region not demarcated posteriorly by a groove; 3) anal vein reduced to posterior half of clavus; 4) labial segment elongate, surpassing apices of metacoxae; 5) presence of two membrane cells; 6) absence of antero-lateral pronotal setae. Termatomiris Ghauri, 1975 Termatomiris Ghauri 1975: 615. Type species.—7ermatomiris probosci- docoris Ghauri, 1975, by original designa- tion. Specimen examined.— Female. QLD: Lankelly, B. Gray, 1i-vii- 1972 (NHM; para- type). Remarks. — Ghauri (1975) described this Australian taxon in the Termatophylini, VOLUME 97, NUMBER 2 primarily because of the porrect head. He indicated that it was similar to the mirine genus Proboscidocoris Reuter, but was clear- ly a member of the Deraeocorinae on the basis of the basally cleft claws and setiform parempodia. 7ermatomiris is removed from the Termatophylini, because of the elongate first labial segment, and transferred to the Deraeocorini. Termatomiris differs from termatophy- lines by the following characters: 1) first la- bial segment longer than bucculae; 2) om- matidia small; 3) absence of interocular se- tae; 5) absence of antero-lateral pronotal se- tae; 6) pronotum bipartite; 7) without rows of punctures on pronotum and hemelytra; 8) two membrane cells; 9) dorsal margin of mesepimeron not angulate; 10) metatho- racic spiracle not exposed; 11) mesepime- ron without evaporative areas; 12) peri- treme dorsally expanded and raised. PHYLOGENY OF THE GENERA OF TERMATOPHYLINI The 29 characters and the 68 character states utilized in this analysis are those of head, eyes, labium, antennae, pronotum, scutellum, hemelytra, pterothoracic pleura, parempodia and male genitalia. The char- acter and character state descriptions are presented in Table 2, and the data matrix for outgroups and Termatophylini is given in Table 3. One cladogram of 66 steps (Fig. 131) was produced, with a consistency in- dex of 0.59 and a retention index of 0.69. The character state changes for the ancestral nodes and terminal taxa are given in Table 4. The following discussion of sister-group relationships is based on the character state changes at the designated ancestral nodes. Discussion is restricted to synapomorphic characters, except for Node 9, which is only supported by homoplasious characters. Node 9.—The sister-group relationship of Hesperophylum and Conocephalocoris + Termatophylini is based on the following characters: elongate head (1-1) and the por- rect head (9-2). Both characters are labile 319 within the Termatophylini, and other Der- aeocorinae, not included in this analysis. The elongate head state is found in the basal genera of the Termatophylini, and the apo- morphic genus Arygrotelaenus, in which it is independently derived. The porrect head has been ascertained by the degree of de- clivity of the clypeus, which is also variable within the Termatophylini. Both these head characters are partially correlated, however in Termatophyloides, the clypeus is mod- erately declivent and the head is elongate. The porrect head is found in other Deraeo- corini, such as Termatomiris and Eurychil- opterella, and my analysis suggests that this character state has independently arisen of- ten in the Deraeocorinae. From this it is evident that previous definitions of the Ter- matophylini (Carvalho 1955a), based on the porrect head, are artificial, and therefore, Conocephalocoris and Hesperophylum are removed from the Termatophylini. Node 8.—The sister-group relationship of Conocephalocoris and the Termatophy- lini is based on the following synapomor- phies: interocular setae present (8-1) and the absence of a sclerotized process (28-1). The presence of interocular setae is diagnostic for all termatophylines, except for Arygro- telaenus, where the lack of setae is inter- preted as a secondary loss. The presence of interocular setae appears to be widespread within the Miridae, however in most of the Deraeocorinae, the eyes appear to lack these setae, and Conocephalocoris is the only non- termatophyline Deraeocorinae examined which has interocular setae. The examina- tion of the male genitalia outside of the ter- matophylines has been limited in this study, and the significance of the sclerotized pro- cess is not understood. Node 7.—The Termatophylini are de- fined as a monophyletic group in this anal- ysis by the following synapomorphies: short first labial segment (3-1), the presence of antero-lateral pronotal setae (12-1) and the condition of the metathoracic spiracle (21- 1). The first labial segment is extremely ab- 320 Table 2. 1) Head shape 2) Bucculae shape 3) Labial segment | 4) Gula vestiture PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Characters and character states used in cladistic analysis of Termatophylini genera. 0 = transverse, 1 = elongate 0 = parallel-sided, 1 = moderately arcuate, 2 = strongly arcuate 0 = longer than bucculae, | = as long as bucculae 0 = without microsetae, 1 = with microsetae on lateral margins, 2 = with dense matt of microsetae 5) Eye size 6) Eye position 0 = large, 1 = very large 0 = removed from pronotal collar by distance greater than AII width, 1 = contiguous with pronotal collar 7) Ommatidia size 0 = small, | = large 8) Interocular setae 9) Clypeus 10) Male AII shape ~— and swollen 11) Male AII vestiture 0 = absent, 1 = moderately distributed, 2 = densely distributed 0 = vertical, 1 = moderately distributed, 2 = densely distributed 0 = cylindrical, incrassate apically, | = fusiform and flattened, 2 = fusiform 0 = setiform only, 1 = setiform intermixed with flattened scale-like setae 12) Antero-lateral pronotal se- 0 = absent, | = present tae 13) Pronotal collar 0 = posterior margin not punctate, | = punctate 14) Callosite region 15) Callosite region 16) Scutellum 17) R+M vein flexion line 18) Hemelytral membrane 19) Membrane cell 20) Mesepimeron shape 21) Metathoracic spiracular opening 22) Scent efferent system evap- orative areas 23) Peritreme 0 = posterior margin not punctate, 1 = punctate 0 = midline without groove, | = with punctate groove 0 = lateral margins not punctate, | = punctate 0 = longer than median flexion line, 1 = equal to or shorter than median 0 = concolorous, | = bicoloured 0 = broad, | = narrow 0 = elongate, 1 = broad 0 = not evident, pleura separated, 1 = visible as a lanceolate/oval opening, 2 = not visible, recessed 0 = contiguous with mesepimeric evaporative areas, | = not contiguous with mesepimeric evaporative areas 0 = anteriad to midwidth of evaporative areas, | = at midwidth of eva- poarative areas, 2 = caudad of midwidth of evaporative areas, 3 = along posterior margin of metepisternum 24) Parempodia 25) Male genital opening 26) Gonoporal cavity 27) Lobal scletites 28) Sclerotized process 29) Vesica apex breviated in the Termatophylini, and ter- minates caudally at the posterior margin of the bucculae. This condition has not been reported in any other Miridae, nor any other Deraeocorinae examined in the course of this study. The first labial segment in the Miridae is the most robust of all the seg- ments, and greatly surpasses the posterior margin of the bucculae, and usually reaches the posterior margin of the head. Schuh and Stys (1991) in a cladistic review of the Cim- icomorpha indicate that the elongate first 0 = setiform, | = setiform, apex weakly flattened, 2 = flattened, ribbon-like O = left margin without tubercle, | = left margin with tubercle 0 = absent, | = present 0 = absent, 1 = elongate, 2 = sickle-shaped 0 = present, 1 = absent 0 = without fields of spines, 1 = with fields of spines labial segment is a synapomorphy for the Miroidea, and that the short to absent first labial segment is in the ground plan for the Cimicomorpha. The short first labial seg- ment of the Termatophylini is undoubtedly of independent origin, and cannot be pres- ently supportive of any alternative taxo- nomic arrangement of the tribe, outside of the Miroidea. Curiously, despite there being no doubt about the carnivorous habit of the Termatophylini (Callan 1975), their labium is extremely slender and fragile. The strong- VOLUME 97, NUMBER 2 Table 3. Character matrix of four outgroups, eight termatophyline genera and 29 characters. Missing da- tay=\?. Ambracius Bothynotus Hesperophylum Conocephalocoris Arygrotelaneus Democoris Kundakimuka Termatophyllella Termatophylidea Termatophylina Termatophyloides Termatophylum ly robust and elongate first labial segment of Conocephalocoris and Hesperophylum supports their removal from the Termato- phylini. The presence of antero-lateral pronotal setae appears to be unique to the Termatophylini, and is very similar in form to that found in some Anthocoridae, but Table 4. List of character state changes at ancestral nodes and terminal taxa for Figure 131. 9: 1-1, 9-2 8: 8-1, 28-1 We 2-1, 3-1, 12-1, 16-0, 21-1 6: 4-1, 7-1 >: 9-1, 14-1, 16-1, 20-1, 23-2 4: 1-0, 2-2, 6-1 3: 5-i, 8-2, 13-1 2: 21-2, 22-1, 23-1, 27-1 ile 2-1, 4-2, 10-2, 11-1, 17-1, 18-1, 24-2, 25-1, 27-2, 29-1 Ambracius 16-0 Hesperophylum 10-1 Conocephalocoris 13-1, 23-0, 29-1 Termatophylina 14-1, 17-1, 23-3, 27-1 Termatophylidea 2-0, 26-0, 28-0 Termatophyloides 15-1 Termatophylella 19-1 Kundakimuka 24-1 Democoris 19-1 Termatophylum 15-1 Arygrotelaenus 1-1, 2-0, 5-0, 8-0, 9-2, 13-0, 14-0 00000000011111111112222222222 1234567890 1234567890 123456789 020000000000000000000?100?7000 020000000000000100000?100?7000 120000012000100100000?000?7011 10120110221100011101211211211 02111112100111010011211001110 02111112100111010001102101010 021101111001010100111020077?? 101100112001000000001?1000000 11100001200101001000103001110 1111001110010111000110200???? OMT AeA O22 Aa must be considered to be independently de- rived. The condition of the metathoracic spi- racle has undoubted phylogenetic value in the Miridae (Cassis 1984, Cassis in prepa- ration). The spiracular opening in the Ter- matophylini is clearly visible, although sometimes recessed (Node 8), and is unlike that in any other Deraeocorinae. In the Ter- matophylini the dorso-caudal arm of the dorsal margin of the mesepimeron is highly modified, with the posterior angle recurved cephalically, and subsiding below the dorsal margin, at times partially fused with the an- terior margin of the metepisternum. The spiracular opening in termatophylines, ex- cept in Termatophylidea, is bounded by evaporative bodies, which presumably function as a barrier to allosomes from en- tering the respiratory system. In other Der- aeocorinae, such as Deraeocoris (Fig. 122), Hyaliodes (Fig. 120) and Trilaccus (Fig. 121), the mesepimeron and metepisternum are confluent, and the spiracular opening is as- sumed to be intersegmental, deeply re- cessed, and not visible or differentiated, as in the Termatophylini. Also, the dorso-cau- dal arm of the dorsal margin of the mes- epimeron of these non-termatophylines is not angulately recurved and recessed, and there is no evidence of evaporative areas. 322 In Bothynotus (Fig. 119) (Clivinemini) evaporative bodies are present on the ven- tro-caudal angle of the mesepimeron, and there is evidence that the dorsocaudal arm is recurved and subsided. However, the pleura are clearly separated in Bothynotus, which is suggestive of a different character state to the Termatophylini, and similar to that found in Conocephalocoris and Hes- perophylum. The structure of the mesepimeron, and particularly the metathoracic spiracle is very similar to that found in the Dicyphini (Fig. 123), and somewhat similar to the Phylinae (Fig. 126) and Orthotylinae (Fig. 125). This form of the spiracular opening cannot be considered to be plesiomorphic for the Mir- idae, because of its absence in the plesio- morphic subfamilies Isometopinae (Fig. 128) and Cylapinae (Fig. 129), and is con- sidered to be an important synapomorphy for the Termatophylini, and elsewhere in the Miridae. The striking similarity of the spiracular opening in the Termatophylini and Dicyphini is suggestive of a close re- lationship between these tribes. This has been previously suggested by Cobben (1968) whose examination of the termatophyline, Termatophylidea opaca, indicated similar- ities with the Dicyphini in the aero-micro- pylar structure of the eggs. More generally Wagner (1955) considered the Dicyphini as a sister taxon to the Deraeocorinae, pri- marily on the basis of the male genitalia. However, on the basis of the pretarsal struc- ture (Schuh 1976), the Termatophylini are considered to belong to the Deraeocorinae, and the relationship of the Dicyphini to the Deraeocorinae requires examination, and is beyond the scope of this work. Node 6.—Termatophylina in this analy- sis is the most plesiomorphic of the ter- matophylines, on the basis of the small, flat- tened ommatidia. The remainder of the ter- matophylines form a clade of seven genera on the basis of the enlarged and convex om- matidia (7-1) and the presence of microse- tae on the lateral margins of the gula (4-1). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The presence of enlarged ommatidia has not been examined in other suprageneric group of Miridae and its importance requires fur- ther examination. The gula of termatophy- lines is usually weakly concave, with trans- verse ridges, and this clade is characterized by the occurrence of microsetae on the gula, which are absent in Termatophylina. Node 5.—This clade is represented ba- sally by the Western Hemisphere genera Termatophyloides and Termatophylella, and the four Old World genera Kundakimuka, Democoris, Termatophylum and Arygrote- laenus. It is defined by the following syna- pomorphies: the broad mesepimeron (20- 1) and the peritreme posteriad to midwidth of the scent efferent system (23-2). The broad mesepimeron is correlated to the strongly angulate dorsal margin (Figs. 57, 60). In Termatophylina (Fig. 59) and Termatophy- lidea (Fig. 58) the dorsal margin of the mes- epimeron is weakly arcuate, which is more typical for other Deraeocorinae, where the dorsal margin of the mesepimeron is almost linear to very weakly arcuate (Figs. 119- 122). The angulate dorsal margin of the mesepimeron is rarely encountered in the Miridae, occurring in the Eccritotarsini (Fig. 130). This character state exists in other non- mirid cimicomorphans, such as the Antho- coridae (Fig. 124). The dorsal margin of the mesepimeron is fundamentally different in the Dicyphini (Fig. 123), Orthotylinae (Fig. 125) and Phylinae (Fig. 126), where the evaporative areas bridge to the postalare, producing a dissected dorsal margin. The position of the scent efferent system peritreme in relation to the evaporative ar- eas has some phylogenetic value in some of the mirid suprageneric groups. In the Iso- metopinae (Fig. 128) the peritreme is con- tiguous with the anterior margin of the met- episternum, and in the Eccritotarsini (Fig. 130) it is situated on the posterior margin of the metepisternum. This character shows variability within some mirid suprageneric groups, such as Termatophylini. In the most basal termatophyline, Termatophylina, the 323 VOLUME 97, NUMBER 2 ‘gpoesids o1oes0yi eau = § Sawejued = d ‘umusajsidaj}ow = sui ‘uoJowIdosoul = dU ‘sBaie SAT}RIOdPAD = Bd “(IULIODOIRIIq) “ds sOJ0aDAIG “TZ “(luLTWOTUINIeS) “ds sNIIDjIA TL “TZ ‘(1urpotjeAp) ‘ds sapoypdy ‘OT ‘(turmoutAt[O) ‘ds snjoudyjog ‘6] | ‘MOIA [esaje] ‘aeULIODOIeIBq, SUT|AYdo}eULIA} uou Jo vingjd o1I9eIOYIOINg “ZZI-6I] ‘S34 324 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 123-130. Pterothoracic pleura of Anthocoridae and non-deraeocorine Miridae. 123, Dicyphus sp. (Bryocorinae: Dicyphini). 124, Temnostethus sp. (Anthocoridae). 125, undetermined orthotyline species. 126, undetermined phyline species. 127, undetermined mirine species. 128, undetermined isometopine species. 129, Cylapus sp. (Cylapinae). 130, undetermined eccritotarsine species. ea = evaporative areas; me = mesepimeron; ms = metepisternum; p = peritreme; s = metathoracic spiracle. VOLUME 97, NUMBER 2 10 ©) 8 28 14 17 23 27 @) | 2 3 12 16 21 2 26 28 325 Ambracius Bothynotus Conocephalocoris La are ae SS oe ee Hesperophylum 19 13 23 29 Termatophylina ©) eT Maee wkiaG ar ee 47 © [iaraa eee 5-0 hi a ees | 9 14 16 20 23 ig a aA arn Fig. 131. = = ee oe a 24 = cai eee | 5 8 LO 19 | 21 22 23 27 Democoris 125 8 9 13 14 2 4 1011 17 18 24 25 27 29 Arygrotelaenus CHHHHHHHHH©: L{}—_____— Termatophylum Cladogram depicting phylogenetic relationships of the genera of Termatophylini. Black bars = synapomorphies; stippled bars = homoplasious forward; white bars = homoplasious reversal). peritreme is contiguous with the posterior margin of the metepisternum (Figs. 59, 71). The peritreme caudad of the midwidth of the scent efferent system occurs in the three most basal genera of Node 5, and is reversed (23-1) for the clade of Node 2. Node 4.—This clade of five genera (T7er- matophylella + Kundakimuka + Demo- coris + Arygrotelaenus + Termatophylum) is defined by the following synapomorphy: the eyes contiguous with the pronotal collar (6-1). This character is partly correlated to very large eye size, however, in Termato- phylella the eyes (6-0) are not as large, but are still contiguous with the pronotal collar (Fig. 9). Node 3.—This clade of four genera (Kun- dakimuka + Democoris + Arygrotelaenus + Termatophylum) is defined by the fol- lowing synapomorphies: very large eyes (5- 1) and densely distributed interocular setae (8-2). In this clade the genera have the eyes occupying almost the entire lateral aspect of the head, with the entire posterior margin contiguous with the collar in lateral view. The interocular setae are densely distrib- uted in this clade, and are nearly always greatly elongate, greatly surpassing the max- imum height of the ommatidia. Node 2.—This clade of three genera (De- mocoris + Arygrotelaenus + Termatophy- lum) is defined by the following synapo- morphies: the metathoracic spiracular opening recessed (21-2), and the scent ef- ferent system not contiguous with the mes- epimeric evaporative areas (22-1). In these genera the metathoracic spiracular opening is not visible when the pleura are observed in lateral view (Figs. 60, 68) and its presence is indicated by a short band of evaporative areas. In postero-lateral view, the spiracle is observed as a small, recessed opening, which is most obvious in Democoris, and is very small in Jermatophylum and Ary- grotelaenus. In many mirids the evapora- tive areas of the mesepimeron and met- episternum are contiguous. The scent effer- ent system, however, is often reduced, and in this clade, the separation of the two evap- orative areas is considered phylogenetically significant. Node 1.—The sister-group relationship of Termatophylum and Arygrotelaenus is strongly supported by the following syna- pomorphies: the gula with dense matt of microsetae (4-2), the male second antennal segment fusiform and swollen (10-2), the male second antennal segment with seti- 326 form and flattened scalelike setae (11-1), the parempodia broadly flattened and ribbon- like (24-2), the hemelytral membrane bi- colored (18-1), the left margin of the male genital opening with a small tubercle (25- 1), and the lobal sclerites sickle-shaped (27- 2): The vestiture of the gula of this clade is characterized by a dense matt of microsetae (Figs. 22, 23), occupying the entire region. This is somewhat variable in Termatophy- lum, but is found in the type species, T. insigne. The vestiture of the male second antennal segment of Arygrotelaenus and Termatophylum is composed of setiform, semierect setae, intermixed with flattened, scalelike setae (Fig. 30). This does not occur in any other termatophyline, and is possibly correlated with the presence of flattened, scalelike setae in Arygrotelaenus and some species of Termatophylum. The flattened, ribbonlike parempodia of Arygrotelaenus (Figs. 89, 95) and Termatophylum (Figs. 93, 94, 99, 100), represents the first reported variation of the deraeocorine setiform par- empodial type, and strongly supports the sister-group relationship of these two gen- era. These parempodia retain longitudinal striations which also supports their deri- vation from a setiform type. Schuh (1976) has indicated that in the Miridae the la- mellate type has been derived from the se- tiform type, and this case is evidence that this has occurred in the Miridae on more occasions than previously thought. This clade is supported by two make genitalic characters. The most significant of these is the presence of sickle-shaped lobal sclerites (Figs. 111, 116-118), whereas in all other termatophylines the lobal sclerites are lin- ear, with the apex often tapered. DISCUSSION I conclude that the Termatophylini be- long to the Deraeocorinae, which are pres- ently defined by the presence of basally cleft claws, and the presence of setiform par- empodia. Schuh (1976) supported the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON monophyly of the Deraeocorinae on the ba- sis of the pretarsal structure, and also in- dicated that most deraeocorines have a pol- ished and punctate pronotum, and a ten- dency towards reduction in femoral tricho- bothrial numbers. The deraeocorine pretarsus, however, remains the only syna- pomorphy for the subfamily, and no addi- tional support has been gained from my ob- servations of the pterothoracic pleura. De- spite the existence of some variation in the parempodia in the termatophylines, with Arygrotelaenus and Termatophylum having flattened parempodia, other characters do not strongly support an alternative taxo- nomic arrangement. The similarity of the Dicyphini and Termatophylini metathorac- ic spiracle is out-weighed by the great dif- ferences in their pretarsal structure. The male genitalia of the Termatophylini is con- sistent with those of other Deraeocorinae (Kelton 1959), although the secondary gon- ophore appears to be considerably reduced. The parameres are strongly asymmetrical, with the right paramere minute, which is again reminiscent of the Dicyphini, al- though this condition also occurs in other Deraeocorinae, such as the Hyaliodini and Clivinemini. Schuh (1976), Akingbohungbe (1974, 1978), and Stonedahl and Cassis (1991) pointed to the inadequacies of the defini- tions of the deraeocorine tribes. Akingbo- hungbe (1974) made an analysis of various genera of Hyaliodini, and supported Knight (1943) in considering them as a subfamily, primarily on the presence of an anal tube. However, he recognized two groups within the hyaliodines, in which one group did not contain the anal tube, and no other char- acter presented as a synapomorphy for the group. The glassy appearance of the he- melytra in many Hyaliodini strongly resem- bles the condition in the termatophyline Termatophylidea, which is superficially similar to Hyaliodes, although the heads are very different. The hemelytra of the hy- aliodines and termatophylines are similar VOLUME 97, NUMBER 2 in structure, with the R+M and anal veins punctate in many of the genera of both tribes. This also occurs in other deraeocorines in- cluding Conocephalocoris, Eurychilopterel- la and Hesperophylum, and some Clivine- mini. The expanded embolium is also shared by the former two tribes, although it is somewhat reduced in some termatophy- lines, such as Arygrotelaenus, Termatophy- lum and Termatophylina. It is clear that the glassy hemelytra and enlarged embolium are not sufficient characters for defining the Hy- aliodini (Carvalho 1955b), and the tribe re- mains without a reasonable diagnosis. The Clivinemini were defined by Car- valho (1952, 1955a) as having the “‘pro- notum with an impressed line running from [the] antero-lateral corner to the posterior margin of [the] calli.” This is also the case in the Termatophylini, with the line some- times marked by a row of punctures. This groove demarcates the callosite region, which is generally much shorter in the Cli- vinemini than in the Termatophylini. There appears to be little character support for the definition of the Clivinemini as a suprage- neric group. I have indicated that the me- sepimeron and metepisternum are widely separated in the clivenemines, but this also occurs in some Deraeocorini, and thus does not define the tribe exclusively. Carvalho (1952) erected the Saturniom- irini as a deraeocorine tribe on the basis of characters of the pronotum, eyes, and he- melytral membrane. He indicated that the pronotum was constricted anteriorly, but this is variable within the tribe, as pointed out by Akingbohungbe (1978), and in my examination of a number of undescribed saturniomirines from Australia. Carvalho (1952, 1955a) also indicates that the calli are large and fused, which is mostly true within the tribe but also occurs in some Cli- vinemini and some Termatophylini. The one-celled membrane is variable within the Saturniomirini, and Akingbohungbe (1974) and Schuh (1976) have indicated that this character has little taxonomic value in the 327 Miridae. Curiously, the one-celled mem- brane is invariant in the Termatophylini. Carvalho (1952) reported that the eyes of saturniomirines are semistylate, which ap- pears to be consistent in all members of the tribe examined. At present it is difficult to evaluate this character because not enough material within the remainder of the sub- family was available for examination. Carvalho (1952, 1955a) defined the Der- aeocorinae chiefly on the basis of absent characters. His key couplet read as follows: “‘pronotum not constricted anteriorly; calli not prominent and fused; neither are eyes semistylate; membrane usually with two cells.” It is obvious that this definition does not provide any diagnosis or synapomorphy for the nominate tribe of the Deraeocorinae. Stonedahl and Cassis (1991) suggested that the tribe was paraphyletic, and was basically a taxon that contains the genera that could not be placed within the more narrowly de- fined tribes. They established the sister- group relationship of the Deraeocorinae genera Angerianus and Fingulus, and sug- gested that a broad survey of the male gen- italia would possibly enlighten the situation. I did not have any material of Surinamel- lini, and cannot comment on the definition of this tribe. It is evident that aside from the present definition of the Termatophylini, no other deraeocorine tribe can be established as a monophyletic group. Undoubtedly, supra- generic groups occur within the other der- aeocorines, such as the anal tube bearing Hyaliodini, but a thorough analysis of the genera is required to provide a useful clas- sification. The similarity of a few Termatophylini, such as Kundakimuka and Termatophylum, with some Anthocoridae is undoubtedly due to convergence. The most striking similar- ities include the abbreviated first labial seg- ment, the short antennae, with the third and fourth segments greatly reduced, and the presence of large, forward projecting setae, on the antero-lateral corners of the prono- 328 tum. However, termatophylines possess femoral trichobothria, a mirid-type pretar- sus, the hemelytra! membrane with a closed cell, and typical mirid male genitalia. There is also no evidence that any of the terma- tophylines have copulatory organs associ- ated with traumatic insemination, typical of most Cimicoidea. The similarities that do exist between the termatophylines and an- thocorids have obviously evolved indepen- dently, and there is no reason to remove the Termatophylini from the Deraeocorinae. LITERATURE CITED Akingbohungbe, A. E. 1974. Nymphal characters and higher classification analysis in the Miridae (He- miptera: Heteroptera) with a subfamily key based on the nymphs. Canadian Entomologist 106: 687- 694. 1978. A new hyaliodine genus for Megacoe- lum nigroscutellatum Distant with new combi- nations and tribal reassignment for some other species of Miridae (Heteroptera). Journal of Nat- ural History 12: 87-95. Betts, C. R. 1986. The comparative morphology of the wings and axillae of selected Heteroptera. Jour- nal of Zoology London (B)1: 255-282. Callan, E. McC. 1943. Natural enemies of the cacao thrips. Bulletin of Entomological Research 34: 313- SA |e . 1975. Miridae of the genus Termatophylidea [Hemiptera] as predators of cacao thrips. Ento- mophaga 20: 389-391. Carayon, J. 1971. Notes et documents sur l’appareil odorant metathoracique des Hemipteres. Annales de le Societe Entomologique de France (NS)7: 737- 770. Carvalho, J.C. M. 1951. On the major classification of the Miridae (Hem. Het.). Transactions of the 9th international Congress of Entomology (Am- sterdam) 1: 133-134. . 1952. On the major classification of the Mir- idae (Hemiptera). 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A phylogenetic analysis of Lygo- coris Reuter (Heteroptera: Miridae) with notes on life histories and zoogeography. Master’s thesis. University of Connecticut, Storrs. 69 pp. Cobben, R. H. 1968. Evolutionary Trends in Het- eroptera. Part I. Eggs, architecture of the shell, gross embryology and eclosion. Centre for Agri- cultural Publishing and Documentation Mede- deling, Wageningen. No. 151, 475 pp. Common, I. F. B. 1990. Moths of Australia. Mel- bourne University Press, Melbourne. Davis, N. T. 1961. Morphology and phylogeny of the Reduvoidea (Hemiptera: Heteroptera). Part II. Wing Venation. Annals of the Entomological So- ciety of America 54: 340-354. Ghauri, M. S. K. 1975. Anomalous Miridae (Het- eroptera) for Australasia. Journal of Natural His- tory 9: 611-618. Kelton, L.A. 1959. Male genitalia as taxonomic char- acters in the Miridae (Hemiptera). Canadian En- tomologist, Supplement 11: 1-72. Knight, H. H. 1927. 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Bemerkungen zum System der Mir- idae (Hem. Het.) Deutsche Entomologische Ziet- schnift 2: 230-242. 1963. XII. Heteropteran. Annalen des Na- turhistorischen Museum Wien 66: 477-487. 1970. Die Miridae Hahn, 1831, des Mittel- meerraumes und der Makaronesischen Inseln (He- miptera, Heteroptera). Entomologische Abhan- dlungen (Supplement) 37: 1-273. Wiley, E.O. 1981. Phylogenetics. Theory and Prac- tice of Phylogenetics and Systematics. John Wiley & Sons, New York. Wootton, R. J.and C. R. Betts. 1986. Homology and function in the wings of Heteroptera. Systematic Entomology 11: 389-400. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 331-339 REVIEW OF THE NEW WORLD LACE BUG GENERA ACANTHOCHEILA STAL AND CARVALHOTINGIS NEW GENUS (HETEROPTERA: TINGIDAE) RICHARD C. FROESCHNER Department of Entomology, National Museum of Natural History, Washington, D.C. 20560. Abstract.—Acanthocheila Stal is revised and redefined with removal of the new genus Carvalhotingis. Of the 16 species formerly cataloged in Acanthocheila, 8 are retained but reduced to 7 with 1 new synonymy: A. abducta White, A. armigera Stal (= A. nigrescens Drake and Bondar), A. dira Drake and Hambleton, A. exquisita Uhler, A. sigillata Drake and Bruner, A. spinicosta Van Duzee, and A. thaumana Drake and Cobben. The other 8 species are transferred to Carvalhotingis and reduced to 5 with the following new com- binations and new synonymies: C. comitis Drake, C. hollandi Drake (= comentis Drake, = denieri Monte, = rustica plana Drake, = rustica rustica Monte), C. nexa Drake, C. tumida Drake, and C. visenda Drake and Hambleton. Keys to the 2 genera and their species are included. Key Words: mies, key In honor of the memory of a long time friend and colleague— José Candido de Melo Carvalho—and in recognition of his many fine contributions to the study of biology, the following new genus, Carvalhotingis, is named after him. My attempts to divide the currently cat- aloged lace bug tribe Tingini into groups of genera based on external characters re- vealed that some of the accepted genera are actually composite, encompassing more than one genus. The present paper deals with one of them by dividing Acanthocheila Stal into two genera. The genus Acanthocheila was cataloged by Drake and Ruhoff (1965) with sixteen species and one “‘variety”’’; the distributions given below are mostly from that catalog, but include subsequently encountered country records indicated by an asterisk. Heteroptera, Tingidae, New World, new genus, new species, new synony- Until now Acanthocheila was recognized among American tingid genera by the com- bination of the uniseriate paranotum whose free margin bears a row of long (almost as long as or longer than width of an eye), stout spines, plus the costal margins being parallel for a short distance from the base and thence widened. However, my studies of other structures showed this definition included two genera: the second is here described as a new genus Carvalhotingis. Initially this separation was suggested by the presence or absence of oc- cipital spines (Figs. 1, 2); then additional contrasting characters were found. The two genera show different structure of the sub- antennal plate on the side of the head: in Carvalhotingis (Fig. 4) that plate is a simple, narrow strip passing vertically next to the eye and not approaching the side of the head: 332 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. in Carvalhotingis (Fig. 4) that plate is a sim- ple, narrow strip passing vertically next to the eye and not approaching the margin of the clypeus, while in Acanthocheila (Fig. 3) that plate has its anterior margin convexly expanded forward to almost or quite reach the margin of the clypeus. Modifications of the bucculae further support the separation: all species of Acanthocheila have the ante- rior ends of the bucculae incurved anterior to the clypeus and, except in A. sigillata Drake and Bondar, in contact with each oth- er beyond the clypeus (Fig. 3). In A. sigillata the anterior ends of the bucculae are dis- tinctly incurved anterior to the clypeus but Acanthocheila armigera (Stal). Natural length 3.7 mm. they do not meet; in contrast, the bucculae in Carvalhotingis are parallel and do not reach the apex of the clypeus (Fig. 4). In both genera there were serious prob- lems in defining and separating certain of the described species—hence, it was nec- essary to synonymize several of them. These two genera may be conveniently separated by the following couplet: Head with prominent occipital spines. Bucculae strongly incurved beyond apex of clypeus ...... Ne eR ME eine Seamer Ts Acanthocheila Stal Head without occipital spines (Caution: the long hairs may sometimes be matted in groups and look VOLUME 97, NUMBER 2 333 Fig. 2. Carvalhotingis tumida (Drake). Natural length 4.0 mm. like spines). Bucculae parallel, not incurved an- LERIORLV SF eae erat cal: Carvalhotingis, new genus Genus Acanthocheila Stal (Figs. 1, 3, 5) Monanthia (Acanthocheila) Stal, 1858: 61. Type-species:— Monanthia (Monanthia) Stal, designated by Van Duzee, 1916: 26. Acanthochila [{sic].—Stal, 1872: 127. Diagnosis. — As defined and keyed above. Characters.— Length, 2.4-5.0 mm. Dor- sal surface with few fine hairs or none. Ma- cropterous; hemelytral axes subparallel to diverging, apices moderately to strongly separated. Head.—Short, with 2-3 cephalic spines: occipital spines elongate, each reaching at least to base of antennal tubercle; with or without anteromedian spine or tubercle; su- praclypeals always absent. Eyes a little less to slightly more than half as wide as inter- ocular space. Bucculae anteriorly surpass- ing, incurved, and, except in A. sigillata, contiguous beyond apex of clypeus. Labium nearly or quite reaching meso-metasternal suture. Antennal segment I three-fourths to subequal to interocular width, almost twice 334 Figs. 3, 4. anterior to clypeus, and convex anterior margin of subantennal plate. 4. Carvalhotingis tumida. Anterior view of head showing widely separated bucculae, and straight anterior margin of subantennal plate. as long as II, III thinnest. Subantennal plate convex anteriorly, nearly or quite reaching margin of clypeus (Fig. 3). Pronotum.— Without cysts; median ca- rinae low, reaching transverse anterior mar- gin of pronotum, virtually without cells. Lateral carinae absent or represented by weak to vague calloused lines on posterior projection. Paranotum narrow to moder- ately broad, uniseriate, with marginal row of 5-10 long, stout, tapering spines. Poste- rior pronotal projection extending to basal third or midlength of discoidal area, acutely triangular, converging to a broadly truncat- ed or distinctly emarginate apex. Hemelytron.—Except for oblique basal part of costal area, nearly flat. Discoidal area, except at apex, distinctly separated from su- tural area; with 2—5 cells across widest part. Subcostal area faintly convex, horizontal, with 2-6 rows of cells, outermost row of cells sometimes narrowed and delimited by thickened veins. Costal margin convex to concave on basal fourth; margin with 0-3 small spines on basal seventh, or with a row PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 3. Acanthocheila armigera. Anterior view of head showing bucculae incurved and contiguous of distinct spines extending from base to beyond apex of discoidal area. Costal area with 1-3 rows of cells. Hypocosta uniseriate and distinct for only a short distance at base, becoming obsolete before midlength of ab- domen. Peritreme variable, elevated and trans- versely auriculate to obsolete. Metapleural flange a narrow, transverse, single row of cells. Sternal laminae present on all three sterna; on prosternum parallel, on meso- sternum distinct to very weak, widely sep- arated near base, ventral margin with or without a premedian, distinct angulation; on metasternum low but distinct, outwardly convex, apically more or less approaching each other. Abdomen basoventrally con- vex. Type species.—Monanthia (Acanthoch- eila) armigera Stal, designated by Van Du- Tee 1OV62526; Known geographic distribution.— The known range of this essentially Neotropical genus is from Argentina north through South and Central America, and the West Indies VOLUME 97, NUMBER 2 to southern Texas and the southern tip of Florida. Comments.—All species of the genus as here constituted, except A. abducta White (A. dira Drake and Hambleton known only from female holotype), show some sexual dimorphism in the subcostal area: Females have it noticeably wider than it is in the males and usually have one or more addi- tional rows of cells; females are darker and more strongly marked than males. Other variations in range of color of in- dividuals are also evident. The pronotum may vary from yellowish tan to virtually black. The hemelytra may vary from vir- tually colorless, with a few parts of veins lightly embrowned, to having most of the cells in the discoidal and subtending sub- costal areas dark brown. In the latter case the brown color extends almost or quite to the costa as a broad band across the basal third of the costal area. Within the band crossing the costal area the cells are usually much reduced in size and form more rows than are evident basal to or beyond the band. The first two antennal segments may be deep black, brown, or pale yellow; accompanying this, the apical two thirds of each femur may be decidedly darkened. The presence (Fig. 5) or absence of the premedian angulation on the lower margin of the mesosternal lamina 1s especially in- teresting when correlated with the distri- bution of the several species. All but one of those species with such an angulation are continental forms from South and Central America, including A. armigera, which ranges north into the southern United States and onto the Antilles. Those species without such an angulation appear restricted to the Bahamas and the West Indies, including 4. exquisita Uhler which was described from Key West, Florida (perhaps it was an intro- duction there), but subsequently found to inhabit the Bahamas. The exception to the above generalizations is the more recently described A. thaumana Drake and Cobben which does have such an angulation but ap- Fig. 5. ing small angulation on mesosternal carina. Acanthocheila armigera. Lateral view show- pears restricted to the Leeward Islands of St. Eustatius and St. Martin. The specimens that run to armigera in the following key to species vary in several and perplexing ways. This variability was recognized and, in part, described by Cham- pion (1897: 28) and some subsequent au- thors. Several of these variations may be present or absent in a series with identical label data or may be found in different com- binations, sometimes on the two sides of the same specimen. At this time the signif- icance of these variations is not evident. But recognition of them as simply “variations,” leads to the conclusion that A. armigera must be the senior synonym of A. nigrescens Drake and Bondar, new synonymy; further study may prove otherwise. It might be helpful to comment on some of these variations. The paranotum may ter- minate at the anterior margin of the collar or conspicuously project cephalad; its 7-11 marginal spines may be equally spaced or variously separated. Most specimens with the simple row of spines on the margin of the paranotum also have the head showing a distinct angulation or a small but evident spine anterodorsally (best seen in profile); in contrast, those specimens with the an- terior 2—3 paranotal spines crowded togeth- er usually have the anterodorsal outline of the head (in profile) smoothly decurved, but sometimes they show a distinct angulation. These combinations of differences may be evident on specimens in a series bearing the same label data, or the paranotal spine num- 336 ber and arrangement may differ on the two sides of one individual. None of these pat- terns show any association with geography. The lateral discal carinae may be absent, or may show a trace of the posterior ends as a vaguely calloused longitudinal ridge projecting anteriorly from the lateral margin of the pronotum’s posterior projection (Fig. 1). These variations do not appear to be suf- ficiently constant and contrasting to use for separating A. armigera into two or more species. LIST OF ACANTHOCHEILA SPECIES Acanthocheila abducta White Acanthocheila abducta White, 1879: 485. Acanthocheila kahavalu Kirkaldy, 1905: 216. Synonymized by Drake and Ru- hoff, 1965: 56. Known distribution: Bolivia, Brazil, Peru. Acanthocheila armigera (Stal) Monanthia (Acanthocheila) armigera Stal, 1858: 61. Monanthia (Acanthocephala) spinuligera Stal, 1858: 61. Synonymized by Monte, 1938: 128. Acanthochila (sic) armigera. —Stal, 1873: OF. Acanthocheila nigrescens Drake and Bon- dar, 1932: 88. NEw SYNONYMyY. Known distribution: Argentina, Bolivia, Brazil, Colombia, Costa Rica*, Cuba, Ecuador, El Salvador*, Guatemala, Haiti, Honduras, Jamaica, Mexico, Panama, Peru, Puerto Rico, Trini- dad, Venezuela, United States of America. Acanthocheila dira Drake and Hambleton Acanthocheila dira Drake and Hamble- ton, 1945: 359. Known distribution: Guatemala Acanthocheila exquisita Uhler Acanthocheila exquisita Uhler, 1889: 143. Known distribution: Bahamas. United States of America. Acanthocheila sigillata Drake and Bruner PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Acanthocheila sigillata Drake and Bru- ner, 1924: 147. Known distribution: Cuba. Acanthocheila spinicosta Van Duzee Acanthocheila spinicosta Van Duzee, 1907: 20. Known distribution: Dominican Repub- lic, Haiti, Jamaica, Puerto Rico, St. Thomas (Virgin Islands). Acanthocheila thaumana Drake and Cob- ben Acanthocheila thaumanni Drake and Cobben, 1960: 71. Known distribution: St. Eustatius, St. Martin (Virgin Islands). KEY TO SPECIES OF ACANTHOCHEILA 1. Costa with a series of strong spines reaching to or beyond apex of discoidal area ........... 2 — Costal area with no spines or with 1-3 confined to‘basal:seventh iar os. erie a ee 4 2. Midline of head with tubercle or a strong, pro- jecting, tapering spine above base of clypeus 3 — Midline of head with no tubercle or spine exquisita 3. Cells of discoidal and subcostal areas subo- paque, rounded, abruptly much smaller than cells on apical half of hemelytron. Paranotum narrow, its cells elongate, no wider than outer limiting vein sigillata - Cells of discoidal areas hyaline, similar to or only slightly smaller than cells in apical half of hemelytron. Paranotum wide, its cells rectan- gular to transverse, larger ones about 3x as wide as outer limiting vein ........... spinicosta 4. Midline of head with a very long (almost as long as antennal segment I), tapering, subver- tical spine between eyes. Paranotum at level of calli vertically reflexed, bringing marginal spines to a vertical or inflexed position — Midline of head without a long spine, some- times with a short, conical tubercle. Paranotum and marginal spines not reflexed, directed obliquely outward’ - 4. eee 5 5. Basal seventh of costal margin with 1-3 short, obliqueispinesi 45-4. eee eee thaumana — Costal margin with no spines .............. 6 6. Discoidal area open apically, not closed by a slightly elevated vein connecting the parallel inner and outer limiting vein. All antennal seg- ments, occipital spines, legs (except sometimes tarsi), veins of paranotum and basal two-thirds or more of costal vein wholly dull black abducta dira VOLUME 97, NUMBER 2 — Discoidal area closed apically by a noticeably elevated cross vein connecting inner and outer limiting veins and forming acute angle with latter in a more-or-less prominent pale spot. Antennal segment III, occipital spines, legs, and veins of paranotum and costa in noticeable part vellowatophyaline wena ese me cera ee armigera Genus Carvalhotingis, NEw GENUS (Figs. 2, 4) Diagnosis. — As defined and keyed above. Characters. — Length 2.8-3.8 mm. Dorsal surface and appendages with many long, pale, erect or recumbent hairs. Hemelytral axes virtually parallel, apices slightly sepa- rated. Head.—Short, without cephalic spines (often with matted groups of decurved hairs resembling spines). Eye one-third to one- half as wide as interocular width. Bucculae parallel, not attaining apex of clypeus. La- bium almost reaching or surpassing meso- metasternal suture. Antenna slender, seg- ment I two-thirds to fully as long as inter- ocular width, about twice as long as II. Sub- antennal plate with anterior margin vertical, not approaching margin of clypeus (Fig. 4). Pronotum.—Anteromedian cyst, when present, confined to area anterior to calli. Median carina simple or slightly elevated with a single row of small, obscure cells. Lateral carinae absent or only vaguely in- dicated on posterior projection. Paranotum no wider than an eye, narrowing posteriorly, uniseriate, lateral margin with a single row of 7-10 long, stout, tapering spines. Poste- rior projection of pronotum reaching basal third or midlength of discoidal area, acutely triangular, apex sharp or roundly truncated. Hemelytron.— With or without a discal elevation. Discoidal area reaching nearly or quite to midlength of hemelytron, 2-4 cells wide, inner limiting vein scarcely or not at all elevated. Subcostal area distinctly oblique, 3-4 cells wide, outermost row of cells punctiform and set off by thickened vein. Costal margins concavely constricted along basal twelfth, thence gently convex: marginal row of long, tapering spines and 337 interspersed hairs extending well beyond apex of discoidal area. Costal area (beyond basal constriction) with 1-2 rows of cells along discoidal area, 2 rows beyond. Hy- pocosta uniseriate near base, becoming ev- anescent near midlength of abdomen. Peritreme weakly to distinctly elevated. Metapleural flange very narrowly expanded, with or without a row of cells. Sternal lam- inae present on all 3 sterna, low: on meso- sternum close together near base, thence well separated and parallel or converging; on metasternum broadly to narrowly cordate, apices variously separated. Abdomen con- vex basoventrally. Type species.—Acanthocheila tumida Drake, here designated. Known geographic range.—Central and South America from Mexico to Argentina. Comments.—This genus is named in honor of Dr. José C. M. Carvalho, of Rio de Janeiro, Brazil. Examination of the five cataloged taxa that run to the first half of couplet 4 in the key below [comentis Drake, denier Monte, hollandi Drake, rustica rustica Monte, and rustica plana Drake] failed to find constant fundamental characters for their separation, and this coupled with the variation exhib- ited in a series of 28 specimens labeled “Vi- cosa, Brazil, Minas Geraes, 23-5-34, E. H. Hambleton” associated with a code number **111” leaves no alternative to synonymyz- ing all of them. In that series of 28 speci- mens, the basal fourth of the expanded cos- tal area may be wholly uniseriate, may have one to several extra interspersed individual cells on one side (as in holotype of Acan- thocheila rustica plana) or on both sides or a second partial row; occasionally an extra cell may be found in the costal area near the apical fourth of the discoidal area. List OF CARVALHOTINGIS Species Carvalhotingis comitis (Drake). NEw COMBINATION. Acanthocheila comitis Drake, 1948: 23. Known distribution: Mexico, Guatemala. 338 Carvalhotingis hollandi (Drake). NEw COMBINATION. Acanthocheila hollandi Drake, 1935: 16. Acanthocheila denieri Monte, 1940: 287. NEW SYNONYMY. Acanthocheila rustica Monte, 1942; 91. NEw SYNONYMY. Acanthocheila comentis Drake, 1953: 13. New SYNONYMY. Acanthocheila rustica plana Drake, 1953: 13. NEw SYNONYMY. Known distribution: Argentina, Brazil, Paraguay. Carvalhotingis nexa (Drake). NEw COMBIN- ATION. Acanthocheila nexa Drake, 1936: 701. Known distribution: Argentina, Brazil. Carvalhotingis tumida (Drake). Nrew COMBINATION. Acanthocheila tumida Drake, 1924: 94. Known distribution: Bolivia, Brazil. Carvalhotingis visenda (Drake and Ham- bleton). NEw CoMBINATION. Acanthocheila visenda Drake and Ham- bleton, 1934: 442. Known distribution: Argentina, Brazil, Peru. KEY TO CARVALHOTINGIS SPECIES (ignore outer row of punctiform cells in subcosta) 1. Discal elevation of hemelytron strongly inflat- ed, inflation laterally rising almost or quite ver- tically from inner limiting vein of costal area. Collar medially with distinctly elevated, in- flated cyst usually as high as or higher than interhumeral convexity — Discal elevation of hemelytron not or scarcely inflated, laterally rising obliquely from inner limiting vein of costal area. Collar sometimes visibly tectate but not inflated, distinctly lower than interhumeral convexity 2. Costal area in basal third or more with a single row of large, rectangular cells ............. nexa — Costal area with 2 irregular rows of cells to base PCa oe ne ae Ree ee aa e NE Sere visenda 3. Subcostal area along basal half or more of dis- coidal area coriaceous with punctiform cells, thence (ignore outer row of punctiform cells) abruptly with large hyaline cells ........ tumida PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON -— Subcostal area along basal third or fourth of discoidal area with punctiform cells, thence (ignore outer row of punctiform cells), on apical two-thirds, cells gradually enlarging ........ 4 4. Dorsal outline of discal elevation, in lateral VIEW allowsCONVeXItys “ne eee eee hollandi - Dorsal outline of discal elevation, in lateral view, a blackened, abrupt, erect, tabular pro- jection: <2 Av, cate e ese oR 8 eae comitis ACKNOWLEDGMENTS The author is grateful for the special sup- port he received from the National Science Foundation in the form of a grant (NSF- GB-791) to study Tingidae and other Het- eroptera in European Museums in 1968; this grant plus the kind cooperation of Dr. L. Brundin made possible study of Stal’s types in the Riksmuseet, Stockholm. For prepa- ration of this paper, special acknowledg- ment goes to Y. T. Sohn (Illustrator, Dept. Entomol., Smithsonian Inst., Washington, D.C.) for the excellent illustrations; and to J. E. McPherson (Dept. Entomol., Southern Illinois University, Carbondale), R. M. Smith (Texas A&M University, College Sta- tion), P. J. Spangler (Dept. Entomol., Smithsonian Inst., Washington, D.C.), and A. G. Wheeler, Jr. (Pennsylvania Depart- ment of Agriculture, Harrisburg) for helpful reviews of the manuscript. LITERATURE CITED Champion, G. C. 1897. Insecta: Rhynchota (Hemip- tera-Heteroptera). Jn Godman and Selvin. Biol- ogia Centrali-Americana. London. 2:1-32. Drake, C. J. 1924. A new species of Acanthocheila from Bolivia (Hemiptera-Tingitidae). Bulletin of the Brooklyn Entomological Society 19: 94. 1935. American Tingitoidea (Hemiptera) in the Natural History Museum of Vienna. Sondar- Abdruck aus ““Konowia”’ 14: 9-20. . 1936. Some Tingitoidea from Argentina (He- miptera). Travaux de l'Institut Zoologique de l’A- cadémie des Sciences de l?URSS 3: 699-701. . 1948. New American Tingidae (Hemiptera). Boletin de Entomologia Venezolana 7: 20-25. . 1953. New Neogaean Tingidae (Hemiptera). Great Basin Naturalist 13: 13-16. Drake, C. J. and G. Bondar. 1932. Concerning Bra- zilian Tingitidae, Hemiptera. Boletim do Museu Nacional, Rio de Janeiro 8: 87-96, | plate. VOLUME 97, NUMBER 2 Drake, C. J.and S.C. Bruner. 1924. Concerning some Tingitidae occurring in the West Indies (Hemip.). Memorias de la Sociadad Cubana de Historia Nat- ural ‘“‘Felipe Poey” 6: 1-13. Drake, C. J. and R. H. Cobben. 1960. The Heter- optera of the Netherlands Antilles—V, Tingidae (lace bugs). Studies on the Fauna of Curacao and Other Caribbean Islands 10:(54): 67-97. Drake, C. J. and E. J. Hambleton. 1934. Brazilian Tingitidae (Hemiptera). Part I. Revista de Ento- mologia, Rio de Janeiro 4: 435-451. 1945. Concerning Neotropical Tingitidae (Hemiptera). Journal of the Washington Academy of Sciences 35: 356-367. Drake, C. J. and F. A. Ruhoff. 1965. Lacebugs of the world, a catalog (Hemiptera: Tingidae). United States National Museum Bulletin 243: i-vii, 1- 634, frontispiece, plates 1-56. Kirkaldy, G. W. 1905. Quelques tingides nouveaux ou peu connus [Hem.]. Bulletin Société Entomo- logique de France 15: 216-217. Monte, O. 1938. Tingitideos Neotropicos. Boletim Biologico, new series 3: 127-132. 1940. Tingitideos novos ou pouco conheci- dos da Fauna Americana. Arquivos de Zoologia do Estado de Sao Paulo 11: 283-300. . 1942. Apontamentos sobre Tingitideos (He- miptera) Americanos, especialmente do Brasil. 339 Arquivos do Instituto Biologico Sao Paulo, 13: 91-98. Stal, C. 1858. Bidrag til Rio Janeiro-Traktens He- miptera-fauna. I. Kongliga Svenska Vetenskaps- Akademiens Handlingar 2(7): 1-84. . 1872-1873. Enumeratio Hemipterorum: Bi- drag till en foreteckning ofver alla hittils kanda Hemiptera, jemte systematiscka meddelanden. Kongliga Svenska Vetenskaps-Akademiens Han- dlingar, 1872, Part 2, 10(4): 1-159; 1873, Part, 11(2): 1-163. Uhler, P. R. 1889. Observations upon the Heterop- tera collected in southern Florida by Mr. E. A. Schwarz. Proceedings of the Entomological Soci- ety of Washington 1: 142-144. Van Duzee, E. P. 1907. Notes on some Jamaican Hemiptera, a report on a collection of Hemiptera made on the island of Jamaica in the spring of 1906. Bulletin of the Buffalo Society of Natural Sciences 8(5): 3-79. 1916. Check List of the Hemiptera (Except- ing the Aphididae, Aleurodidae and Coccidae) of America North of Mexico. New York Entomo- logical Society, New York. xi + 2 + 111 pp. White, F. B. 1879. Descriptions of New Hemiptera (I). Journal of the Linnaean Society of London, Zoology 14: 482-489. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 340-345 PROBOSCIDOTYLUS CARVALHOI, A NEW GENUS AND SPECIES GF SEXUALLY DIMORPHIC PLANT BUG FROM MEXICO (HETEROPTERA: MIRIDAE: ORTHOTYLINAE) THOMAS J. HENRY Systematic Entomology Laboratory, Plant Sciences Institute, Agricultural Research Ser- vice, United States Department of Agriculture, % National Museum of Natural History, MRC-168, Washington, D.C. 20560. Abstract.—The new orthotyline genus and species Proboscidotylus carvalhoi are de- scribed from specimens collected in Veracruz, Mexico. This new mirid, with the male possessing an elongate tubercle at the apex of the tylus, represents one of the most unusual cases of sexual dimorphism in the subfamily Orthotylinae. An adult habitus drawing, illustrations of male genitalia, and scanning electron micrographs of selected structures are furnished to aid in recognition, and relationships are discussed. Key Words: phism, Mexico, Veracruz The subfamily Orthotylinae is one of the largest and most poorly studied groups in the Miridae, especially in the Neotropics. Five tribes are recognized (Austromirini, Halticini, Ceratocapsini, Nichomachini, and Orthotylini), but this level of the classifi- cation is unstable (Schuh 1974, Henry 1994) and a worldwide analysis is needed to es- tablish the monophyly of these orthotyline tribes. Discovery of a bizarre new genus and species fitting tato Schuh’s (1974) Zanchius group (Orthotylini) adds new and corrob- orative character information that should be useful to future phylogenetic investiga- tions of the subfamily. Herein, I describe the new genus and new species Proboscidotylus carvalhoi, provide an adult habitus drawing, illustrations of male genitalia, and SEM micrographs of pertinent structures, and discuss the rela- tionship of this peculiar new genus to other orthotyline genera. This paper and the new species described within it are dedicated to the memory of Insecta, Heteroptera, Orthotylinae, new genus, new species, sexual dimor- Dr. José Candido de Melo Carvalho, who published on the Miridae for more than 50 years. Landmarks in an outstanding career were his 1113-page world catalog of the Muiridae (1957-1960), keys to the mirid gen- era of the world (1955), and many papers in which he described more than 385 genera and 2080 species (Carvalho and Froeschner 1987, 1990, in press). It is hard to imagine that José’s voluminous contributions to our knowledge of this large and diverse family will ever be surpassed. Proboscidotylus Henry, NEw GENUS Type species. — Proboscidotylus carvalhoi Henry, new species. Diagnosis.—This genus can be recog- nized by the strongly swollen frons (Figs. 2, 4), prominent tylus, and the scattered black scalelike setae on the hemelytra (Fig. 6). Most distinctive, however, are the peculiar autapomorphic structures found only in males. The elongate, apically setose, tuber- cle arising from the apex of the tylus (Fig. VOLUME 97, NUMBER 2 341 Fig. 1. Dorsal habitus of Proboscidotylus carvalhoi, 2, 3) and the tuberclelike process bearing a field of stout bristlelike setae on the genital capsule (Figs. 8, 9) are unique in the sub- family Orthotylinae and support the mono- phyly of Proboscidotylus. Description. —Orthotylinae: Orthotylini. Small, delicate (Fig. 1), length 2.64-3.04 mm. Head (Figs. 2-4) broader than long; male. interocular area in female flattened, in male weakly depressed; frons abruptly and strongly swollen in both sexes but more so in females (Fig. 4), then flattened to meet base of pronounced, broadly rounded tylus; male with apex of tylus extended into an elongate, apically rounded tubercle, having long dorsally directed setae distally (Figs. 2, 342 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-9. Proboscidotylus carvalhoi. 2, head, lateral aspect of male (150). 3, lateral aspect of male tylus (260 x). 4, head, lateral aspect of female (151 x). 5. ostiolar area (359 x). 6, scalelike setae of hemelytra (1610 x). 7, claw (1040 x). 8, male genital capsule, caudal aspect (426 x). 9, male genital capsule, lateral aspect (447 x). VOLUME 97, NUMBER 2 343 Figs. 10-12. Male genitalia. 10, left paramere. 11, right paramere. 12, vesica. 3), tubercle absent in female; eye promi- nent, oblong oval, strongly faceted, occu- pying all but lowermost edge of head in lat- eral aspect, posterior margin nearly touch- ing anterior margin of pronotum, set with scattered short setae, posterior lateral mar- gin with a row of stout, erect setae. Rostrum extending nearly to bases of metacoxae. An- tenna slender; segment I shortest, stoutest; segment II slender, more than 2 times length of segment III in males, less than 2 times length of segment III in females; segment III longer than segment IV. Pronotum tra- peziform, collar absent, much broader than long, with a transverse impression through middle separating calli from disc; posterior width much greater than anterior width; an- terior angles rounded; lateral margins straight and angled outward posteriorly, margins weakly carinate; pleural area flat- tened, subquadrate; posterior margin deep- ly sinuate; calli weakly swollen. Scutellum flattened, equilateral; mesoscutum wide, about two thirds as long as scutellum, broadly exposed. Hemelytron hyaline, weakly sinuate laterally through middle in males, more nearly straight or weakly rounded in females; set with relatively long, semierect simple setae, sparsely intermixed with flattened, apically acute, scalelike, black setae (Fig. 6) on clavus, corium, and cuneus; cuneus about as long as wide; membrane hyaline, broadly rounded apically, with two closed cells. Ventral surface weakly shining, with scattered simple setae. Ostiolar area (Fig. 5) small, auricle roughly triangular, margins bordered with modified evapora- tive or mycoid surface. Legs slender; tibial spines short, indistinct; tarsi three seg- mented; claws (Fig. 7) slender, apically re- curved, parempodia broad and convergent. Male genital capsule (Figs. 8, 9) small, api- cally pointed, with a broad process or tu- bercle arising below basal margin of aper- ture bearing a field of stout, apically blunt, bristlelike (and likely glandular) setae. Left paramere (Fig. 10) elongate, extended trans- versely into an apically acute, slender arm; right paramere (Fig. 11) oval, apically acute; vesica (Fig. 12) membranous, bearing a sin- gle, slender, apically acute and curved spic- ulum, secondary gonopore absent or indis- tinct. Remarks.—Relationships of the New World Orthotylinae are insufficiently known to fully place Proboscidotylus in a phylo- genetic context. This genus belongs in the tribe Orthotylini as it is now defined and can be assigned to Schuh’s (1974) Zanchius group, members of which have a general delicate body structure, flattened appear- ance, hyaline hemelytra, and a vesica lack- 344 ing spiculi (although some taxa appear to have at least one spiculum, including Pro- boscidotylus carvalhoi). 1 add to these di- agnostic characters a broadly exposed mesoscutum (resulting in part from a deeply sinuate posterior pronotal margin) and a blunt, apically extended process on the male genital capsule below the caudal edge of the aperture. This combination of characters is unique in the Orthotylinae and suggests that members of this group warrant placement in a separate tribe. Such an action, however, needs to await a comprehensive study of the subfamily. Neotropical genera that Schuh (1974) placed in the Zanchius group are Brasiliom- iris Carvalho, Hyalochloria Reuter, Itacoris Carvalho, Jobertus Distant, and Paraproba Distant. To this list, in addition to Probos- cidotylus, 1 add Saileria Hsiao and Diaphn- ocoris Kelton. In Knight’s (1968) key to western U.S. genera, Proboscidotylus will run to either Squamocoris Knight or Melanotrichus Reu- ter based on head and eye characteristics and the presence of scalelike setae on the hemelytra. In Carvalho (1955), it will key to Parthenicus Reuter, if the head is said to lack a well-defined posterior margin, or, if the posterior margin is considered well de- fined, to J/nacora Reuter having black, scalelike, dorsal setae and antennal segment I subequal to the width of the vertex. Only a few New World orthotyline genera bear black, scalelike hemelytral setae (e.g. Brooksetta Kelton, I/nacora, Ilnacorella Knight, Macrotyloides Van Duzee, and Par- thenicus). Proboscidotylus, however, ap- pears to have little to do with these taxa based on overall morphology and male gen- italia. Strong sexual dimorphism is uncommon in the Orthotylinae. Although hemelytral brachyptery or microptery is common in females of some taxa (e.g. Ceratocapsus Reuter, sensu lato), other structural differ- ences between the sexes are rare. The pri- mary exception occurs in the Neotropical PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON genus Hyalochloria Reuter, in which males always have one or two stout, often de- curved spines on the basal '4 of the second antennal segment (Henry 1978). Probosci- dotylus provides another striking example of sexual dimorphism, with the male bear- ing an elongate tylar process and a blunt genital tubercle covered by a field of stout apically blunt and, perhaps, glandular setae. Proboscidotylus carvalhoi Henry, NEW SPECIES Figs. 1-12 Diagnosis.— This species is recognized by the overall pale brownish-yellow colora- tion, strongly swollen frons (Figs. 2, 4), and hyaline hemelytra possessing scattered black, scalelike setae (Fig. 6), as well as by the generic characters discussed above. Description. — Male (n = 4): Length 2.68- 2.84 mm, width 1.20-1.26 mm; overall col- oration pale brownish yellow. Head: Weak- ly shining, with scattered erect setae; width 0.64-0.66 mm, vertex (interocular width) 0.24-0.28 mm, length of setigerous tubercle on tylus about 0.10 mm, apex with a ciuster of long, erect, simple setae. Rostrum: Length 0.74-0.78 mm, extending nearly to bases of metacoxae. Antenna: Segment I, length 0.28-0.30 mm; II, 1.20—1.24 mm; III, 0.56— 0.64 mm; IV, 0.34-0.40 mm. Pronotum: Length 0.28-0.30 mm, basal width 0.84—- 0.86 mm. Scutellum and mesoscutum with scattered erect and semierect simple setae. Hemelytron: Translucent pale brownish yel- low, clothed with numerous semierect sim- ple setae, intermixed with flattened, apically acute, scalelike, black setae on clavus, co- rium, and inner angle of cuneus. Ventral surface: Shiny pale brownish yellow, with scattered simple setae, especially on abdo- men. Male genitalia: Genital capsule small, tapered apically, ending in a broad setig- erous process or tubercle; left paramere (Fig. 10), right paramere (Fig. 11), and vesica (Fig. 12) as described under genus. Female (n = 8): Length 2.64-3.04 mm, VOLUME 97, NUMBER 2 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; III, 0.74-0.78 mm; IV, 0.40-0.46 mm. Pro- notum: Length 0.28-0.32 mm, basal width 0.84-0.88 mm. Very similar to male in overall shape and coloration, differing in lacking the elongate, setigerous tubercle at the apex of the frons and having a more strongly swollen frons (Fig. 4). Host.— Unknown. Etymology.—The specific epithet of this unusual species is named in memory of my good friend and colleague Dr. José C. M. Carvalho. Type data.—Holotype 3, Mexico: Vera- cruz, Los Tuxlas Estac. Biol., N of Cate- maco, 16-19 Sept. 1989, E. Barrera, T. J. Henry, and I. M. Kerzhner colls., taken at incandescent porch light (USNM [U.:S. Na- tional Museum of Natural History, Wash- ington, D.C.]). Paratypes: 4 46, 8 92, same data as for holotype (UNAM [Universidad Nacional AutOnoma de México, México, D.F.], USNM). Other specimens examined.—1 4, 1 2, same data as for holotype, prepared for scanning electron microscope investiga- tions; 1 6, with abdomen missing (USNM). ACKNOWLEDGMENTS I thank H. Brailovsky (UNAM) for his hospitality and invitation to visit Mexico in 1989, and E. Barrera (UNAM) for his com- panionship and guidance (with I. M. Ker- zhner) to many interesting collecting sites in the Mexican states of Oaxaca, Puebla, and Veracruz. Alexander Konstantinov (Washington, D.C.) illustrated the adult habitus, and R. C. Froeschner (USNM), Ronald W. Hodges (Systematic Entomolo- 345 gy Laboratory (SEL), PSI, ARS, USDA, % USNM), James Pakaluk (SEL, % USNM), and A. G. Wheeler, Jr. (Pennsylvania De- partment of Agriculture, Harrisburg) kindly reviewed the manuscript. LITERATURE CITED Carvalho, J.C. M. 1955. Keys to the genera of Mir- idae of the world (Hemiptera). Boletim do Museu Paraense Emilio Goeldi XI(II): 1-151. 1957-1960. Catalogue of the Miridae of the world. Arquivos do Museu Nacional, Rio de Ja- neiro. Part I. Cylapinae, Deraeocorinae, Bryocor- inae 44(1): 1-158 (1957); Part II. Phylinae 45(2): 1-216 (1958); Part III. Orthotylinae 47(3): 1-163 (1958); Part IV. Mirinae 48(4): 1-384 (1959); Part V. Bibliography & Index 51(5): 1-194 (1960). Carvalho, J.C. M. and R. C. Froeschner. 1987. Tax- onomic names proposed in the insect order Het- eroptera by José Candido de Melo Carvalho from 1943 to January 1985, with type depositories. Journal of the New York Entomological Society 95: 121-224. . 1990. Taxonomic names proposed in the in- sect order Heteroptera by José Candido de Melo Carvalho from January 1985 to January 1989, with type depositories. Journal of the New York En- tomological Society 98: 310-346. . Taxonomic names proposed in the insect or- der Heteroptera by José Candido de Melo Car- valho from January 1989 to January 1993. Journal of the New York Entomological Society (In press). Henry, T. J. 1978. Review of the Neotropical genus Hyalochloria, with descriptions of ten new species (Hemiptera: Miridae). Transactions of the Amer- ican Entomological Society 104: 69-90. 1994. Revision of the genus Schaffneria Knight (Heteroptera: Miridae). Proceedings of the Entomological Society of Washington 96: 701-712. Knight, H. H. 1968. Taxonomic review: Miridae of the Nevada Test Site and the western United States. Brigham Young University Science Bulletin 9(3): 1-282. Schuh, R. T. 1974. The Orthotylinae and Phylinae (Hemiptera: Miridae) of South Africa with phy- logenetic analysis of the anti-mimetic tribes of the two subfamilies for the world. Entomologica Americana 47: 1-322. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 346-349 REUTEROSCOPUS CARVALHOI N. SP., A NEW FOSSIL PLANT BUG (HETEROPTERA: MIRIDAE: PHYLINAE) J. Maldonado Capriles and George O. Poinar, Jr. (JMC) Department of Crop Protection, College of Agricultural Sciences, Mayagiiez Campus of the University of Puerto Rico, Mayagitiez, Puerto Rico 00681; mailing address: Urb. Aponte 6 I 1, Cayey, Puerto Rico 00736. (GOP) Division of Entomology and Plant and Soil Microbiology, 201 Wellman Hall, University of California, Berkeley, California 94720. Abstract. —The fossil mirid Reuteroscopus carvalhoi n. sp. is described from Dominican amber and compared to extant species of the genus. Key Words: The mirid genus Reuteroscopus Kirkaldy, 1905, occurs in the Greater and Lesser An- tilles, northern coast of South America (specimens from the last two places in JMC’s collection), Central America, and in North America as far north as Canada. It includes about 40 species that live on weeds and a few species of trees. The fossil species described in this paper is very similar to many extant species, es- pecially in the typical color pattern of the hemelytra. In the description we use the same colors used by Knight (1965) to fa- cilitate comparison. The only morphological difference we no- ticed between this fossil and modern species of Reuteroscopus is the shape of the eyes. In the dorsal view, instead of being hemispher- ical they are somewhat elliptical. As in mod- ern species, the eyes extend backward for a short distance along the lateral margins of the pronotum, but in this species they ex- tend a little more forward, thus, occupying a slightly larger area of the vertex. In lateral view (Fig. 2), the anterior margin is concave to receive the insertion of the antenna. Ex- tant species have vertical, oval eyes, some of the them with the posterior margin shal- Miridae, Reuteroscopus carvalhoi n. sp., fossil, Dominican Amber lowly indented, and the antennae are in- serted close to or are slightly separated from the anterior margin. The eyes of specimens from the Antilles, northern South America, Central America and southern United States were examined. There are few mirid fossil records. As cit- ed by Carvalho (1959), Scudder (1890) de- scribed 14 species as mirids from the Flor- issant (Cenozoic, Oligocene). Carvalho (1959) indicated that Scudder’s taxa look more like anthocorids or at least cannot be assigned to annectant genera. Scudder placed his species in Aporema Scudder, Clostero- coris Uhler, Fuscus Distant, Poecilocaspus Reuter, Capsus Fabricius, and Hadronema Uhler. Despite such assignments the char- acters he used are insufficient to place them in subfamilies and tribes. Also cited by Car- valho, Germar and Berent (1856) described 13 species, mostly from Prussian amber, placing all of them in what they loosely called Phytocoris. Our new species is in a piece of amber originating from mines in Cordillera Sep- tentrional between Puerto Plata and Santi- ago in northern Dominican Republic. These mines are in the Altamira facies of the El VOLUME 97, NUMBER 2 347 Figs. 1-4. Reuteroscopus carvalhoi Maldonado and Poinar, n. sp., holotype, female. 1, habitus, dorsal view. 2, head, lateral view. 3, hind femur, lateral external view. 4, abdomen, lateral view. Scale lines represent 1.0 mm. Mamey Formation and the estimated age ranges between 25 and 40 million years (up- per Oligocene-upper Eocene) (Lambert et al. 1985). Reuteroscopus carvalhoi Maldonado and Poinar, NEw SPECIES Figs. 1-4 Female. Overall coloration of body prob- ably pale green in life, hemelytra, hind fe- mur and abdomen conspicuously orna- mented with black. Head, pronotum, lateral angles of mesoscutum, base and apex of cla- vus, most of corium including its apex (mes- ad to cuneus), embolium, and cuneus pale green. Excluding membrane, remaining parts of fore wings black (Fig. 1); mem- brane— basal half apparently black, most of inner cell blackish, outer cell grayish, cau- dad of cells to apex of membrane grayish. Antennal segments: I black basally, apical half pale because of its silvery pilosity; II fuscous, basally and apically black; III and IV uniformly dark fuscous, slightly darker than II. Rostrum probably fuscous, at least not pale. Pronotum laterally, most of me- sopleura except for pale area around base of mesocoxa, and metapleura black. Osteo- lar peritreme pale greenish. Abdominal sterna pale green along upper margin; first three segments black above the pale green below, next three with a small pale green basal area, the black areas almost contigu- ous with the pale green ovipositor (Fig. 3). Legs: anterior—coxa mostly pale green, in- ner and outer surfaces with a small black central area; trochanter pale green; femur pale green, ventral edge thinly lined with black; tibia with basal 73 blackish, apically pale fuscous; tarsi pale fuscous; middle— coxa and trochanter green, fuscous apically, tibia fuscous, slightly lighter than last two 348 antennal segments; hind—coxa and tro- chanter pale green; femur black, basal and apical fourths pale green, the latter with an incomplete, irregular ring and two or three small black spots (Fig. 4). Pilosity. — Head, pronotum, light areas of corium, embolium, cuneus, and costal mar- gin with decumbent fine pilosity; dark areas of fore wing with yellow or greenish fine, decumbent pilosity; the dark areas seem to have some pale green, scalelike, adpressed setae. Extant species have only one kind of setae on the dark areas of the hemelytra, so, these “‘scalelike”’ setae are either an illusion due to refraction, or at least a second type of setae, not necessarily scalelike. Spinelike setae of hind tibia not arising from black spots. Measurements (in mm).—Head length 0.13, width across eyes 0.50, eyes moder- ately large, ellipsoidal, lower margin far from ventral margin of head, length 0.31; pro- notum length 0.31, humeral width 0.81; mesoscutum exposed; scutellum width 0.50, length 0.37. Antennal segments: I, 0.13; II, 0.56; IIT, 0.31; IV, 0.50; all short decumbent setose. Rostrum 1.56, reaching IV sternum, i.e., surpassing base of ovipositor. Legs— fore: femur 0.53, tibia 0.53; middle: femur 0.62, tibia 0.68; hind: femur 0.76, tibia 1.15. Tarsi as in Phylinae, linear, about % length of corresponding tibia, parempodia appar- ently small, not visible, claws elongate, slen- der. Hemelytra well surpassing abdominal apex, length 1.87, greatest width 1.00. Ab- domen length 1.1. Total length of body 2.31. Holotype female, from Dominican Am- ber, DOMINICAN REPUBLIC; the amber piece containing the specimen is designated HE-4-47 and is deposited in the Poinar col- lection of Dominican amber maintained at the University of California, Berkeley. We take pleasure in naming the species after J. C. M. Carvalho, the father of modern mir- idology. Discussion.—The morphological differ- ences between the eyes of this fossil and modern species do not seem to warrant es- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tablishing a new genus. Considering that R. carvalhoi is described from a female 2.31 mm long and that males are usually shorter than females evidently this is the smallest species in the genus. The best way to define species in Reuteroscopus is by means of the shape of the vesica of the male genitalia. Knight (1965: 102) points out that differ- ences in coloration, whether the black of the hemelytra is sharply (the “‘ornata” group as in R. carvalhoi) or diffusely separated from the pale areas, the relative lengths of anten- nal segment II and humeral width of pro- notum, and the relative thickness of anten- nal segment II and thickness of protibia can be used to separate females. Reuteroscopus carvalhoi n. sp. runs in both Knight’s (1965) and Kelton’s (1964) key to R. aztecus Keiton after skipping characters related to male genitalia in the latter key. The females of R. aztecus are 3.78-4.06 long, with green abdominal sterna and infuscated cuneus. The females of R. femoralis Kelton, from Mexico and R. hamatus Kelton are slightly under 3.00 long, but in both the rostrum extends to the tip of hind coxae. Other species in the genus are 3.00 or longer and differ in coloration. In R. carvalhoi the humeral width is 1.4 times greater than the length of antennal segment IJ, whereas in the two species mentioned above, the hu- meral width is 1.0—1.1 greater than antennal segment II, and their abdominal sterna are yellow. LITERATURE CITED Carvalho, J. C. M. 1959. Catalogo dos mirideos do mundo. Part IV. Mirinae. Arquivos do Museu Na- cional, Rio de. Janeiro 48: 1-384. Germar, E. F. and G. C. Brendt. 1856. Die im Bern- stein befindlichen Hemipteren und Orthopteren der Vorwelt. Jn G. C. Brendt, die im Bernstein befindlichen Reste der Vorwelt gesammelt in Ver- bindung mit Mehrern, bearbeitet und herausge- geben von... G. C. Berendt. Bd. 2, Abth. 1, IV, 1-40 pp., 4 pls. Berlin. Kelton, L. A. 1964. Revision of the genus Reuter- oscopus Kirkaldy 1905 with description of eleven new species (Hemiptera: Miridae). Canadian En- tomologist 96(11): 1421-1433. VOLUME 97, NUMBER 2 349 Lambert, J. B., J. S. Fryerard, and G. O. Poinar, Jr. (Hemiptera, Miridae). Iowa State Journal of Sci- 1985. Amber from the Dominican Republic: ence 40(2): 101-120. Analysis by nuclear magnetic resonance spectros- Scudder, S. H. 1890. The tertiary insects of North copy. Archaeometry 27: 43-51. America. Report United States Geological Survey Knight, H. H. 1965. A new key to species of Reu- Territories 13: 1-464, 28 plates. teroscopus Kirk. with descriptions of new species PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 350-372 A PHYLOGENETIC REVIEW OF THE POTAMOBATES FAUNA OF COLOMBIA (HETEROPTERA: GERRIDAE), WITH DESCRIPTIONS OF THREE NEW SPECIES JOHN T. POLHEMUS AND DAN A. POLHEMUS (JTP) University of Colorado Museum, 3115 S. York St., Englewood, Colorado 80110; (DAP) Department of Natural Sciences, Bishop Museum, P.O. Box 19000-A, Honolulu, Hawaii 96817. Abstract.—Three new species of Potamobates are described from Colombia and sur- rounding regions: P. manzanoae n. sp. from the Rio Anchicaya drainage in Valle de Cauca Province; P. anchicaya n. sp. from the Rio Anchicaya area and Panama; and P. carvalhoi n. sp. from the Rio Claro in Antioquia Province, and Venezuela. These new species are compared with other previously described taxa in the genus, and a set of phylogenetically inferred intrageneric species groupings is proposed, accompanied by a key and checklist for all known species, and a species level phylogeny. Distribution maps are provided for species occurring in Colombia, and country-level distributional data are given for all other known species. Key Words: Due to its equatorial position and moun- tainous character, Colombia supports one of the richest and yet most poorly investi- gated aquatic Heteroptera biotas in South America. In the current report we review the region’s fauna of Potamobates water striders, describe three new species, provide a key and checklist to all known species, and present a preliminary phylogenetic analysis of the genus at the species level. The three new taxa described herein bring the total number of documented species in the genus to 15, which we assign to four intrageneric species groups, based primarily on apomorphic characters of the male and female abdominal terminalia. Country-lev- el distributions and complete nomenclatur- al citations for all described species are also provided, plus detailed distribution maps for all species occurring in Colombia. The genus Potamobates was described by Champion (1898), and redescribed by Drake Potamobates, Gerridae, Colombia, taxonomy, phylogeny distribution, key and Harris (1934), Kuitert (1942) and Mat- suda (1960). Members of this genus are moderately large, muscular water striders that are generally found actively skating on the midstream waters of swift, rocky streams at low to intermediate elevations. The genus is wholly Neotropical, ranging from south- ern Mexico to Peru, and appears to repre- sent the Western Hemisphere ecological equivalent of the Oriental genus Ptilomera. Distribution data and keys to species were previously published by Drake and Harris (1934) and Kuitert (1942), but these authors did not have access to all of the known spe- cies, and their omissions plus the three new species described below have necessitated the development of the revised key pre- sented herein. All measurements in the descriptions be- low are given in millimeters. CL numbers following localities in the material exam- ined sections refer to a data coding system VOLUME 97, NUMBER 2 used by the authors to cross reference spec- imens, ecological field notes, and habitat photographs. Potamobates carvalhoi, NEw SPECIES Figs. 1-6, 19 Diagnosis.—Potamobates carvalhoi oc- cupies a phylogenetic position immediately basal to the insular South American P. wil- liamsi group (see Figs. 30, 31), but is set aside in its Own monotypic intrageneric grouping because of the unusual and diag- nostic character of the abdominal termina- lia in both males and females (see subse- quent group analysis). The completely hid- den female tergite VIII is unique within the genus. Description (all measurements in milli- meters).—Apterous male: Ground color blackish, faintly shining, covered with very short dark pubescence; head except vertex along eyes and longitudinal oval black spot ahead of eyes, median longitudinal wedge on pronotum, figures on mesonotum if pres- ent (Fig. 2), propleura, mesopleura, metace- tabulae, coxae, fore trochanters, dorsum of fore femur except distally, pygophore and venter brownish yellow to leucine. Antero- lateral angles of pronotum, longitudinal stripe on lateral margins and posterolateral angles of mesonotum, lateral margins of ab- dominal tergites, and coxal cavities dorsally covered with short bright golden pubes- cence. Legs black to brown, middle and hind femora lighter beneath, antennae black. Legs set with short inconspicuous black spines. Length of head (1.22) more than four times narrowest interocular space (0.28); width of eye 0.72; eyes extending posteri- orly about % length of propleura. Rostrum short, barely reaching onto mesosternum. Pronotum short (1.05), width (1.89) about equal to head across eyes. Mesonotum long (2.33), widest across mesocoxae (2.89). Metanotum short (0.83), widest across metacoxae (2.39). Abdomen excluding gen- ital segments relatively short (2.33), genital segments long (2.05). Tergites III—-V about 351 of equal length (0.17-0.22), tergites II and VI longer (0.28), tergite VII longest (0.67); VIII long (1.44), large, slightly modified, ventrally asymmetrical (Figs. 2, 5, 6); proc- tiger modified, with long projection on left side, terminating in a “bird-head”’ shape; male terminalia as shown in Figs. 5, 6. Con- nexiva angulate caudally, not produced; raised along entire length. Antennae long, slender, segments I and IV slightly stouter than II and III; length of segments I-IV: 1.83; 0.61; 0.67; 1.00. An- terior femur stout, tapering distally; anterior tibia slightly flattened, not dilated. Mea- surements of legs as follows: Femur, tibia, tarsal 1, tarsal 2 of fore leg, 2.55: 2.61: 0.14: 0.50; of middle leg, 9.43: 6.94: 3.33: 0.94; of hind leg, 9.82: 6.60: 0.44: 0.28. Length, mean 9.68 (N = 6, min. 9.50, max. 9.86). Width, mean 2.93 (N = 6, min. 2.84, max. 3.04). Apterous female: Body more robust than male (Fig. 1), otherwise similar in general size, coloration, and other features except abdominal terminalia very differently formed. Connexiva produced posteriorly into long slender digitate lobes, usually con- vergent (Fig. 3). Abdominal ventrite VIII produced symmetrically, lateral lobes large, folded over tergite VIII which is hidden (Figs. 3, 4). Length, mean 8.84 (N = 10, min. 8.61, max. 9.14). Width, mean 3.13 (N = 10, min. 3.02, max. 3.29). Macropterous male: Similar to apterous form in most respects, except pronotum long, with posterior lobe raised and broadly margined with brownish yellow, humeri prominent. Wings light brown, reaching be- yond tip of abdomen, veins blackish brown, without pubescence, with five closed cells. Length, 11.28 (N = 1). Width, 3.02 (N = 1). Macropterous female: Unknown. Etymology.— The name carvalhoi honors José Candido de Melo Carvalho for his many 352 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Potamobates carvalhoi n. sp. 1, Female, dorsal habitus (legs omitted). 2, Male, dorsal habitus (legs omitted). 3, 4, Female terminal abdomen. 3, Dorsal view. 4, Ventral view. 5, 6, Male terminal abdomen, showing distal part of segment VII, all of VIII, IX & X partially hidden within VIII. 5, Lateral view. 6, Oblique ventral view. VOLUME 97, NUMBER 2 splendid contributions to our knowledge of Heteroptera. Ecological notes.—P. carvalhoi was taken in midstream, skating in areas of slow, smoothly flowing current. Material examined.— Holotype, apterous male and allotype, apterous female (USNM), COLOMBIA: Antioquia Prov., Rio Claro, 13 km W. of Dorodal, water temp. 25 °C, 250im. CE240352 21 viel9s9. Je Ti. dé D. A. Polhemus. Paratypes as follows (all apter- ous unless noted): COLOMBIA: Antioquia Prov.: 4 males, 5 females, 1 macropterous female, same data as holotype (JTPC, USNM); 2 males, 5 females, Quebrada La Negra, small tributary to Rio Claro, 13 km W. of Dorodal, 250 m, CL 2406, 21.vii.1989, J. T. & D. A. Polhemus (JTPC). VENE- ZUELA: Barinas Prov.: 2 males, 4 females, Barinitas, Dec. ’42, P. Anduzee (det. C. J. Drake as P. unidentatus) (JTPC, USNM). Potamobates anchicaya, NEw SPECIES Figs. 7-13, 21 Diagnosis.— Potamobates anchicaya is most closely related to P. tridentatus Esaki, but compared to the latter anchicaya may be separated by the narrower silvery pu- bescent stripe laterally on the mesonotum; the less pronounced incision between the “teeth” on the lobe of male abdominal seg- ment VIII, and in having this lobe of dif- ferent shape (Fig. 10; compare with Esaki 1926, Fig. le); by the emarginate ventral incision of male abdominal segment VIII being narrower and shallower, and the lat- erally directed process of the proctiger larg- er, longer and of different shape (Fig. 12: compare with Esaki 1926, Fig. 1d); and by female ventrite VIII extending posteriorly about the length of ventrite VI (vs. about 2 the length of ventrite VI in tridentatus), un- less strongly bent ventrad (vs. never stongly bent ventrad), and with the lateral lobes less asymmetrical, having both right and left lobes developed (vs. left lobe developed), with one or both sometimes wrapped over 353 tergite VIII basally (vs. left lobe wrapped over or at least alongside). Description (all measurements in milli- meters).—Apterous male: Ground color blackish, faintly shining, covered with very short dark pubescence; head except vertex along eyes and often longitudinal oval black spot ahead of eyes, median longitudinal wedge on pronotum, figures on mesonotum (Fig. 13), propleura, mesopleura, metace- tabulae, coxae, fore trochanters, dorsum of fore femur except distally, pygophore and venter brownish yellow to leucine. Antero- lateral angles of pronotum, longitudinal stripe on lateral margins and posterolateral angles of mesonotum, lateral margins of ab- dominal tergites, and coxal cavities dorsally covered with short bright golden pubes- cence. Legs black to brown, middle and hind femora lighter beneath basally, antennae black. Legs set with short inconspicuous black spines. Length of head (1.05) about four times narrowest interocular space (0.28); width of eye 0.58; eyes extending posteriorly about ¥% length of propleura. Rostrum short, bare- ly reaching onto mesosternum. Pronotum short (0.83), width (1.39) about equal to head across eyes. Mesonotum long (2.44), widest across mesocoxae (2.55). Metanotum short (0.78), widest across metacoxae (2.05). Ab- domen excluding genital segments relative- ly short (2.22), genital segments long (2.00). Tergites II-VI about of equal length (0.22- 0.28), tergite VII about twice as long (0.55); VIII long (1.39), large, highly modified, with a lobe on left side bearing two large “‘teeth”’ (Fig. 10); proctiger highly modified, with long projection on left side (Fig. 12); male terminalia as shown in Figs. 10-12. Con- nexiva angulate caudally, not produced; raised along entire length. Antennae long, slender, segments I and IV slightly stouter than IJ and ITI; length of segments I-IV: 1.55; 0.57; 0.50; 0.89. An- terior femur stout, tapering distally; anterior tibia slightly flattened, not dilated. Mea- surements of legs as follows: Femur, tibia, 354 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 11 Figs. 7-12. Potamobates anchicaya n. sp. 7-9, Female terminal abdomen. 7, Dorsal view, without caudal projection. 8, Dorsal view. 9, Lateral view. 10-12, Male terminal abdomen. 10, Lateral view. 11, Dorsal view. 12, Oblique ventral view. tarsal 1, tarsal 2 of fore leg, 2.33: 2.11: 0.14: 0.44; of middle leg, 8.55: 5.77: 2.39: 0.61: of hind leg, 9.26: 4.83: 0.36: 0.22. Length, mean 9.36 (N = 10, min. 9.24, max. 9.59). Width, mean 2.50 (N = 10, min. 2.31, max. 2.75). Apterous female: Body more robust than male, otherwise similar in general size, col- oration, and other features except dark markings on head usually more extensive, and abdominal terminalia very differently formed. Connexiva produced posteriorly into long slender digitate lobes, usually al- VOLUME 97, NUMBER 2 Ripenlis: sal view. Potamobates anchicaya n. sp. Thorax, dor- most parallel distally, sometimes divergent (Figs. 7, 8). Abdominal ventrite VIII pro- duced asymmetrically, embracing or par- alleling tergite VIII which may be either truncate (Fig. 7) or produced into a sinuate spine (Figs. 8, 9). Length, mean 8.59 (N = 10, min. 8.08, max. 9.24). Width, mean 2.70 (N = 10, min. 2.58, max. 2.84). Macropterous male: Similar to apterous form in most respects, except pronotum long, with posterior lobe raised and broadly margined with brownish yellow, humeri prominent. Wings light brown, reaching be- yond tip of abdomen, veins blackish brown, without pubescence, with five closed cells; wings often mutilated, presumably to facil- itate mating, sometimes without even basal stub remaining. Length, mean 10.42 (N = 6, min. 10.12, max. 10.57; de-alated specimens not mea- sured). 355 Width, mean 2.56 (N = 6, min. 2.49, max. 2.66). Macropterous female: Similar to mac- ropterous male in most respects, except slightly broader, and wings reaching far be- yond tip of abdomen. Length, mean 10.46 (N = 5, min. 10.39, max. 10.66; de-alated specimens not mea- sured). Width, mean 2.79 (N = 5, min. 2.66, max. 2.84). Etymology.—The name anchicaya, a noun in apposition, refers to the type lo- cality, the valley of the Rio Anchicaya. Ecological notes.—Potamobates anchi- caya was taken in midstream on swift, smoothly flowing reaches above riffles, in company with Metrobates fugientis Drake and Harris. Potamobates horvathi was also present at the type locality, but was found on the slower waters of pools both above and below riffles, and not in the midstream areas favored by P. anchicaya. Remarks.—Potamobates species are highly variable in the development and shape of the bizarre modifications of the abdominal terminalia in both males and fe- males, which led previous workers to over- look or misidentify specimens of P. anchi- caya in their collections. Potamobates an- chicaya specimens from Panama, for ex- ample, were identified as P. tridentatus Esaki by both Drake (in Drake Collection, USNM) and Hungerford (1937b). For many years specimens of anchicaya from the Canal Zone of Panama in the Polhemus collection have been marked as “‘not tridentatus,”’ but a sat- isfactory delineation was possible only with good series of both species available from a number of localities. Material examined.— Holotype, apterous male and allotype, apterous female (USNM), COLOMBIA: Valle de Cauca Prov., Rio Tatabro, 7 km E. of Sabaletas, water temp. 24°C, CL2436.30.vil989.Jo2& DLA. Polhemus. Paratypes as follows (all in JTPC, all apterous unless noted): COLOMBIA: Valle de Cauca Prov.: 4 males, 3 macrop- 356 terous males, 8 females, 3 macropterous fe- males, 3 nymphs, same data as holotype; 14 males, 1 female, swift rocky tributary to Rio Anchicaya, east of CL 2436, 100 m, water temp. 24 °C, CL 2435, 30.vii.1989, J. T. & D. A. Polhemus; 11 males, 11 fe- males, 2 nymphs, small forest stream near Bajo Calima research station, NE of Bue- naventura, 40 m, water temp. 25 °C, CL 2437, 30.vii.1989, J. T. & D. A. Polhemus; 1 male, 2 females, Bajo Calima, 3.111.1987, M. R. Manzano (NNC); 4 males, 2 females, Rio Tatabro, 7 km E. of Sabaletas, 1.11.1987, #87004, Gonzales (NNC; JTPC); 3 males, 3 females, Gorgona, Quebrada Acueducto, 30.x11.1989, M. Baena (NNC); 1 male, 1 macropterous female, Rio San Cipriano, 40 m, 7.vili.1989, M. R. Manzano (UVCC). PANAMA: Panama Prov. (all collected by JTP unless noted): 1 male, i macropterous male, La Cascada, small stream and water- fall on Madden Highway, about 4 km S. of Maria Eugenia, trib. of Rio Pedro Miguel, ~100;m, ‘water temp: .24 °C, CL 2771, 31.x11.1992, J. T. Polhemus & A. R. Gil- logly; 5 males, 1 macropterous male, 3 fe- males, 2 macropterous females, 1 nymph, Rio Juan Grande, off Pipeline Road, nr. Gamboa, km 2, ~30 m, CL 2784, 5.1.1993; 4 males, | female, 1 nymph, Rio Agua Sa- lud, off Pipeline Road, km 17, ~5 m, CL 27185, 5.1993: 3.males, 3 females, 2 nymphs, unnamed stream off Pipeline Road, km 13.8, ~5 m, CL 2786, 5.1.1993; 1 fe- male, unnamed stream off Pipeline Road, km 15, ~5 m, CL 2789, 6.1.1993; 7 males, 6 females, small unnamed stream off Pipe- line: Road). km. 10:6, ~5) m).CL. 2790: 6.1.1993; 1 female, small unnamed stream off Radar Site road S. of Gamboa, ~50 m, CL 2795, 7.1.1993; 1 male, 2 macropterous males, 2 macropterous females, stream crossing road between Gamboa Rd. and Cascada on Madden Hwy., ~50 m, CL 2796, 7.1.1993; 1 male, Rio Indio, Altos del Cerro Azul, E. of Panama City, 560 m, CL 2799, 8.1.1993; 2 females, stream on main road past Altos del Cerro Azul, 26.5 km from Pan Am Hwy. nr. Villa Linda, E. of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Panama City, ~900 m, CL 2800, 8.1.1993: 1 apterous male, N. of Portobello, CL 1296, 2.1.1970; 1 apterous female, Canal Zone, 10.11.1939, C. J. Drake; 1 apterous male, 1 apterous female, Pearl Islands, San Jose, 30.v11.1944, J. P. E. Morrison. San Blas Prov.: 7 males, | macropterous male, 2 fe- males, 3 macropterous females, Rio Nusa- gandi, west of Nusagandi, ~200 m, water temp. 245°C, Cio 27725 11.1993: J. T. Pol- hemus & A. R. Gillogly. Potamobates manzanoae, NEw SPECIES Figs. 14-18, 20 Diagnosis.— Potamobates manzanoae is a sister species to P. unidentatus Champion (see Figs. 30, 31), but compared to the latter manzanoae may be separated by the shorter yellowish stripe medially on the mesono- tum, which does not extend anteriorly much beyond the middle (vs. reaching to prono- tum in unidentatus); the laterally directed process of proctiger, which is larger, longer and of different shape (process in uniden- tatus small, usually hidden); and the unique, bizarre female abdominal terminalia, not resembling those of any other species (Figs. 14, 15). Description (all measurements in milli- meters).—Apterous male: Ground color blackish, faintly shining, covered with very short dark pubescence; head except vertex along eyes and longitudinal oval black spot ahead of eyes, median longitudinal wedge on pronotum, figures on mesonotum (sim- ilar to P. anchicaya but somewhat smaller; see Fig. 13), propleura, mesopleura, metace- tabulae, coxae, fore trochanters, dorsum of fore femur except distally, and venter brownish yellow to leucine. Anterolateral angles of pronotum, longitudinal stripe on lateral margins and posterolateral angles of mesonotum, lateral margins of abdominal tergites, and coxal cavities dorsally covered with short bright golden pubescence. Legs black to brown, middle and hind femora lighter beneath basally, antennae black. Legs set with short inconspicuous black spines. VOLUME 97, NUMBER 2 Figs. 14-18. Potamobates manzanoae n. sp. 14, 15, Female terminal abdomen. 14, Dorsal view. 15, Lateral view. 16-18, Male terminal abdomen. 16, Dorsal view. 17, Ventral view. 18, Lateral view. Length of head (1.17) about four times narrowest interocular space (0.28); width of eye 0.61; eyes extending posteriorly about ¥ length of propleura. Rostrum short, bare- ly reaching onto mesosternum. Pronotum short (0.83), width (1.44) about equal to head across eyes. Mesonotum long (2.72), widest across mesocoxae (2.66). Metanotum short (0.88), widest across metacoxae (2.33). Ab- domen excluding genital segments relative- ly short (2.55), genital segments long (2.00). Tergites II-V about of equal length (0.17- 0.22), tergite VI longer (0.33), tergite VII longest (0.67); VIII long (1.33), large, mod- ified, asymmetrical beneath (Fig. 17); proc- tiger highly modified, with long projection on left side (Fig. 17); male terminalia as shown in Figs. 16-18. Connexiva angulate caudally, not produced; slightly raised along entire length. Antennae long, slender, segments I and IV slightly stouter than II and III; length of segments I-IV: 1.83; 0.67; 0.67; 1.00. An- terior femur stout, tapering distally; anterior tibia slightly flattened, not dilated. Mea- surements of legs as follows: Femur, tibia, tarsal 1, tarsal 2 of fore leg, 2.77: 2.60: 0.11: 0°44: of middle les. 91052 6.22-°2:77- 0.72: of hind leg, 9.54: 5.38: 0.50: 0.28. Length, mean 9.72 (N = 2, min. 9.41, max. 10.04). Width, mean 2.58 (N = 2, min. 2.49, max. 2.66). Apterous female: Body more robust than male, otherwise similar in general size, col- oration, and other features except dark 358 markings on head usually less extensive, and abdominal terminalia very differently formed. Connexiva truncate, not produced (Fig. 14). Abdominal ventrite VIII strongly produced posteriorly, long, asymmetrical, twisted, spatulate; tergite VIII long, pro- duced, acuminate distally (Figs. 14, 15). Length, 9.24 (N = 1). Width, mean 2.84 (N = 1). Macropterous male: Similar to apterous form in most respects, except pronotum long, with posterior lobe raised and broadly margined with yellowish, humeri promi- nent. Wings light brown, reaching beyond tip of abdomen, veins blackish brown, with- out pubescence, with four closed cells, distal cell open. Length, mean 10.39 (N = 2, min. 10.30, max. 10.48). Width, mean 2.53 (N = 2, min. 2.49, max. 2.58). Macropterous female: Similar to mac- ropterous male in most respects, except slightly broader, and wings reaching far be- yond tip of abdomen. Length, 10.39 (N = 1). Width, 2.75 (N = 1). Etymology.— The name manzanoae hon- ors Maria Rosario Manzano for her contri- butions to our knowledge of the aquatic Heteroptera of Colombia. Ecological notes.—P. manzanoae was taken in midstream in areas of swift, smooth current. Material examined.— Holotype, apterous male and allotype, apterous female (USNM), COLOMBIA: Valle de Cauca Prov., swift rocky tributary to Rio Anchicaya, water temps 24 °C, Ch 2435,.50-vii. 1989. Je Dak D. A. Polhemus. Paratypes as follows (all in JTPC, all apterous unless noted): CO- LOMBIA: Valle de Cauca Prov.: 1 male, 1 macropterous male, | female, | macropter- ous female, same data as holotype; | male, 1 female, Rio Danubio, 35 km W. of Quer- emial; Cl 2432, 30.vi.1989..J. 1. & DA. Polhemus. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Potamobates tridentatus Esaki Fig. 19 The range of P. tridentatus extends from northern Costa Rica southward to Chiriqui Province in northern Panama. Repeated collections southward have not revealed any intervening populations between Chiriqui Province and the region of the Panama Ca- nal Zone, where P. anchicaya is common. Esaki (1926) provided splendid figures of the abdominal terminalia of both males and females that illustrate the differences be- tween these two species (see discussion un- der P. anchicaya). Wheelwright and Wilkinson (1985) stud- ied the ecology of P. tridentatus in the Osa Peninsula of Costa Rica. Material examined. —(all in JTPC, all ap- terous unless noted). COSTA RICA: Gua- nacaste Prov.: 1 male, 12.vii.1957, D. R. Leach. Puntarenas Prov.: 1 male, 2 females, Canas, Rio Lagarto, 9.xi.1981, E. Berrera & H. Brailovsky; 2 males, | macropterous male, 2 females, | macropterous female, river 24 km N of Esparta, CL 1264, 24.xi1.1969, J. T. Polhemus. PANAMA: Chiriqui Prov.: 5 males, 1 female, Balnearo, river 3.8 km N of David, CL 2819, 13.1.1993, J. T. Polhemus & A. R. Gillogly; 1 female, small stream ~110 km S of Da- vid, CL 1289, 31.xii.1969, J. T. Polhemus. Potamobates horvathi Esaki Fig. 20 This is the commonest and most wide- spread species in the genus, with a range from southern Mexico to Colombia. It oc- curs in both the Atlantic and Pacific drain- ages, usually at low elevations. Matsuda (1961) studied the relative growth of all nymphal instars and adults of this species in a Panamanian population. Material examined.—(all in JTPC, all ap- terous unless noted). BELIZE: | male, Rio Grande, Nov. 1931; 2 females, Punta Gor- da, 1932. COLOMBIA: Valle de Cauca Prov.: 4 males, 4 females, Rio Tatabro, 7 km E. of Sabaletas, water temp. 24 °C, CL VOLUME 97, NUMBER 2 359 Fig. 19. Distributions of Potamobates species. Solid triangles: P. tridentatus Esaki. Open triangle: P. carvalhoi n. sp. 2436, 30.vii.1989, J. T. & D. A. Polhemus; 3 males, 1 macropterous male, | female, 2 macropterous females, Rio San Cipriano, 40 m, 7.vil.1989, M. T. Manzano (UVCC). COSTA RICA: Guanacaste Prov.: 4 males, 2 macropterous males, 4 females, 3 macrop- terous females, 1 nymph, Rio Sanzapote, 6 km N of La Cruz, nr. Nicaragua border, CL 1307, 8.1.1970, J. T. Polhemus. Puntarenas Prov.: 6 males, 2 macropterous males, 4 females, | macropterous female, 1 nymph, mven 24 km N of Esparta; (CL 1264; 24.x11.1969, J. T. Polhemus. GUATEMA- LA: | male, Esquintla; 2 males, 1 female, Los Amates; 2 males, 7 females, | macrop- terous female, small river, Azucho, 24 km S of Esquintla, CL 1252, 20.xii.1969, J. T. Polhemus; 5 males, 2 macropterous males, 1 female, Stain Creek, 11 km N of Zacapa, CL 1316, 11.1.1970, J. T. Polhemus; 5 males, 1 nymph, stream, E of Quirigua, CL 1317, 11.1.1970, J. T. Polhemus. HONDURAS: 1 male, Lancetilla, 22.111.1936, John Deal. NICARAGUA: 1I5 males, 4 females, small Stream, 3 “km™N of Esteli; \CL 1262. 23.x11.1969, J. T. Polhemus. PANAMA: Bocas del Toro Prov.: 2 males, 5 females, deep stream on road to oil tanks, S. of Chi- riqui Grande, sea level, CL 2829, 14.1.1993, J. T. Polhemus & A. R. Gillogly; 1 male, 1 female, vegetated stream, 5.1 km S. Punta Pena. 50am, Cle2830) 14:7.19938 I. 1. Polhemus & A. R. Gillogly. Chiriqui Prov.: 1 macropterous female, Balnearo, river 3.8 km N of David, CL 2819, 13.1.1993, J. T. Polhemus & A. R. Gillogly; Colon Prov.: 2 360 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 20. Distributions of Potamobates species. Solid triangles: P. horvathi Esaki. Open triangle: P. manzanoae n. sp. males, 2 females, 2 km N of Maria Chiquita, 27.11.1993, A. R. Gillogly; 2 males, 4 fe- males, 3 nymphs, N of Portobello, sea level, CL 1296, 2.1.1970, J. T. Polhemus. Herrera Prov.: 1 male, | macropterous male, 3 fe- males, Rio Paritas, 2.8 km S of Oct, CL 2802, 9.1.1993, J. T. Polhemus & A. R. Gil- logly; Panama Prov.: 3 males, | macrop- terous male, 3 females, 2 macropterous fe- males, Rio Cabra, ~3 km N of jet. to Altos Cerro, Azul, ~20 mz CL, 2797, 741.1993.1; T. Polhemus; 7 males, 16 females, 5 km N of San Carlos, CL 1300, 4.1.1970, J. T. Pol- hemus; 2 males, Panama City, 11.1939, C. J. Drake; 1 macropterous male, Canal Zone, 10.11.1939, C. J. Drake; 9 males, 7 females, 3 nymphs, E of Panama City, CL 1295, 2.1.1970, J. T. Polhemus. Potamobates unidentatus Champion Fig. 22 This species is common and widespread throughout Costa Rica and Panama, occur- ring in both the Pacific and Atlantic water- sheds, in contrast to P. tridentatus which has so far been found only in Pacific drain- ages. We did not collect this species during our surveys in Colombia, and to our knowl- edge it has been reported from the country only once, at Mamatoco, Magdalena Prov- ince (Esaki 1926). Matsuda (1961) studied the relative growth of all nymphal instars and adults of this species in a Costa Rican population. Material examined. —(all in JTPC, all ap- terous unless noted). COSTA RICA: Car- VOLUME 97, NUMBER 2 Figs, 21: P. anchicaya n. sp. tago Prov.: 1 male, 1 female, km 20, Tur- rialba—Siquirres Rd., 22.11.1982, E. Ber- rera. Heredia Prov.: 19 males, 1 macrop- terous male, 18 females, 1 macropterous female, Ojo de Agua, nr. San Antonio, CL 1269, 25.x11.1969, J. T. Polhemus. Limon Prov.: 1 male, 2 females, Rio Santa Clara, Guapiles, 26.iv.1984, E. Berrera. Puntare- nas Prov.: 1 male, 2 macropterous males, 3 females, 3 macropterous females, 1 nymph, Rio Java, nr. San Vito de Java, CL 1283, 29.xii.1969, J. T. Polhemus. PANAMA: Bocas del Toro Prov.: 2 males, 4 females, stream 6.6 km N. of Continental Divide on rd. to Chiriqui Grande, ~600 m, CL 2824, 13.1.1993, J. T. Polhemus & A. R. Gillogly; 6 males, 4 females, tributaries to Rio Hor- nitos, Reserva la Fortuna, 1400 m, CL 2825, Distributions of Potamobates species. Solid triangles: P. vivatus Drake & Roze. Open triangle: 14.1.1993, J. T. Polhemus & A. R. Gillogly. Coclé Prov.: 1 male, 2 females, stream in forest below El Valle, CL 1299, 3.1.1970, J. T. Polhemus. Panama Prov.: 2 males, 1 fe- male, 1 macropterous female, Rio Indio, Altos Cerro Azul, 558 m, CL 2799, 8.1.1993, J. T. Polhemus; 1 male, small stream nr. Villa Linda, W. of Altos Cerro Azul, 26 km from Pan Am Hwy, ~900 m, CL 2800, 8.1.1993, J. T. Polhemus. San Blas Prov.: 1 male, | female, Rio Nusagandi, W. of Nu- sagandi, ~200 m, CL 2772, 1.1.1993, J. T. Polhemus & A. R. Gillogly; 4 males, 9 fe- males, small stream nr. Nusagandi, ~500 mm: CEI27 742711993: J: Polhemus: 2 males, 4 females, waterfall ~5 km E. of Nu- sagandi, ~450 m, CL 2775, 2.1.1993, J. T. Polhemus & A. R. Gillogly. 362 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 22. Distribution of Potamobates unidentatus Champion. Potamobates vivatus Drake and Roze Big. 21 This species was originally described from two localities in the coastal ranges of north- ern Venezuela. We did not collected this species in Colombia, and to our knowledge it has been taken in this country at only a single locality, the Serrania de la Macarena in Meta Province, an isolated sandstone massif lying to the east of the main Andean uplift (Fig. 21). Material examined. —(all in JTPC, all ap- terous unless noted). COLOMBIA: Meta Prov.: 7 males, 1 macropterous male, 5 fe- males, 4 nymphs, Rio Santo Domingo, Ser- rania de la Macarena, 24.i1.1989, C. Murillo (JTPC, UVCC). VENEZUELA: Distrito Federal: 1 male paratype, Cerro el Avila, 16.viii.1952, J. A. Roze; Carabobo Prov.: 1 female paratype, Belén, 27.vi.1953, J. A. Roze. KEY TO THE SPECIES OF POTAMOBATES Species keys were last published for the genus Potamobates by Drake and Harris (1934) and Kuitert (1942). The former in- cluded only six, and the latter only ten, of the fifteen presently known species, there- fore a new key is provided below, including all described taxa. A fully illustrated key, though desirable, was beyond the scope of this paper, whose primary focus is the fauna of Colombia; the present key thus makes reference to previously published figures, with new illustrations being provided only for those species in which one or both sexes have never before been figured. 363 VOLUME 97, NUMBER 2 ey 27 Figs. 23-29. Potamobates spp.. abdominal terminalia. 23, P. osborni Drake & Harris, female, lateral view. 24, P. bidentatus Champion, female, lateral view. 25-27, P. vivatus Drake & Roze; 25, female, lateral view; 26, male, lateral view; 27, male, ventral view. 28, 29, P. williamsi Hungerford, female; 28, lateral view; 29, postero- dorsal view. 364 2(1). 3(2). 4(3). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pronotum medially with a roughly par- allel sided linear longitudinal yellow stripe. Male abdominal segment VIII modified, segments IX, X rotated a maximum of 1} Meee 8 Ben cht cs cco ndiy Tose D Pronotum medially with a roughly tri- angular or wedge shaped yellow mark. Male abdominal segment VIII modified (except thomasi), segments IX, X rotated variably in each species, from about 10° to about: 900 sao ee nee 5 Male abdominal segment VIII not mod- ified; segments IX, X not rotated. Female with short sharp connexival spines, and a pair of ventral prolongations; anal cone not tubular (see Hungerford 1937, pl. II; habitus, Andersen 1982, fig. 438) ...... dee JG CM sv SEN ee thomasi Hungerford Male abdominal segment VIII modified, with an asymmetrical tumescence or tooth (Fig. 17); segments IX, X rotated at most 30° to 45°. Female without connexival spines or ventral prolongations; anal cone tubular (Bigss23) 24) san petri aor 3 Mesonotum of apterous forms without light markings, except golden or silvery setae forming stripes or spots. Male ab- dominal segment VIII broad, almost par- allel sided, distally modified asymmetri- cally. Female abdominal segment VIII dorsally produced into long spine; ven- trally anal cone long, curved, tubular (Figs. 23, 24) Mesonotum of apterous forms with me- dian yellow longitudinal stripe (rarely completely dark). Male abdominal seg- ment VIII narrow, tapering posteriorly, not expanded distally, with only a small tooth (see Drake and Harris 1934, pl. XXVI f, g). Female abdominal segment VIII dorsally triangular, ventrally with a short finger-like protuberance (anal cone) (see Matsuda 1960, fig. 471) .. horvathi Esaki Vertex of head largely yellow, with a large elongate black region anteriorly, mostly anterad of eyes. Male abdominal segment VIII modified, with an asymmetrical tu- mescence (see Drake and Harris 1934, pl. XXV h). Female abdominal segment VIII dorsally with moderately long process (Fig. 18) RR ae Bae ty se osborni Drake and Harris Vertex of head largely black, with a small elongate yellow spot between eyes. Male abdominal segment VIII modified, with an asymmetrical lobe bearing two blunt teeth (see Drake and Harris 1934, pl. 5(1). 6(5). 7(5). 8(7). 9(8). XXVI a). Female abdominal segment VIII dorsally with moderately long process (Fig. DAY Pa aiik cree ee bidentatus Champion Male abdominal segment VIII modified, segments IX, X rotated a maximum of 45°. Female without connexival prolon- gations Male abdominal segment VIII modified, segments IX, X rotated about 90°. Female with large connexival prolongations ..... 7 Male abdominal segment IX distally light; process of proctiger spatulate, usually hid- den; segments IX, X rotated 5° to 10° (see Matsuda 1960, fig. 479). Female abdom- inal segment VIII dorsally triangular; ven- trally with a triangular structure (see Mat- suda 1960, fig. 472) .. unidentatus Champion Male abdominal segment IX distally dark; process of proctiger acuminate, exposed (Fig. 17); segments IX, X usually rotated 30° to 45°. Female abdominal segment VIII dorsally produced into long spine-like structure; ventrally with an asymmetrical spoon-shaped structure (Figs. 14, 15) .. PA ee ene le eet Se manzanoae 0. sp. Male abdominal segment VIII broad, short, slightly modified (Figs. 2, 5, 6). Fe- male abdominal terminalia truncate (Figs. 1 3244) Y Oe ee ee ne oe carvalhoi n. sp. Male abdominal segment VIII longer, modified, with spinose or dentate lobes on right side (Figs. 10-12). Female ab- dominal segment VIII dorsally with short to long median digitate or spine-like pro- longation (polymorphic, sometimes short), but never truncate (Figs. 7-9) Distal lobe on right side of male abdom- inal segment VIII forming a single acute angle posteroventrally (Figs. 26, 27). Fe- male abdominal segment VIII dorsally produced into a long posterodorsally di- rected’ spine(Fig425) nanos vivatus Drake Distal lobe on right side of male abdom- inal segment VIII variably formed, but not forming a single acute angle postero- ventrally. Female abdominal segment VIII dorsally with short or long process, but not posterodorsally directed Distal lobe on right side of male abdom- inal segment VIII forming a posteriorly directed acute angle, plus a posteroven- trally directed twisted spatulate lobe (see Polhemus and Polhemus 1983, fig. 1). Fe- male abdominal segment VIII dorsally short, triangular; ventrally with a long process extending posteriorly far beyond VOLUME 97, NUMBER 2 10(9). 11(10). 12(11). tip of dorsal triangular plate on left side (see Hungerford 1937, pl. II) .......... OCR abet oan woytkowskii Hungerford Distal lobe on right side of male abdom- inal segment VIII variably formed, but not forming a single acute angle postero- ventrally. Female abdominal segment VIII dorsally with short or long process; ven- trally with short or long process, but not extending posteriorly far beyond dorsal SIUCIIIC Ret Ae Thr rete eee Distal lobe on right side of male abdom- inal segment VIII forming a posteriorly directed short spine, plus a posteroven- trally directed acute angle (see Polhemus and Polhemus 1983, fig. 2). Female ab- dominal segment VIII dorsally produced into a long posteriorly directed finger-like process; ventrally with a long twisted pro- cess on left side extending posteriorly half the length of dorsal process (see Polhemus and Polhemus 1983) fig. 3). 2 ..o-6-...- 10 ph Ls, Sei toe spiculus Polhemus & Polhemus Distal lobe on right side of male abdom- inal segment VIII variably formed, but not forming a short spine, plus an acute angle. Female abdominal segment VIII dorsally with short or long process; ven- trally with short flap-like process not ex- tendine far posteriorly...) 5 ya.) Distal lobe on right side of male abdom- inal segment VIII forming a posteriorly directed twisted spatulate protuberance (see Hungerford 1937, pl. II). Female ab- dominal segment VIII dorsally triangular; ventrally with a short flap-like lobe on left side (see Matsuda 1960, fig. 460) ...... Sire RA Ah aC lca peruvianus Hungerford Distal lobe on right side of male abdom- inal segment VIII forming two ventrally or posteroventrally directed spines. Fe- male abdominal segment VIII dorsally with short or long process; ventrally with short flap-like process not extending far POStEHIOrIYseet ye tece ee as oe taee Distal lobe on right side of male abdom- inal segment VIII forming two postero- ventrally directed teeth, ventral one lon- ger; process of proctiger bifurcate distally (see Matsuda 1960, fig. 479). Female ab- dominal segment VIII dorsally with short stout process, directed posteroventrally; ventrally with short flap-like process cov- ering left side of dorsal process (see Mat- 12 suda 1960, fig. 473) .. variabilis Hungerford Distal lobe on right side of male abdom- 365 inal segment VIII forming two ventrally or posteroventrally directed spines of about equal length; process of proctiger not bifurcate distally. Female abdominal segment VIII dorsally with short or long process; ventrally with short flap-like pro- CESS) je ios oe oR Pee accel: 13 13(12). Distal lobe on right side of male abdom- inal segment VIII forming two widely spaced posteroventrally directed stout spines (see Hungerford 1937, pl. II). Fe- male abdominal segment VIII dorsally with short triangular process; ventrally with short flap-like process usually cov- ering left side and part of right, almost hiding dorsal process (Figs. 28, 29) .... cha ran asehd ra eae williamsi Hungerford - Distal lobe on right side of male abdom- inal segment VIII forming two ventrally directed closely spaced spines of about equal length. Female abdominal segment VIII dorsally with short or long process; ventrally with short flap-like process ... 14 14(13). Distal lobe on right side of male abdom- inal segment VIII forming two ventrally directed stout spines with a pronounced rounded incision between (see Drake and Harris 1934, pl. XXVI c, d, e). Female abdominal segment VIII dorsally with short or long process, never bent ventrad; ventral short flap-like process with left lobe only developed (see Drake and Harris 1934. DIsexcK VID) tease tridentatus Esaki - Distal lobe on right side of male abdom- inal segment VIII forming two ventrally directed stout spines with a shallow tri- angular incision between (Figs. 10-12). Female abdominal segment VIII dorsally with short or long process, often bent ven- trad; ventral short flap-like process with both right and left lobes developed (Figs. EU) epee roi en Cio eio orc anchicaya n. sp. POTAMOBATES SPECIES GROUPS AND SPECIES CHECKLIST When this study began, we initially di- vided the genus Potamobates into species groups based on the complexity of the male and female abdominal terminalia, which exhibit the most salient species-specific characters in this genus. We then arranged these groups in what we considered to be rough phylogenetic order, with those exhib- 366 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON iting the largest number of plesiomorphic character states discussed first in the an- notated checklist that follows. The geo- graphic progression of phylogenetically in- creasing complexity from north to south suggested an ancestral stock in Mexico, and a secondary radiation southward through Mesoamerica into South America. The ge- nus is currently known on this latter con- tinent only in the Andean mountain chain from Colombia to Peru, and in the coastal sierras of northern Venezuela, inhabiting swift clear streams associated with moun- tainous terrain. The checklist below, arranged by groups, contains all known species and includes the references for the original descriptions of all previously described species. Kirkaldy (1899) tentatively identified material from Ecuador as P. bidentatus and P. unidentatus; this material, in the Turin Museum, has not been restudied, but the identifications are almost certainly incorrect, and the records are not included here, because many new species have since been described from South America. New distributional records for described species are indicated with an asterisk (*). P. thomasi group Diagnosis.— Male genitalia simple, only slightly modified; segment IX not rotated; abdominal segment VIII not modified; proctiger with modest modification. Female with small connexival spines; abdominal segment VII ventrally modified, with two (1 + 1) lateral prolongations; VIII, IX not modified. thomasi Hungerford 1937: 63. Mexico (western) P. unidentatus group Diagnosis.— Male genitalia relatively simple; segment IX slightly rotated, or ro- tated a maximum of 30° to 45°; abdominal segment VIII at most slightly modified ven- trally. Female without significant connexi- val prolongations; abdominal segment VIII dorsally without or with modest median prolongation, ventrally without or with only a modest prolongation; anal cone tubular, longest in Mexican species, shortest in wni- dentatus. bidentatus Champion 1898: 155. Eastern Mexico (Catemaco area). [Note: P. bi- dentatus, collected by “‘Sallé, in Mexi- co” (1831-1834), remained an enigma ever since its original description, with the exact collection locality unknown, until Brailovsky and coworkers redis- covered it in the Sierra de Tuxtla, ona peninsula near Catemaco, where it is apparently endemic. The female, pre- viously unknown, is figured herein (Fig. 24). P. osborni also occurs in the same general area, but further inland.] horvathi Esaki 1926: 254. Belize, Colom- bia*, Costa Rica, Guatemala, Hondu- ras, Mexico (Chiapas), Nicaragua*, Panama manzanoae J. & D. Polhemus, n. sp. Co- lombia osborni Drake & Harris 1928: 25. Eastern Mexico unidentatus Champion 1898: 155. Co- lombia, Costa Rica, Panama P. carvalhoi group Diagnosis.— Male genitalia relatively simple, only slightly modified, very broad, short; segment IX rotated almost 90°; ab- dominal segment VIII slightly modified ventrally; proctiger modified, prolongation long, complex. Female tergite VII de- pressed, connexival prolongations long; ventrite VII prolonged, upturned distally; abdominal segments VIII, [IX small, un- modified, symmetrical, withdrawn. carvalhoi J. & D. Polhemus, n. sp. Co- lombia, Venezuela P. tridentatus group Diagnosis.— Male genitalia highly modi- fied, segment IX strongly rotated; abdom- inal segment VIII modified, with spines or modest to large lobe-like extension on right side (sometimes divided); proctiger with a plate-like or long curved spine-like exten- sion, distally always spine-like. Female with long connexival prolongations; abdominal VOLUME 97, NUMBER 2 367 C.linearis anchicaya tridentatus williamsi peruvianus spiculus vivatus variabilis woytkowskii carvalhoi manzanoae unidentatus bidentatus osborni horvathi thomasi C. linearis anchicaya tridentatus peruvianus spiculus vivatus variabilis woytkowskii williamsi carvalhoi manzanoae unidentatus bidentatus osborni horvathi thomasi C.linearis anchicaya peruvianus woytkowskii variabilis spiculus vivatus tridentatus williamsi carvalhoi manzanoae unidentatus bidentatus osborni horvathi thomasi Fig. 30. Three most parsimonious trees generated from preliminary phylogenetic analysis of Potamobates species (see Appendices | and 2 for explanation). 368 vc Fig. 31. Preferred cladogram for Potamobates species, with numbering refers to that employed in Appendix 1). Numbers in PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ol al 9'€ Ol ‘6 to apomorphic state along a given branch. Ol 9% C. linearis woytkowskii peruvianus variabilis spiculus vivatus williamsi anchicaya tridentatus carvalhoi manzanoae unidentatus horvathi bidentatus osborni thomasi characters used in its construction (character dicate characters changing from plesiomorphic VOLUME 97, NUMBER 2 369 South America Peru Peru Peru Peru Colombia, Venezuela Ecuador Colombia, Panama Costa Rica, Panama Colombia, Venezuela Colombia Colombia, Panama, Costa Rica Colombia to Southern Mexico Southeastern Mexico Southeastern Mexico Western Mexico Fig. 32. Taxon-area cladogram based on preferred phylogeny for Potamobates species (see Fig. 31). segment VIII modified, dorsally short, tri- angular or with a long median spine-like prolongation, ventrally with one or two short to long flap-like lobes, or with complex spoon-like prolongation. anchicaya J. & D. Polhemus, n. sp. Co- lombia (western), Panama peruvianus Hungerford 1936: 178. Peru spiculus J. & D. Polhemus 1983: 286. Peru tridentatus Esaki 1926: 251. Costa Rica, Panama variabilis Hungerford 1938: 85. Peru vivatus Drake & Roze 1954: 228. Colom- bia* (eastern), Venezuela williamsi Hungerford 1932: 228. Ecuador woytkowskii Hungerford 1937: 144. Peru 370 We have subsequently tested our hypoth- esis of species groupings via a preliminary cladistic analysis with a more comprehen- sive character set (see Appendices 1 and 2), which reveals that our groupings based on genitalic characters alone represent three monophyletic clades plus one basal para- phyletic grade (the unidentatus group). Our initial attempts to further discriminate two separate groupings among the South Amer- ican species forming a clade at the far end of the tree (the tridentatus group) are not supported phylogenetically, since the entire suite of characters used contains systems that vary independently to such a degree that all groupings based on genitalic struc- tures alone end up being polyphyletic in at least one of the most parsimonious trees. The north to south biogeographic pro- gression in genitalic character development alluded to previously was supported based on both our initial intuitive group hypoth- esis, and on the results of the phylogenetic analysis. In addition, the taxon-area clado- gram derived from the phylogenetic anal- ysis (Fig. 32) strikingly illustrates Colom- bia’s pivotal role as a crossroads or Neo- tropical biogeography, which has acted as both a conduit between South America and Mesoamerica, and as an insular center of regional diversification. In marked contrast to the Peruvian Potamobates species, which all arise from a single clade, the Colombian Potamobates fauna contains members of all but one of the currently recognized intra- generic clades, including all the constituents of the paraphyletic unidentatus group as de- fined herein. ACKNOWLEDGMENTS We are indebted to the following col- leagues for the loan or gift of material: M. R. Manzano, University de Valle de Cauca, Cali, Colombia (UVCC), and Nico Nieser, Tiel, The Netherlands (NNC). Holotypes of all new species described herein are depos- ited in the National Museum of Natural History, Smithsonian Institution, Washing- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ton, D.C. (USNM). All remaining material examined in the course of this study is held either in the foregoing institution, or in the J. T. Polhemus Collection, Englewood, Col- orado (JTPC). The field surveys that cata- lyzed this investigation were funded by Na- tional Geographic Society Grant 4092-89, and we gratefully acknowledge their contin- ued support for research into the system- atics and zoogeography of aquatic Heter- optera. LITERATURE CITED Andersen, N. M. 1982. The Semiaquatic Bugs (He- miptera, Gerromorpha). Phylogeny, Adaptations, Biogeography and Classification. Scandinavian Science Press, Klampenborg, Denmark, Ento- monograph Vol. 3, 455 pp. Champion, G. C. 1897-1901. Insecta Rhynchota (Hemiptera—Heteroptera). Vol. II. Jn Goodwin and Salvin, eds., Biologia Centrali-Americana. Lon- don, xvi + 416 pp., 22 pls. Drake, C. J. and H. M. Harris. 1928. Three new gerrids from North America (Hemip.). Proceed- ings of the Biological Society of Washington 41: 25-30. 1934. The Gerrinae of the western hemi- sphere (Hempitera). Annals of the Carnegie Mu- seum 23: 179-240. Drake, C. J. and J. A. Roze. 1954. New Venezuelan Gerridae (Hemiptera). Proceedings of the Biolog- ical Society of Washington 67: 227-230. Esaki, T. 1926. On some new species of Potamobates (Hemiptera: Gerridae). Annales Musei Nationalis Hungarici 23: 251-257. Hungerford, H. B. 1932. A new Potamobates (Ger- ridae). Bulletin of the Brooklyn Entomological So- ciety 27: 228-230. 1936. A new Potamobates from Peru, S. A. (Hemiptera, Gerridae). Bulletin of the Brooklyn Entomological Society 31: 178-180. 1937a. A new Potamobates from Mexico (Hemiptera—Gerridae). Journal of the Kansas En- tomological Society 10: 63-65. . 1937b. Asecond new Potamobates from Peru, S. A., with notes on other species (Hemiptera— Gerridae). Bulletin of the Brooklyn Entomological Society 32: 144-147. 1938. A third new Potamobates from Peru, S. A. (Hemiptera—Gerridae). Journal of the Kansas Entomological Society 11: 85-87. Kirkaldy, G. W. 1899. Viaggio del Dr. Enrico Festa nell’Ecuador e regione vicine. Bolletino dei Musei VOLUME 97, NUMBER 2 di Zoologia ed Anatomia comparata della Univ- ersia di Torino 14(350): 1-9. Kuitert, L. C. 1942. Gerrinae in the University of Kansas Collections. University of Kansas Science Bulletin 28: 113-143. Matsuda, R. 1960. Morphology, evolution and a clas- sification of the Gerridae (Hemiptera—Heterop- tera). University of Kansas Science Bulletin 41: 25-632. . Studies of relative growth in Gerridae Hemip- tera-Heteroptera). I-III. Annals of the Entomo- logical Society of America 54: 578-598. Polhemus, J. T. and D. A. Polhemus. 1983. A new species of Potamobates from Peru (Hemiptera- Heteroptera: Gerridae). Journal of the Kansas En- tomological Society 56: 286-288. Swofford, D. L. 1990. PAUP: Phylogenetic Analysis Using Parsimony, Version 3.0m. Computer pro- gram distributed by the Illinois Natural History Survey, Champaign, Illinois. Wheelwright, N. T. and G.S. Wilkinson. 1985. Space use by a Neotropical water strider (Hemiptera: Gerridae): Sex and age-class differences. Biotro- pica 17: 165-169. APPENDIX | Phylogenetic analysis of Potamobates species Characters and polarity.—The set of characters listed below were employed in a preliminary analysis of Potamobates phy- logeny. For each character, two or more dis- crete states were defined and symbolized by numbers. The most primitive (plesiomor- phic) state was assigned a zero (0) value, and each successive derived (apomorphic) state a value of | or greater using successive integers. Multistate characters (those with more than two states) were ordered in linear transformation series, with 0 the most ple- siomorphic state and each subsequent state assumed to be more apomorphic. Character polarities were generally determined a priori by complexity, with a 0 state being the least complex and therefore presumably the most plesiomorphic. Ina few cases characters were also ordered by comparison with the out- group (Cylindrostethus linearis). 1. Male distal genital segments (abdomi- nal segments IX, X): 0 = not rotated: 1 = rotated 5° to 10° max.; 2 = rotated 371 30° to 45° max.; 3 = rotated approx. 90°: 2. Modifications to male first genital seg- ment (abdominal segment VIII): 0 = not modified; 1 = distal ventral or lat- eral angle or angles; 2 = distal tumes- cence on left side; 3 = distal angulate lobe on left side; 4 = distal dentate lobe on left side. 3. Median pronotal markings, male, fe- male: 0 = roughtly linear stripe; 1 = triangular or wedge shaped mark; 2 = absent. 4. Female connexival spines: 0 = short spines, acuminate; | = absent; 2 = long, thickened. 5. Female anal cone: 0 = not tubular; | = tubular, short; 2 = tubular, long. 6. Mesonotum, medial markings: 0 = dark, no light marks; 1 = longitudinal light stripe; 2 = complex figures. 7. Female ventrite VII: 0 = unmodified; 1 = modified, with prolongations. 8. Modification of male proctiger: 0 = simple angle; 1 = long process, distally simple; 2 = long process, distally “dog head” shaped; 3 = long process, distally bifurcate; 4 = long process, distally strongly twisted (corkscrew fashion), at least 90°. 9. Female first dorsal genital segment (ter- gite VIII): 0 = triangular; 1 = posterior extension digitate, short; 2 = posterior extension spinose, long. 10. Female first ventral genital segment (ventrite VIII): 0 = triangular; 1 = trun- cate, hidden; 2 = short flaps, often asymmetrical; 3 = posterior extension spatulate, elongate, not twisted; 4 = posterior extension complex, twisted, narrow, spoon-shaped, long. Computer analysis.—A cladistic analysis of Potamobates species was performed us- ing the program PAUP 3.0.m (D. L. Swof- ford, Illinois Natural History Survey). This program generates phylogenies using the principle of maximum parsimony, and eliminates trees that are identifical due to 372 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON zero length branches. The ingroup taxa for the analysis were the 15 species in the genus Potamobates, while the outgroup for the analysis was Cy/lindrostethus linearis, an- other Neotropical member of the subfamily Cylindrostethinae. Using the rooting option on PAUP, the outgroup was constrained to be a monophyletic sister group to the in- group. The PAUP analysis was carried out using the branch and bound option, using the fur- thest addition sequence and collapsing zero length branches to yield polytomies. This analysis produced three most parsimonious trees, depicted in Fig. 30. These trees had lengths of 47, consistency indices of 0.53, and retention indices of 0.52. Two of the three trees were fully resolved, while one of the trees (tree 2 of Fig. 30) had an internal polytomy at the base of the clade containing the insular South American species. Based on evolutionary considerations involving genitalic characters and biogeographic par- simony, one of the two fully resolved trees (tree 1 of Fig. 30) was chosen as a preferred phylogeny, and is shown (with some branches swapped) in Fig. 31. A taxon-area cladogram based on this tree is given in Fig. 32, and depictes a clear biogeographic pro- gression from plesiomorphic clades occu- pying Mesoamerica to apomorphic clades in South America. APPENDIX 2. Coding of characters for phyloge- netic analysis. Character matrix for Potamobates spe- cies (character states are ordered sequentially for char- acters 1-10 as given in Appendix 1). Taxon Character States Cylindrostethus linearis' 0000100001 Potamobates anchicaya 3412020122 Potamobates bidentatus 2101200221 Potamobates carvalhoi 3212020201 Potamobates horvathi 2101210000 Potamobates manzanoae 2111020024 Potamobates osborni 2101210121 Potamobates peruvianus 3312000402 Potamobates spiculus 3412000423 Potamobates thomasi 0000111000 Potamobates tridentatus 3412010122 Potamobates unidentatus 1111120002 Potamobates variabilis 3412000312 Potamobates vivatus 3312010422 Potamobates williamsi 3412020102 Potamobates woytkowskii 3412020403 ' Outgroup. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 373-378 CARVALHOISCA, A NEW GENUS OF ORTHOTYLINI FROM MEXICO (MIRIDAE, HETEROPTERA) J. C. SCHAFFNER AND P. S. F. FERREIRA (JCS) Department of Entomology, Texas A&M University, College Station, Texas 77843, U.S.A.; (PSF) Departamento de Biologia Animal Universidade Federal de Vicosa, 36570 Vicosa, Minas Gerais, Brasil. Abstract.—The orthotyline genus Carvalhoisca and the species C. jacquiniae from the state of Oaxaca and C. michoacanus from the state of Michoacan are described as new. A scanning electron micrograph of the stridulatory device of C. jacquiniae is presented. Illustrations of the male genitalia are provided. It is not obvious to what other genera Carvalhoisca may be related. Key Words: The genus described herein represents an- other new genus of the rich and diverse or- thotyline fauna found in the arid regions of southern Mexico. The two species of this rather unusual genus are found on plants of the genus Jacquinia Linnaeus, which is widely distributed through the New World tropics. Measurements are given in mm. Carvalhoisca n. gen. Characterized by the small size, minutely granulate surface, wide and sharply decli- vent head, rather quadrate pronotum, deep- ly incised cuneal fracture, and the presence of a stridulatory device involving the hem- elytron and hind femur. Body appearing minutely granulate, not shining, vestiture consisting primarily of scattered decumbent hairs. Head sharply declivous, almost as wide as width of pro- notum; vertex with posterior margin sharp- ly angulate, frons slightly rounded; clypeus weakly delimited from frons, recurved; ju- gum, clypeus, lorum, buccula, and gena small relative to size of head; eyes large, touching pronotum; antennal socket contig- Heteroptera, Miridae, Orthotylini, Carvalhoisca, new genus, new species uous with anteroventral margin of eye, length of antennal segments in ascending order I-IV-III-II, vestiture semidecumbent; rostrum extending beyond hind coxae. Pronotum flat; calli weakly delimited; collar extremely narrow, clearly delimited; anterior and posterior corners of pronotum angulate; lateral margins angulate, straight; posterior margin convex; mesoscutum cov- ered; scutellum convex; corium curved downward along costal margin; embolium only delimited basally, costal margin with well-developed stridulitrum (Fig. 2); cuneal fracture deeply incised, moderately deflect- ed; femora only slightly incrassate, with three setae or trichobothria longer than diameter of femur, hind femur with plectrum (Fig. 2); tibiae lacking erect spines or setae. Apex of abdomen of male somewhat pointed; parameres small. Type species: Carvalhoisca jacquiniae n. sp. We take great pleasure in naming this ge- nus in honor of Dr. J. C. M. Carvalho, Mu- seo Nacional, Rio de Janeiro, Brazil, whose contributions to our knowledge of the mirid fauna of the neotropics are unexcelled. 374 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON This distinctive genus exists at couplet 82 with the genera 4/mixia Reuter and Asery- mus Distant in Carvalho’s key to the mirid genera of the world (1955) but bears no par- ticular resemblance to either. Its relation- ship to other New World genera is not readi- ly apparent, but it is superficially similar to Adfalconia Distant. Due to the small size of the insect, the stridulatory device is difficult to see with an ordinary light dissecting microscope. This type of stridulatory apparatus has been re- ported as occurring in other mirid subfam- ilies. The host plant for these two mirid species are members of the genus Jacquinia (Theo- phrastaceae). These plants have stiff narrow leaves, the tips of which are acuminate, im- parting some protection for the bugs that feed rather openly among the leaves. Both nymphs and adults of the two species were taken. The chlorotic spots on the thick leaves caused by the feeding were readily apparent and were somewhat like the damage seen on orchids caused by members of the bry- ocorine genus Tenthecoris Scott or that caused by Caulotops Bergroth or Haltico- toma Townsend on Yucca Linnaeus. Approximately 50 species of Jaquinia are known from tropical America, including the West Indies. Nine species have been re- ported from Mexico. Inasmuch as the Car- valhoisca species described are from the northern areas of the plant distribution, it is likely that additional species of this mirid genus will be found. The crushed fruit of this plant has been reported (Standley 1923) to be widely used by Indians to stupefy fish and also as a medicinal plant. Carvalhoisca jacquiniae n. sp. (Figs. 1, 2) Male (measurements of holotype given first followed in parentheses by means and ranges, n = 20): Length, 1.88 (1.93, 1.74— 2.04); width, 0.96 (0.98, 0.90-1.04). Head length, 0.16 (0.14, 0.12—0.16); width through eyes, 0.68 (0.70, 0.68-0.74); vertex width, 0.30 (0.30, 0.28-0.30). Length of antennal segment I, 0.24 (0.23, 0.20-0.24); II, 0.72 (0.75, 0.70-0.82); III, 0.36 (0.39, 0.36-0.42); IV, 0.28 (0.30, 0.28-0.34). Pronotal length, 0.32 (0.33, 0.32-0.34); width, 0.76 (0.78, 0.74-0.84). Cuneal length, 0.36 (0.34, 0.32- 0.36); width, 0.22 (0.24, 0.22-0.24). General coloration black with append- ages pale yellowish white. Head black downward to level of antennal insertions, juga and area of clypeus between juga red- dish brown, lora and area of clypeus be- tween lora pale; antenna pale, faint reddish brown coloration on base of antennal seg- ment I; rostrum pale, dark fuscous at apex. Thorax, including scutellum and hemely- tron, dark fuscous to black, membrane of hemelytron paler along outer margin; bases of mid and hind coxae dark fuscous, re- mainder of legs pale. Abdomen dark fus- cous. Morphological characters are given for genus. Genitalia similar to those of C. mi- choacanus (Figs. 3-5). Female (means followed in parentheses by ranges, n = 20): Length, 1.96 (1.84—- 2.08); width, 1.03 (0.92-1.12). Head length, 0.17 (0.12-0.22); width through eyes, 0.74 (0.68—-0.78); vertex width, 0.35 (0.34—0.36). Length of antennal segment I, 0.22 (0.20- 0.24); II, 0.61 (0.54—-0.66); III, 0.34 (0.30- 0.36); IV, 0.30 (0.26-0.34). Pronotal length, 0.34 (0.30-0.36); width, 0.83 (0.74-0.88). Cuneal length, 0.32 (0.28-0.36); width, 0.26 (0.24—0.28). Similar to male in form and color. Holotype: 6, MEXICO: Oaxaca, 2.1 mi. nw. Totolapan, July 11-17, 1981, [D. S.] Bogar, [J. C.] Schaffner, [T. P.] Friedlander. Deposited in the collection of the Instituto de Biologia, Universidad Nacional Autono- ma de Mexico, Mexico, D.F. Paratypes: 24 6, 76 2, same data as ho- lotype: 4, 4 2, same locality as holotype, Au- gust 7, 1980; (J. CG] Schafiner and [ie -P5] Friedlander; 8 3, 43 2°, Oaxaca, 10 mi. e. Totolapan, elev. 4000 ft., July 20, 1987, [P. W.] Kovarik, [J. C.] Schaffner. Deposited in 375 VOLUME 97, NUMBER 2 ass EGA SS coaiiien SS i eens SNS nd ee ducal \ WSK bes ne Nites ~~ ee a” a eS ee eee CQsSsSRET See Ne ES “SSR Carvalhoisca jacquiniae, dorsal habitus. Fig. 1. 376 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the National Museum of Natural History, Washington, D.C.; in the collection of the Department of Entomology, Texas A&M University, College Station, Texas; in the J. C. M. Carvalho collection, Rio de Janeiro, R.J.; and in the collection of the Instituto de biologia, U.N.A.M., Mexico, D.F. The host plant for both nymphs and adults of this species is Jacquinia seleriana Urb. & Loes. Carvalhoisca michoacanus n. sp. (Figs. 3-5) Male (measurements of holotype given first followed in parentheses by means and ranges, n = 20): Length, 1.60 (1.60, 1.44- 1.66); width, 0.96 (0.94, 0.90-0.98). Head length, 0.14 (0.12, 0.10-0.14); width through eyes, 0.76 (0.75, 0.74-0.78); vertex width, 0.34 (0.33, 0.32-0.34). Length of antennal segment I, 0.22 (0.21, 0.20-0.22); II, 0.68 (0.65, 0.60-0.68); IIT, 0.38 (0.35, 0.34-0.40); IV, 0.26 (0.24, 0.24—0.28). Pronotal length, 0.34 (0.34, 0.34-0.36); width, 0.82 (0.81, 0.80-0.84). Cuneal length, 0.32 (0.32, 0.30- 0.34); width, 0.20 (0.21, 0.20-0.22). General coloration black with append- ages pale yellowish brown. Head black, juga and base of tylus reddish black, lora and apical half of tylus pale yellowish brown; antenna pale yellowish brown, apex dark fuscous; rostrum pale yellowish brown. Thorax including scutellum and hemelytron black, membrane of hemelytron black be- coming abruptly pale near margin; thoracic sterna reddish brown to reddish black; bases of mid and hind coxae dark fuscous, re- mainder of legs light yellowish brown. Ab- domen reddish brown to reddish black. Morphological characters as given for ge- nus. Genitalia as in Figs. 3-5. Female (means followed in parentheses by ranges, n = 20): Length, 1.72 (1.62- 1.84); width, 1.03 (1.00-1.06). Head length, 0.16 (0.14—0.20), width through eyes, 0.80 (0.78-0.82); vertex width, 0.40 (0.38-0.42). Length of antennal segment I, 0.19 (0.18- Fig. 2. Carvalhoisca jacquiniae, costal margin of hemelytron showing stridulittum and hind femur showing plectrum. 0.22); II, 0.52 (0.48-0.56); THI, 0.34 (0.32- 0.38); IV, 0.25 (0.24—0.30). Pronotal length, 0.36 (0.34—0.38); width, 0.87 (0.86-0.90). Cuneal length, 0.33 (0.32—0.34); width, 0.24 (0.22—0.26). Similar to male in form and color. Holotype: 6, MEXICO: Michoacan, 28.5 miles south of Nueva Italia, July 9, 1985, [R. W.] Jones, [J. C.] Schaffner. Deposited in the collection of the Instituto de Biologia, Universidad Nacional Autonoma de Mex- ico, D.F. Paratypes: 25 6, 54 2, same data as ho- lotype. Deposited in the National Museum of Natural History, Washington, D.C.; in the collection of the Department of Ento- mology, Texas A&M University, College Station, Texas; in the J. C. M. Carvalho collection, Rio de Janeiro, Brasil; and in the collection of the Instituto de Biologia, U.N.A.M., Mexico, D.F. Specimens of this species are nearly al- VOLUME 97, NUMBER 2 377 Figs. 3-5. ways shorter than those of C. jacquiniae. The males range in length from 1.44—-1.66 mm and the females 1.62—1.84 mm. The males of C. jacquiniae are 1.74—2.04 mm in length and females 1.84—2.08 mm. The head width of C. michoacanus is greater than that of C. jacquiniae. The male head width is 0.74-0.78 mm, that of the female, 0.78- 0.82 mm. In the case of C. jacquiniae, the male head width is 0.68-0.74 mm and the female, 0.68—-0.78 mm. The antennal seg- ment lengths are shorter on individuals of C. michoacanus. By using the head width divided by the length of antennal segment II, a ratio for easy separation of the 2 species can be derived. The males of C.jacquiniae have antennal segment II longer than the head width (1.12—1.23), whereas in the case of C. michoacanus the second antennal seg- Carvalhoisca michoacanus. 3. Vesica. 4. Left paramere. 5. Right paramere. ment is shorter than the head width (0.90- 0.97). Antennal segment II is 1.39-1.67 times as long as the head width in females of C. jacquiniae individuals but only 1.13- 1.28 as long for specimens of C. michoa- canus. We are unable to separate the species on the basis of the genitalia. The host plant for both nymphs and adults of this species is Jacquinia pungens A. Gray. ACKNOWLEDGMENTS We are grateful to Helga Bhacktar, Texas A&M University, College Station, for her preparation of the S.E.M. micrograph and to Dr. J. Heraty, USDA, Systematic Ento- mology Laboratory, Washington, D.C., for the habitus drawing. We also thank Dr. H. R. Burke, Texas A&M University, College Chel P= Station, for reviewing the manuscript. Our 378 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sincere thanks are also extended to Dr. P. A. Fryxell, University of Texas, Austin, for identifying the host plants. LITERATURE CITED Carvalho, J. C. M. 1955. Keys to the genera of the Miridae of the world (Hemiptera). Boletim do Mu- seu Paraense Emilio Goeldi 11: 1-151. Standley, P. C. 1923. Trees and shrubs of Mexico. Contributions from the U.S. National Herbarium 23: 1-1721. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 379-395 EUROPIELLA REUTER (HETEROPTERA: MIRIDAE): RECOGNITION AS A HOLARCTIC GROUP, NOTES ON SYNONYMY, AND DESCRIPTION OF A NEW SPECIES, EUROPIELLA CARVALHOI, FROM NORTH AMERICA RANDALL T. SCHUH, PER LINDSKOG, AND I. M. KERZHNER (RTS) Department of Entomology, American Museum of Natural History, New York, New York 10024, U.S.A.; (PL) Department of Entomology, Swedish Museum of Natural History, Stockholm S-104-05 Sweden; (IMK) Department of Insect Taxonomy, Zoological Institute, Academy of Sciences, St. Petersburg, Russia 199034. Abstract.—The subgenus Poliopterus Wagner of Plagiognathus Fieber is placed in syn- onymy with Europiella Reuter. The identity of Holarctic species is clarified. New com- binations are created because of generic synonymy, species are transferred from other genera, and many names are placed in synonymy. Most species for which hosts are known feed on Artemisia, a few feeding on other Asteraceae such as Chrysothamnus, Helichrysum, and Tanacetum, with two species being recorded from the Lamiaceae. Key Words: Reuter (1909) used the name Europiella for two species of phyline Miridae from North America. Since that time many ad- ditional American species have been placed in the genus. Wagner (1949) described the subgenus Plagiognathus (Poliopterus) to contain several species of Palearctic Phyli- nae related to P. albipennis (Fallén). Our individual research efforts have shown that many species placed in Europiella and Pla- giognathus are in fact congeneric, or in a few cases conspecific, that many species from North America that have previously been placed in Europiella do not belong there, and some Palearctic species placed in Pla- giognathus sensu stricto and Chlorillus ac- tually belong to Europiella. There is much confusion regarding the correct application of species-group names, including substan- tial synonymy. In the following pages we provide solutions to these problems and de- scribe a new species. Additional details on Heteroptera, Miridae, Europiella, new species, new synonymy, Holarctic the history of synonymy for the Nearactic fauna, as well as more detailed distributions of the species, can be found in Henry and Wheeler (1988). This paper is presented in honor of our long-time colleague and friend José Can- dido de Melo Carvalho. His influence on the study of the Miridae has been profound, because of his monumental world catalog and his unparalleled descriptive efforts, par- ticularly on the Neotropical fauna. We thank Thomas J. Henry and Michael D. Schwartz for reviewing the manuscript. Europiella Reuter Type species.—Agalliastes stigmosus Uhler 1893. Europiella Reuter 1909: 83 (n. gen., desc.): Knight 1968: 37 (key to spp.). Poliopterus Wagner 1949: 53 (n. subgen. of Plagiognathus, desc.). NEw SYNONYMY. 380 Diagnosis. — Relatively small, length 2-5 mm; some spp. sexually dimorphic, body form varying from elongate and nearly par- allel-sided to broadly ovoid, especially in females. Coloration varying from almost completely black to almost completely green or white, usually with a few dark markings at the bases of spines on the legs. Vestiture of dorsum with reclining simple setae and recumbent, woolly, sericeous setae; head weakly produced and not overlapping an- terior margin of pronotum. Male genitalia varying greatly in size, but pygophore al- ways large relative to total size of abdomen; right paramere truncate apically, with a more or less well-developed protrusion on either side, never lanceolate in form; vesica always with two apical spines, these sometimes elongate and broadened basally (Figs. 12, 13), or much shorter and variously acumi- nate and/or strongly curving (Figs. 6, 7, 9, 10, 15-17); secondary gonopore placed to the side of the chitinous bands of the vesica rather than between them and distinctly proximad of attachment of spines. Often breeds exclusively on Artemisia, occasion- ally on other Asteraceae, and rarely on other plant families. Discussion. — Wagner (1949) described the subgenus Poliopterus, with albipennis (Fallén) as the type. He consistently placed it in the genus Plagiognathus Fieber by vir- tue of its members having two elongate chi- tinous spines apically on the vesica and with the secondary gonopore rather distant from them. Within Plagiognathus, Wagner rec- ognized Poliopterus by the presence of only light colored setae on the dorsum. Wagner (1975) placed 13 species from the Mediter- ranean Region in Poliopterus; the majority of those for which hosts are known feed on species of Artemisia. Members of another assemblage of Pa- learctic species, richly represented in the Far East (Kerzhner 1988a) and including one European species, are green and dissimilar to albipennis in appearance. They were tra- ditionally placed in Plagiognathus sensu PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON stricto, but some species such as Plagiog- nathus alpinus, have been placed in Chlor- illus or Psallus by some authors. The struc- ture of the genitalia indicates that they should be placed in Europiella, as does com- parison with some North American species, which are similar in coloration. All but two of them feed on Artemisia. In the New World there are several spe- cies that are very closely related to albipen- nis, and a number of others that we treat as congeneric. Among these is stigmosa (Uhl- er), the type species of Europiella Reuter 1909. We are therefore treating Poliopterus Wagner as the junior synonym of Euro- piella. The male genitalia of stigmosa are shown in Figs. 12-14 and those of albipen- nis in Fig. 16. The generic limits of Plagiognathus are a subject that can be properly addressed only with a much more wide-ranging analysis of the Holarctic Phylini, something that we do not attempt in this paper. Because subgen- era have been used primarily in the Pale- arctic, a consistent treatment of the world fauna including the use of Palearctic sub- generic concepts would leave many species unplaced as to subgenus. It is for these rea- sons that we treat Europiella as a distinct genus. Knight (1968, 1969, 1970) diagnosed Eu- ropiella without examining the structure of the male genitalia. He included some spe- cies which are clearly congeneric with stig- mosa, the type. He also included many oth- er species (as enumerated by Henry and Wheeler 1988) which bear a superficial re- semblance to stigmosa and its congeners. These species are found in desert areas of western North America feeding primarily on Atriplex, Sarcobatus (Chenopodiaceae), and Lycium (Solanaceae), but nearly all of them belong to a separate lineage. Com- parison of preparations of the male genitalia of paratypes of most of these species and the genitalia of Megalopsallus atriplicis Knight, the type species of Megalopsallus Knight, indicates that most, if not all, of VOLUME 97, NUMBER 2 381 Figs. 1-17. Male genitalia of Europiella spp. 1-5. E. carvalhoi. 1. Vesica. 2. Detail of apex of vesica. 3. Obverse view of distal portion of vesica. 4. Right paramere. 5. Left paramere. 6-8. E. decolor (western North America). 6. Vesica. 7. Apex of vesica, rotated 90 degrees. 8. Right paramere. 9-11. FE. artemisiae (western North America). 9. Vesica. 10. Apex of vesica, rotated 90 degrees. 11. Right paramere. 12-14. E. stigmosa. 12. Vesica. 13. Obverse view of distal portion of vesica. 14. Right paramere. 15-17. Comparative views of apex of vesica of Holarctic species. 15. E. artemisiae (Europe). 16. E. albipennis (Europe). 17. E. decolor (Europe). 382 them are congeneric. We provide below a list of all names for which new combina- tions are formed in Megalopsallus. Some species which are clearly congeneric with stigmosa have often been placed in other genera. We have transferred all of those of which we are aware. In the Holartic there are Artemisia-feed- ing species placed in other genera of Phy- linae (e.g. species placed in Phyllopidea Knight) as well as in other subfamilies of Miridae. Critical examination of these spe- cies indicates that although the host pref- erence of Europiella spp. is helpful in be- ginning to establish a basis for their generic identity, it is often only with examination of the male genitalia that generic placement and specific identity can be determined with certainty. We have organized most of our treatment on geographical grounds, because the per- tinent literature is organized on that basis. IDENTITY AND SYNONYMY OF EUROPIELLA ALBIPENNIS AND TWO RELATED SPECIES Three closely related species of Europiella are widely distributed in the Palearctic, two of these also occurring in the Nearctic. No other species of the genus are Holarctic. All three of these species have been identified by various authors as Plagiognathus albi- pennis. We found that these taxa are reliably distinguished from one another by the form of the vesical appendages as seen in dorsal view (e.g. Figs. 15-17). The size and color are highly variable, but in some regions the species can be recognized by external ap- pearance. Europiella albipennis (Fallen), New ComBINATION Fig. “16 Phytocoris albipennis Fallén 1829: 107 (n. sp., desc.). Agalliastes albipennis: Fieber 1861: 311 (key); Puton 1873: 25 (syn.). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Agalliastes tibialis Fieber 1864: 228 (n. sp., desc.). Agalliastes lanuginosus Jakovlev 1875: 172 (nesp, desc): Plagiognathus albipennis var. tibialis: Ja- kovlev 1877: 279 (syn.). Plagiognathus albipennis: Reuter 1878: 175 (part; descr., variability, syn.). Plagiognathus collinus Wagner 1941: 249 (n. sp., desc., figs.). NEW SYNONYMY. Plagiognathus arenicola Wagner 1941: 252 (n. sp., desc., figs.). NEW SYNONYMY. Plagiognathus (Poliopterus) collinus: Wag- ner 1952a: 197 (key, desc., figs.). Plagiognathus (Poliopterus) arenicola: Wagner 1952a: 197 (key, desc., figs.); Wagner and Weber 1964: 413 (key, desc., figs.); Wagner 1975: 35 (key, desc., figs.). Plagiognathus (Poliopterus) lanuginosus: Josifov 1974: 14, 20 (restored from syn- onymy, list, host); Wagner 1975: 34 (key, desc., figs.). Distribution.— From southernmost Swe- den, southeastern Finland, and Leningrad Province, Russia, at least to Bulgaria and from France east at least as far as East Ka- zakhstan. Hosts.— Living exclusively on Artemisia campestris in Northern Europe; records of Plagiognathus lanuginosus from A. mariti- ma in southern Europe probably refer to E. decolor. Notes.—In this species the dark spots on the hind tibia are usually larger than in re- lated species and the venter 1s black in both sexes, even though the dorsum may be pale. The apex of the vesica is shown in Fig. 16. Fallén (1829) described Phytocoris albi- pennis from four specimens, including male(s) and female(s), all collected by Zet- terstedt in July. Type locality: Sweden, Skane Prov. (Scania), Esper6d. Extant material qualifying as syntypes is present only in the collection of Fallén, not in the collection of Zetterstedt, both collections deposited in the Zoological Museum, Lund University. It consists of one male labelled “P. al/bipennis VOLUME 97, NUMBER 2 [male symbol]” and a small blue square, and one female labelled ‘““P. albipennis [female symbol]”’ plus a few illegible characters, and ““Mellby 12 jun.” on the underside of the label. The labels are in Zetterstedt’s hand- writing, and the small blue square is known to signify that Zetterstedt collected the spec- imen at Esperéd, Mellby Parish. Among these specimens is pin without any speci- men with Fallén’s label “P. a/bipennis [male symbol] [female symbol] Esperéd.” According to Wagner (1941: 256), F. Os- siannilsson informed him that the type was no longer present in the collection of Fallén, but rather that only an empty pin remained. Ossiannilsson in a letter to Lindskog (Jan. 15, 1982) explained that his reply to Wagner was based on the opinion of museum au- thorities that in general only specimens car- rying Fallén’s labels could be considered as his types. It is known that after the death of Fallén in 1830 his collection passed to the possession of Zetterstedt, who notori- ously filled in any missing species with spec- imens from his own collection. Ossiannils- son wrote (in litt.) “In the present case this was apparently fully justified, as I should have realized if time had been taken to look more closely on Zetterstedt’s label and Fal- lén’s text. Fallén described a/bipennis based on four specimens. He probably retained two specimens for his own collection which later were lost somehow, and returned two to the collector Zetterstedt, who in turn placed these in Fallén’s collection after he acquired it. These two specimens should ac- cordingly be considered as syntypes.”’ As Fallén indicated that the types were collected in July, and because Zetterstedt in his handwritten notes in his copy of Fallén’s (1829) work (now in the library of the Zoo- logical Institute, St. Petersburg), indicated that he collected the species at “‘I[ocis] aridis Esp[eroed] 18 jul. freq[uens],” it seems doubtful that the female labelled 12 June belongs to the syntype series although no such doubt exists for the male. We accept these arguments and designate 383 the male as lectotype of P. albipennis and have labelled it accordingly. Wagner (1941) stated that the type spec- imens were presumably lost and designated from his collection ‘“Neotypen,” saying nothing about their number, sex, and lo- cality. Wagner’s “‘neotypes” are not con- specific with the true types (see below) and Wagner’s nomenclatural act is invalid be- cause more than one specimen was desig- nated as the neotype. From the examination of Fallen’s speci- mens we have determined that the name albipennis should be ascribed to a species which is not the most common in the Pa- learctic and which does not occur in North America, in spite of many indications in the literature to the contrary. Agalliastes tibialis was described from Sarepta (now Krasnoarmeysk, part of Vol- gograd) from Frey-Gessner’s collection. The type specimens were apparently examined by Puton and Reuter, and Reuter (1878, pl. 4, fig. 8) published Fieber’s figures of a fe- male. We cannot locate the type(s). Judging from the large, black, tibial spots noted in the original description and the figure, and the pale dorsal surface and black abdomen, this species in a almost certainly identical with albipennis. Agalliastes lanuginosus was described from specimens collected at Sarepta by Christoph and at Akhtubinskaya steppe by Becker. The only known syntype is a badly damaged specimen (probably female) from Sarepta in the Zoological Institute, St. Pe- tersburg. Its remnants include the head with antennae, prothorax, and all legs. Judging from these parts and from the original de- scription, the species is identical with al- bipennis. Plagiognathus arenicola was described from Germany and Poland and P. collinus from Poland. We examined the types and paratypes of both species in the Zoological Museum Hamburg and paratypes in other collections. Some paratypes were dissected 384 for comparison of the male genitalia, and these are conspecific with albipennis. Reuter (1878: 81, 82) described several varieties of P. albipennis, assigning them Greek letters. Reuter did not indicate spec- imens for his varieties and did not label varieties in his collections. Stichel (1934: 282) named these varieties assmanni, al- bella, beckeri, and antennaria. The identity of these names cannot be clarified at pres- ent, and it is clear that Stichel did not base his names on specimens which he himself examined. Europiella decolor (Uhler) Figs. 6-8, 17 ? Phytocoris pallidulus Dahlbom 1851: 211 (n. sp., descr.). Questionable NEw SYNONYMY. Agalliastes decolor Uhler 1893: 380 (n. sp., desc.). Agalliastes apiatus Uhler 1895: 53 (n. sp., desc.). NEw SYNONYMY. Agalliastes signatus Uhler 1895: 55 (n. sp., desc.). NEw SYNONYMY. Chlamydatus bakeri Bergroth 1898: 35 (n. n. for Agalliastes signatus Uhler). Plagiognathus albipennis var. extrema Reuter 1901: 187 (n. var., descr.). NEw SYNONYMY. Chlamydatus uhlerianus Kirkaldy 1909: 390 (unnecessary n. n. for Agalliastes signatus Uhler). Plagiognathus decolor: Reuter 1909: 81 (disc.). Europiella decolor: Van Duzee 1916: 47 (list). Chlamydatus apiatus: Van Duzee 1917: 417 (n. comb.). Psallus waldeni Knight 1923: 468 (n. sp.). NEw SYNONYMY. Psallus bakeri: Knight 1941: 43 (key, desc.). Plagiognathus (Poliopterus) litoralis Wag- ner 1949: 53 (n. sp., desc., figs.); Wagner 1952a: 199 (key, desc., figs.). NEw SYNONYMY. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Plagiognathus extremus: Wagner 1954: 77 (n. status, descr.). Plagiognathus (Poliopterus) litoralis f. abro- tani Wagner 1949: 53 (n. form, desc.). NEw SYNONYMY. Europiella bakeri: Carvalho 1955: 227 (n. comb.); Knight 1969: 86 (disc., redesc., host). Plagiognathus (Poliopterus) abrotani: Wag- ner and Weber 1964: 413 (new status, key, desc., figs.). Psallus artemisicola Knight 1964: 149 (n. sp., desc., host). NEw SYNONYMY. Europiella nigricornis Knight 1968: 40 (n. sp., desc., host). NEW SYNONYMY. Plagiognathus (Poliopterus) larae Kerzhner 1978: 46 (n. sp., desc., figs.). NEw SYNONYMY. Psallus albipennis (not Fallén 1829): Whee- ler and Hoebeke 1982: 696 (n. comb., syn., figs.). Distribution.—Broadly — distributed throughout the Holarctic. Hosts.—Neararctic hosts include Arte- misia californica, A. campestris, A. dracun- culus, A. filifolia, A. ludoviciana, A. nova, A. tridentata, and Chrysothamnus sp. In the western Palearctic known from Artemisia absinthium, A. maritima, and A. abrotani (in botanical gardens). Kurile Islands spec- imens are from A. schmidtiana, belonging to the ‘“‘Maritimae”’ series. Notes.—The name decolor (Uhler) has previously been applied only to the North American fauna. Our studies indicate, how- ever, that the taxon to which this name ap- plies also occurs in the Palearctic. Uhler (1893) indicated in his discussion of decolor that he examined four specimens from American Fork, Utah, as well as larger specimens collected near Los Angeles, Cal- ifornia. We located in the National Museum of Natural History, Washington, D.C., a single female specimen from American Fork that seems to pertain to material originally examined by Uhler. In addition to the data noted above it bears the labels “COTY PE VOLUME 97, NUMBER 2 (by Uhler) Agalliastes decolor Uh.” [ap- parently affixed by H. H. Knight], and “Co- type No. 52835 U.S.N.M.” Uhler (1895) described Agalliastes apia- tus from specimens collected at Fort Col- lins, Manitou, and Steamboat Springs, Col- orado. He indicated that it also occurred in Kansas. The collections of the USNM con- tain 10 specimens associated with the apia- tus identification. One of these was labelled as a cotype by H. G. Barber. It is clearly congeneric with decolor. Uhler (1895) described Agalliastes sig- natus from a single male specimen from Manitou, Colorado. We were not able to find any specimens labelled as signatus after careful checking of the USNM collections. Bergroth (1898) proposed the replacement name bakeri for the preoccupied signatus. Our use of the name bakeri is based on spec- imens identified by H. H. Knight, which are clearly conspecific with decolor. We there- fore treat bakeri as a junior synonym. Wheeler and Hoebeke (1982) correctly synonymized Psallus waldeni Knight with what they called a/bipennis (Fallén), but our examination of specimens and literature in- dicates that the senior synonym is in fact decolor (Uhler), not albipennis (Fallén). We examined type specimens of Plagiog- nathus litoralis (described from Germany and the Netherlands), P. abrotani (from the Hamburg Botanical Gardens), and P. larae (from Kurile Islands), as well as many ad- ditional specimens, and found these nom- inal taxa to be conspecific with decolor from North America. Plagiognathus albipennis var. extrema Reuter 1901 (p. 187) was described from several specimens (length 1.75-2.00 mm) collected by J. Sahlberg at ““Constantinov- skaya”’ (Konstantinovskaya on River Chu, about 20 km N of Beshkek, Kirgizia). Wag- ner (1954) examined a female from the Uni- versity Zoological Museum, Helsinki la- belled ‘““Tokmak” (about 60 km E of Besh- kek). He designated it as lectotype and con- sidered Plagiognathus extremus as a valid 385 species. Additional single male and female specimens identified as this species were found in the collections in Helsinki together with the lectotype. All of them are labelled “Tokmak,” but the male has Sahlberg’s code label indicating that it was collected at Con- stantinovskaya. Other cases of confusion in labelling specimens from these two locali- ties were also encountered. All specimens belong to E. decolor. The identity of Phytocoris pallidulus Dahlbom, 1851 is unclear. Reuter (1878) placed this name in synonymy with P. al- bipennis. Wagner (1941) treated pallidulus as a valid species, but later (Wagner 1952a) considered it as a pale variety of P. albi- pennis sensu Wagner. Phytocoris pallidulus was described from two specimens (appar- ently a male and a female) collected by Dahlbom at Stensuga, Gotland Island, Swe- den, from Umbelliferae. The type speci- mens are apparently lost; according to R. Danielsson (in litt.), Zoological Museum, Lund University, they cannot be located in the collections of Dahlbom, Thomson, or other pertinent collections. Dahlbom’s orig- inal description is inadequate for identifi- cation and was apparently based on teneral specimens. At least the description can hardly be referred to FE. albipennis or E. artemisiae (albipennis sensu Wagner). Lindskog was unable to find either of these species from Gotland Island either in Swed- ish collections or through intensive personal collecting. The only species of Europiella found on Gotland Island is E. decolor which is abundant there on Artemisia absinthium. But, as Dahlbom’s description may refer to some other small Miridae, especially to ten- eral specimens, we prefer to consider Phy- tocoris pallidulus as a possible synonym of E. decolor rather than treat it as a valid species. Europiella artemisiae (Becker), NEw COMBINATION Figs. 9-11, 15 Capsus artemisiae Becker 1864: 487 (n. sp., desc.). 386 Plagiognathus solani Matsumura 1917: 432 (n. sp., desc.). NEw SYNONYMY. Plagiognathus albipennis var. obscura Sahl- berg 1920: 167 (n. var., desc.). NEW SYNONYMY. Plagiognathus albipennis (not Fallén 1829): Wagner 1941: 248 (desc., figs.). Plagiognathus (Poliopterus) albipennis (not Fallén 1829): Wagner 1952a: 198 (key, desc., figs.); Kerzhner 1964: 761 (key, figs.); Kerzhner 1988a: 853 (key, figs.); Li and Zheng 1991: 90 (key, dist., figs.). Plagiognathus (Poliopterus) gracilis Wagner 1956b: 74 (n. sp., desc., figs.). NEw SYNONYMY. Distribution. — This species is widely dis- tributed in the Palearctic, in the north to Scandinavia and Chukotka. Our examina- tion of collections from Canada and the Western United States indicates that al- though this species has not previously been recorded from North America, it ranges from Alaska south to montane areas of the northwestern United States (detailed local- ity records to be published separately). Host. — Palearctic hosts include Artemisia vulgaris, the related A. montana, and A. rub- ripes, aS well as other species. The only known Nearctic host is Artemisia ludovi- clana. Notes. — This is the most common species of Europiella in the Palearctic. Wagner (1941) designated a neotype for what he considered to be Plagiognathus albipennis. But because Wagner misidentified the spe- cies under discussion as Phytocoris albipen- nis Fallén, his neotype(s) belongs to another species, the oldest available name for which appears to Capsus artemisiae Becker. Capsus artemisiae was described from specimens collected by Becker in 1862 at Sarepta in southern Russia, from a plant he identified as Artemisia fragrans. This plant species does not grow in the Lower Volga region, so probably Becker collected from another species of Artemisia. In the Zoo- logical Institute, St. Petersburg, there are six specimens which were received from Beck- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON er: three females (received in 1866), two males (1872), and one female (1874); one additional male collected by Becker was re- ceived with the Jakovlev collection. All males belong to artemisiae (Becker) (albi- pennis sensu Wagner). The male bearing the number 13737 is designated here as lecto- type. In the Naturhistorisches Museum Wien, there are two males received from Becker in 1870 and one female received in 1869. They were examined by Reuter. One male belongs to artemisiae, the other to al- bipennis. Plagiognathus solani Matsumura was de- scribed from Sakhalin and Hokkaido. The name was synonymized with albipennis sensu Wagner by Miyamoto (1977) and should put in synonymy with artemisiae (Becker). Also the varietal name obscura (Sahlberg) given to dark specimens from Finland should be placed here in synonymy. Plagiognathus gracilis was described from Austria and Croatia. The type series was examined by us and paratypes dissected for study of the male genitalia, indicating that gracilis is a junior synonym. NEARCTIC SPECIES Europiella angulata (Uhler) Maurodactylus angulatus Uhler 1895: 53 (n. sp., desc.). Europiella angulata: Knight 1968: 41 (n. comb., disc., dist., host). Europiella yampae Knight 1968: 43 (n. sp., desc., host). NEw SYNONYMY. Distribution.— Montane western United States. Hosts.—Artemisia arbustorum, A. dra- cunculus, A. ludoviciana, A. tridentata (As- teraceae). Notes. — Uhler (1895) mentioned having examined “Only one specimen ... a male ...” [of angulatus]. The specimen here rec- ognized as the holotype of angulata, in the USNM, fits Uhler’s original description, and bears the additional label ““Maurodactylus angulatus Uhl.” Examination of the male genitalia of the VOLUME 97, NUMBER 2 holotype of angulatus and paratype males of E. yampae Knight from the type locality indicates that the two are synonymous. Europiella artemisiae (Becker), see above. Europiella carvalhoi Schuh, NEw SPECIES Figs. 1-5 Diagnosis.— Recognized by the light col- oration of the dorsum with pale simple setae and weakly-flattened, recumbent, sericeous setae, the elongate oval body shape, anten- nal segment one black, segment two black basally and progressively lighter distally, and the form of the male genitalia, the vesica with two relatively short apical spines of nearly equal length, the more ventral spine with a small notch at the apex. Europiella carvalhoi is most similar among described species in body form and coloration to E. unipuncta Knight which has more differ- ently formed vesical spines and E. signi- cornis Knight which has totally pale anten- nae. Description.— Male: Dorsum, including membrane pale, white to tan, tinged with green; neck (when visible) castaneous; tho- racic pleuron and venter and pregenital ab- dominal segments weakly infuscate, pygo- phore somewhat darker; antennal segment one black, segment two black proximally, progressively lighter distally, segments three and four weakly infuscate; legs weakly in- fuscate, tibiae narrowly black at femoral ar- ticulation, tibial spines black with black bases; femora with a few diffuse scattered infuscate areas. Dorsum densely clothed with reclining pale simple setae and weakly flattened, re- cumbent, sericeous setae. Body elongate ovoid; frons bulging; eyes globular, touching anterior margin of pro- notum; posterior margin of vertex sinuous. Pygophore very large, occupying well more than half of abdomen; vesica with two relatively short apical spines of nearly equal length, the more ventral with a small notch at the apex (Figs. 1-3); left paramere as in Fig. 5; right paramere as in Fig. 4. 387 Measurements: Apex tylus Inter- Width Length —cuneal Width — ocular pro- antennal fracture head space notum segment 2 M 8 1.87-2.11 0.67-0.73 0.41-0.47 0.86-0.96 0.60-0.70 F 8 1.65-2.07 0.66-0.74 0.43-0.49 0.82-1.00 0.50-0.68 Holotype.—male, USA: Wyoming: Big Horn Co.: 27 mi. W Burgess Jct. on Rt. 14, 7700 ft., Aug. 12, 1986, Schuh, Schwartz, and Stonedahl; Artemisia tridentata Nutt. (Asteraceae); deposited in the AMNH. Paratypes.— 3 males, 9 females, same data as holotype (AMNH, OSU, USNM). USA: Nevada: Washoe Co.: 7 mi. W of Vya to- ward Decarville, 1800 m, July 2, 1979, R. T. Schuh and B. M. Massie, ex Artemisia sp. (not tridentata) (Asteraceae) (AMNH), 5 males, 9 females. Oregon: Jackson Co.: Ashland, 6500 ft., September 24, 1968, Oman (OSU), 4 males, 13 females; 1 mi. below summit of Mt. Ashland, 6500 ft., September 24, 1968, J. D. Lattin (OSU), 7 males, 7 females. Umatilla Co.: 2 mi. W Tollgate, 4600 ft., August 4, 1986, Schuh, Schwartz, and Stonedahl, Artemisia triden- tata Nutt. (Asteraceae) (AMNH, USNM), 32 males, 59 females. Wheeler Co.: 2 mi. W Mitchell on Rt. 26, June 22, 1979, M. D. Schwartz, G. M. Stonedahl, ex Artemisia tridentata (AMNH), 7 males, 20 females; Mitchell, June 22, 1979, R. T. Schuh, ex Artemisia tridentata wyomingensis (AMNH), 2 males, 8 females. Hosts.—Artemisia tridentata, Artemisia tridentata wyomingensis (Asteraceae). Europiella consors (Uhler), NEw COMBINATION Maurodactylus consors Uhler 1895: 53 (n. Spe desc,): Europiella fuscicornis Knight 1969: 82 (n. sp., desc., host). NEw SYNONYMY. Europiella basicornis Knight 1970: 230 (n. sp., desc.). NEW SYNONYMY. Distribution.— Western North America. Hosts. — Artemisia dracunculus, A. filifol- ia, A. ludoviciana ludoviciana, A. tridentata, 388 Chrysothamnus nauseosus, C. parryi, C. vis- cidiflorus, C. sp. (Asteraceae). Notes. Uhler (1895) described consors on the basis of ‘“‘one specimen, a male... from Colorado.” We have examined a spec- imen in the USNM which fits Uhler’s orig- inal description, and bears the label ““Mau- rodactylus consors col. Uhler.” We therefore have assumed that it is the specimen orig- inally examined by Uhler. Dissections of specimens which are almost certainly con- specific indicate that this is actually a spe- cies of Europiella rather than belonging to the European Maurodactylus Reuter, and it is therefore transferred. Examination of the holotypes of Euro- piella basicornis Knight and E. fuscicornis Knight, a large amount of additional ma- terial, and many dissections of the male gen- italia, indicate that these two nominal spe- cies represent the same species and are both junior synonyms of consors. Europiella decolor (Uhler), see above. Europiella pilosula (Uhler) Atomoscelis pilosulus Uhler 1893: 377 (n. sp., desc.). Tuponia subnitida Uhler 1895: 45 (n. sp., desc.). NEW SYNONYMY. Psallus pilosulus: Van Duzee 1915: 118 (list, n. comb.). Microphylidea pallens Knight 1968: 29 (n. sp., desc.). NEw SYNONYMY. Europiella pilosula: Knight 1968: 44 (n. comb., note). Europiella flavicornis Knight 1969: 82 (n. sp., desc., host). NEw SYNONYMy. Europiella pallida Knight 1969: 83 (n. sp., desc., host). (Syn. by Stonedahl 1990:79). Europiella albata Knight 1969: 85 (n. sp., desc.). NEW SYNONYMY. Distribution.—Interior western United States. Hosts.— Artemisia filifolia, A. tridentata (Asteraceae). Notes. — Uhler (1893) described Atomos- celis pilosulus on the basis of ‘‘Several spec- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON imens. . . collected on Bigelovia near Amer- ican Fork, June 22.’ We have located in the USNM 2 specimens, a male and a female, that bear the correct data. A third specimen, a female, was actually a misidentified ex- ample of unipuncta Knight. Comparison of the male genitalia of pi- losula with those of the holotype of Micro- phylidea pallens Knight indicates that the two are synonymous. Comparison of the male genitalia of pi/osula with those of to- potypic males collected on the same date as the holotype of Europiella albata Knight in- dicates that the two are synonymous. Com- parison of pilosula with E. flavicornis Knight indicates that the two are synonymous. Uhler (1895) described Tuponia subniti- da from Steamboat Springs, Colorado, on the basis of two specimens. We searched the collections of the USNM, but were unable to find any specimens which appeared to have been examined by Uhler. We did find, however, specimens from Ft. Garland, Col- orado, collected by H. H. Knight and la- belled by him as “Tuponia subnitida Uhler, compared with type.” These specimens agree closely with the Uhler’s description of sub- nitida with the exception of the fact that antennal segment one is not “‘blackish”’ as indicated by Uhler, but rather only weakly brown. All other attributes, including size, shape, and distribution agree, however, and we are therefore treating subnitida as a syn- onym of pilosula. Europiella signicornis Knight Europiella signicornis Knight 1969: 84 (n. Sp2)dese: host): Distribution. — Northern Arizona. Host.— Artemisia sp. (Asteraceae). Europiella stigmosa (Uhler) Figs. 12-14 Agalliastes stigmosus Uhler 1893: 379 (n. Sp. desc.) Europiella stigmosa: Reuter 1909: 84 (disc.); Knight 1968: 43 (in part, dist., host). VOLUME 97, NUMBER 2 Distribution.— Western United States. Hosts.—Artemisia californica; Artemisia tridentata (Asteraceae). Notes. —In his original description, Uhler said that ““Specimens [of stigmosa] were col- lected at American Fork, June 22.” We lo- cated three specimens in the USNM which bear these data. One of these, a female, bears an identification label which appears to have been written by Uhler as well as a “lecto- type” label affixed by H. G. Barber, al- though we can find no evidence that Barber ever published this designation. The other two, both males, were in the collection of H. H. Knight and bear his identification la- bels, but there is no direct evidence that they are part of the material examined by Uhler. We have not designated a lectotype, but nei- ther do we consider the identity of stigmosa to be in question. Europiella umbrina Reuter Europiella umbrina Reuter 1909: 85 (n. sp., desc.). Europiella similis Knight 1969: 81 (n. sp., desc., host). NEw SYNONYMY. Europiella stigmosa: Knight 1968: 43 (mis- identification). Distribution.— Western United States. Hosts. — Artemisia californica, A. nova, A. tridentata (Asteraceae). Notes.— Henry and Wheeler (1988) listed umbrina Reuter as a synonym of stigmosa. Comparison of the types of stigmosa, um- brina, and similis Knight, and the male gen- italia of these nominal species, indicates that umbrina is distinct from stigmosa, but that similis is synonymous with umbrina. Europiella unipuncta Knight Europiella unipuncta Knight 1968: 44 (n. sp., desc., host). Distribution.—Interior western United States. Host. — Artemisia tridentata (Asteraceae). 389 Species transferred from Europiella to Megalopsallus Knight 1927 albipubescens Knight 1968 (Europiella), NEw ComBINATION arizonae Knight 1968 (Europiella), NEw COMBINATION balli Knight 1968 (Europiella), NEw COMBINATION brevicornis Knight 1968 (Europiella), NEw COMBINATION franseriae Knight 1969 (Europiella), NEw COMBINATION grayiae Knight 1968 (Europiella), NEw COMBINATION humeralis Van Duzee 1923 (Sthenarus), NEw CoMBINATION knowltoni Knight 1970 (Europiella), NEw COMBINATION lycii Knight 1968 (Europiella), New COMBINATION montanae Knight 1968 (Europiella), NEw COMBINATION monticola Knight 1968 (Europiella), NEw COMBINATION multipunctipes Knight 1970 (Europiella), New COMBINATION nicholi Knight 1968 (Europiella), NEw COMBINATION nigrofemoratus Knight 1968 (Europiella), NEw COMBINATION punctipes Knight 1968 (Europiella), NEw COMBINATION rubicornis Knight 1968 (Europiella, NEw COMBINATION rufiventris Knight 1968 (Europiella), NEw COMBINATION sarcobati Knight 1969 (Europiella), NEw COMBINATION sparsus Knight 1969 (Europiella), NEw COMBINATION stitti Knight 1968 (Europiella), NEw COMBINATION viridiventris Knight 1968 (Europiella), NEw COMBINATION PALEARCTIC SPECIES Europiella albipennis (Fallén), see above. 390 Europiella alpina (Reuter 1875), NEW COMBINATION Plagiognat/iiis alpinus Reuter 1875: 56 (n. sp., descr.); Kerzhner 1964: 761 (key, figs.). Psallus pallidus Reuter 1880: 24 (n. sp., descr.) (syn. by Wagner 1958). Plagiognathus alpinus f. nigrescens Stichel 1934: 279 (n. form, key). Plagionathus alpinus f. simplex Stichel 1956: 332 (n. form, key). Psallus (Psallus) alpinus: Wagner 1956a: 298 (disc., figs.); Wagner 1958: 325 (syn.). Plagiognathus (Chlorillus) alpinus: Ker- zhner 1962: 384 (disc., figs.). Chlorillus alpinus: Wagner 1963b: 155 (disc.,); Wagner 1975: 13 (desc., key, figs.). Distribution.—Europe, western Palearc- tic Asia. Contrary to Wagner (1975), this species is not recorded from North Amer- ica. Host:— Mentha aquatica (Lamiaceae). Europiella artemisiae (Becker), see above. Europiella canoflavida (Qi and Nonnaizab), NEw COMBINATION Plagiognathus (Poliopterus) canoflavidus Qi and Nonnaizab 1993: 31 (n. sp., desc., figs.). Distribution. —China: Inner Mongolia. Note.—We have not examined speci- mens of this species. It is probably synon- ymous with either FE. decolor or E. arte- misiae, based on the published figures of the male genitalia. Europiella decolor (Uhler), see above. Europiella herbaalbae (Wagner), NEw COMBINATION Plagiognathus (Poliopterus) herbaalbae Wagner 1969a: 13 (n. sp., desc., figs.); Wagner 1975: 29 (desc., key, figs.). Distribution. — Algeria, Libya, Tunisia. Host.—Artemisia herba-alba (Astera- ceae). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Europiella gilva (Kulik), NEw COMBINATION Plagiognathus gilvus Kulik 1965: 155 (n. sp., desc., figs.); Kerzhner 1979: 54 (key, syn., figs.); Kerzhner 1988a: 855 (key, figs.). Chlorillus pilosus Wagner 1969b: 34 (n. sp., desc., figs.). Distribution.—Russia (Primorsk Terri- tory). Records from Korea (Josifov and Kerzhner 1972) refer to E. livida. Host.— Rabdosia excisa (Lamiaceae). Europiella kiritschenkoi (Kulik), NEw COMBINATION Plagiognathus kiritschenkoi Kulik 1975: 587 (n. sp., desc., figs.); Kerzhner 1979: 55 (key, figs.); Kerzhner 1988a: 855 (key, figs.); Li and Zheng 1991: 90 (key, distr., figs.). Distribution.—Russia (Primorsk Terri- tory); China (Heilongjiang). Host.— Artemisia sp. (Asteraceae). Europiella leucopus (Kerzhner), New ComMBINATION Plagiognathus leucopus Kerzhner 1979: 50 (n. sp., desc., figs.); Kerzhner 1988a: 854 (key, figs.); Li and Zheng 1991: 90 (key, distr., figs.). Distribution.—Russia (Primorsk Terri- tory); China (Heilongjiang). Europiella livida (Reuter), NEw COMBINATION Plagiognathus lividus Reuter 1906: 73 (n. sp., desc.); Kerzhner 1979: 51 (part; distr., figs.); Kerzhner 1988a: 855 (key, figs.); Li and Zheng 1991: 91 (key, distr., figs.). Distribution.—Russia (Amur Province; Primorsk Territory); Korea; eastern China (to Sichuan). Host.—Artemisia gmelinii (Asteraceae). VOLUME 97, NUMBER 2 Europiella lividella (Kerzhner), NEw COMBINATION Plagiognathus lividellus Kerzhner 1979: 51 (n. sp., desc., figs.); Kerzhner 1988a: 855 (key, figs.); Li and Zheng 1991: 90 (key, distr., figs.). Distribution.— Russia (Far East); China (Heilongjiang; Fujian, Ningxia). Hosts.—Artemisia montana, A. rubripes, and A. vulgaris (Asteraceae). Europiella miyamotoi (Kerzhner), NEw COMBINATION Plagiognathus lividus (not Reuter 1906): Miyamoto 1969: 90 (desc., figs.); Ker- zhner 1979: 54 (part). Plagiognathus miyamotoi Kerzhner 1988a: 955 (n. sp., key, figs.); Kerzhner 1988b: 64 (desc.). Distribution. — Russia (Sakhalin and Ku- rile Islands); Japan. Host.—Artemisia montana (= gigantea) (Asteraceae). Europiella moesta (Reuter), NEw COMBINATION Plagiognathus albipennis var. moesta Reu- ter 1906: 75 (n. var., desc.). Plagiognathus (Poliopterus) moestus: Wag- ner 1954: 75 (n. stat., desc., figs.); Li and Zheng 1991: 90 (key, distr., figs.). Distribution. — China: Sichuan. Collected at high altitudes. Host.— Unknown. Note.—This species is very close to E. artemisiae and does not differ from it in the structure of the male genitalia, except in size. It is, however, markedly larger, very dark, and strongly shining. Europiella nigrocunealis (Putshkov), NEw COMBINATION Plagiognathus (Poliopterus) nigrocunealis Putshkov 1975: 283 (n. sp., desc.). Distribution.— Armenia; Azerbaijan. Host.— Tanacetum vulgare (Asteraceae). 39] Europiella ovatula (Wagner), NEw COMBINATION Plagiognathus (Poliopterus) ovatulus Wag- ner 1952b: 41 (n. sp., desc., figs.); Putsh- kov 1971: 35 (dist., host); Wagner 1975: 31 (key, desc., figs.). Distribution. — Croatia; Ukraine. Hosts.— Helichrysum angustifolium, H. arenarium (Asteraceae). Note.—This species does not differ from E. artemisiae in the structure of the aede- agus, but shows marked differences in size and color, and because the host plants are also different, we regard it as a separate spe- cies. It is closely related to the less well- known E. tomentosa and possibly synony- mous with it. Europiella strawinskti (Sienkiewicz), NEw COMBINATION Plagiognathus (Poliopterus) strawinskii Sienkiewicz 1986: 373 (n. sp., desc., figs.). Distribution. — Romania. Note.—Probably a synonym of E. deco- lor. Europiella strigifemur (Wagner), NEw COMBINATION Europiella (Poliopterus) strigifemur Wagner 1964: 70 (n. sp., desc., figs.); Wagner 1975: 28 (desc., key, figs.). Distribution. — Libya. Host.— Unknown. Note.— We examined only female speci- mens of this species. They clearly belong to the group of species closely related to Eu- ropiella albipennis. Europiella tomentosa (Reuter), NEw COMBINATION Plagiognathus tomentosus Reuter 1888: 236 (n. sp., desc.). Plagiognathus (Poliopterus) ovatulus (not 392 Wagner 1952b): Wagner and Weber 1964: 415 (desc.). Plagiognathis (Poliopterus) tomentosus: Wagner 1975: 31 (key, desc.). Distribution.—Southern France; Sicily. Host.— Helichrysum spp. (Asteraceae). COMMENTS ON ADDITIONAL PALEARCTIC SPECIES Psallus argyrotrichus Fieber 1861. This taxon was described from Germany and Spain. Wagner (1963a) examined four specimens from Spain belonging to the col- lections of Meyer-Diir (now in the AMNH) identified as P. argyrotrichus, designating one of them as the lectotype. He transferred argyrotrichus to Plagiognathus (Poliopterus) and placed P. decolor Lindberg in synonymy with it. We re-examined the material stud- ied by Wagner and have determined that the specimens are actually Compsidolon crotchi (Scott). Contrary to what Wagner (1963a) indicated, the identification label attached to the specimens is written by Meyer-Diir, not by Fieber. Although all of the specimens are badly rubbed, they have remnants of black setae on the hemelytra, which contradicts Fieber’s original descrip- tion. It is clear that the specimens examined by Wagner do not belong to the type series of Fieber and were misidentified. The iden- tity of P. argyrotrichus needs clarification. Plagiognathus bicolor (Jakovlev 1880). This species, the only representative of Plagiognathus (Zophocnemis) Kerzhner 1962, resembles some species of Europiella in coloration and size, but male genitalia are typical of Plagiognathus sensu stricto, and we therefore place it in that genus. Plagiognathus decolor Lindberg 1934. This species was described on the basis of four females from Spain (Lindberg 1934). Wagner (1963a) placed the taxon in syn- onymy with Psallus argyrotrichus Fieber and transferred argyrothichus to Plagiognathus (Poliopterus). The synonymy, taxonomic PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON position, and consequent possible second- ary homonymy of P. decolor Lindberg with Europiella decolor (Uhler) need re-exami- nation. See also notes under Psallus argy- rotrichus. Plagiognathus flavipes Reuter 1875. This species is known from southwestern Europe, living on Lonicera. Wagner (1952a) placed it in Plagiognathus (Poliopterus). We examined about 10 specimens, including some from Corsica, the type locality, iden- tified by Puton. The structure of the male genitalia is typical of Plagiognathus sensu stricto. Plagiognathus servadeii Wagner 1972. Wagner (1972) described this species from northern Italy and placed it in Plagiogna- thus (Poliopterus). Judging from the figure of the right paramere accompanying the original description, it does not belong to Europiella; the structure of the aedeagus will not clearly place the species in either Eu- ropiella or Plagiognathus, and we therefore provisionally leave the species in Plagiog- nathus sensu stricto. Plagiognathus pallescens Zheng and Li 1991. This species was described (Zheng and Li 1991) from China (Sichuan). The type series (of which we examined three specimens la- belled as paratypes), is mixed, as could be presumed from the great variability in the length, males 3.30—4.28, females 3.02-3.22. One male specimen labelled as a paratype from “Szechwan, 2000-2300 m, 10.IX.1963,” measures 4.5 mm in length and contradicts the original description regarding length and collecting data; it bears a handwritten para- type label where the other two specimens available for our examination have printed labels. The genitalia of this specimen are of the species figured with the original descrip- tion. A male from “‘Hsiao-chin” is 4.2 mm long, has genitalia very different in structure from the first discussed specimen, and is possibly not even congeneric with it. Nei- ther of these specimens belong to Europiella and could only be placed in Plagiognathus VOLUME 97, NUMBER 2 sensu stricto with difficulty. The third spec- imen is a female measuring 3.2 mm in length, from ‘‘Pao-shing, 950-1350 m, 1. VII.1963,” these data matching the orig- inal description except for the date; it pos- sibly belongs to the group of species closely related to alpina. Salicarus bimaculatus Zheng and Li 1991. This species was described from Sichuan, China. Judging from the size variability in- dicated in the original description (2.09-3.5 mm), it may be based on a mixed series. Clearly, this is not a species of Salicarus, but judging from the description closely re- sembles species closely related to albipen- nis, especially E. artemisiae. LITERATURE CITED Becker, A. 1864. Naturhistorische Mitteilungen. Bul- letin de la Societe Imperiale des Naturalistes de Moscou 37(2): 477-493. Bergroth, E. 1898. Ueber einige amerikanische Cap- siden. Wiener Entomologische Zeitung 17: 33-35. Carvalho, J.C. M. 1955. Keys to the genera of Mir- idae of the world. Boletim Museu Paraense Emilio Goeldi 11(2): 1-151. 1955. Analecta miridologica: Miscellaneous observations in some American museums and bibliography (Hemiptera). Revista Chilena de En- tomologia 4: 221-227. Dahlbom, A. G. 1851. Anteckningar 6fver Insekter, som blifvit observerade pa Gottland och i en del af Calmare Lan, under sommaren 1850. Kongliga Svenska Vetenskaps-Akademiens Handligar 1850: 155-229. Fallén, C. F. 1829. Hemiptera Sveciae. Sectio prior, Hemelytrata. Officina Berlingiana, Londini Goth- orum. 188 pp. Fieber, F. X. 1860-1861. Die europadischen Hemip- tera. Halbfliigler (Rhynchota Heteroptera). Carl Gerold’s Sohn, Wien. 444 pp. (pp. 1-112, 1860; pp. i-iv, 113-444, 1861). . 1864. Neuere Entdeckungen in europdischen Hemipteren. Wiener Entomologische Monat- schrift 8: 65-86, 205-234, 321-335. Gillette, G. P. and C. F. Baker. 1895. A preliminary list of the Hemiptera of Colorado. Colorado Ag- ricultural Experiment Station Bulletin 31 (Tech. Ser: 1) 137 pp: Henry, T. J. and A. G. Wheeler, Jr. 1988. Family Miridae Hahn, 1833, pp. 251-507. Jn Henry, T. J. and R. C. Froeschner, eds., Catalog of the Het- eroptera or True Bugs, of Canada and the Conti- nental United States. E. J. Brill, Leiden, New York. 393 Jakovlev, B. E. 1875. Bugs, Hemiptera Heteroptera, of the Astrakhan region. Bulletin de la Societe Im- periale des Naturalistes de Moscou 49(3): 145- 174. [In Russian and German. ] . 1877. New bugs, Hemiptera Heteroptera, of the Astrakhan fauna. Second supplement. Bulletin de la Societe Imperiale des Naturalistes de Moscou 52(2): 269-300. [In Russian and German. ] . 1880. Bugs (Hemiptera, Heteroptera) of Rus- sia and neighboring countries. I. Trudy Russ. En- tomol. Obshch. 11: 200-220. Josifov, M. 1974. Die Heteropteren der Bulgarischen Schwarzmeerkiiste. Izvestiya Instituta Zool. Muz., Sofia 39: 5-27. [In Bulgarian, German summary. ] Josifov, M. and I. M. Kerzhner. 1972. Heteroptera aus Korea. I. Teil (Ochteridae, Gerridae, Saldidae, Nabidae, Anthocoridae, Miridae, Tingidae und Reduviidae). Annales Zoologici (Warsaw) 29: 147- 180. Kerzhner, I. M. 1962. Materials on the taxonomy of capsid bugs (Hemiptera, Miridae) in the fauna of the USSR. Entomologicheskoye Obozreniye 41: 372-387. [In Russian.] . 1964. Fam. Miridae. /n Bei-Bienko, G. Ya., ed., Keys to the Insects of the European USSR. 1: 700-765. Moscow and Leningrad, Nauka. [In Russian.] [ English translation: 1967, Israel Pro- gram for Scientific Translation, Jerusalem, pp. 913- 1003.] 1978. Heteroptera of Saghalien and Kurile Islands. Trudy Biol.-Pochv. Inst. Dal’nevost. Nauch. Tsentra Akad. Nauk SSSR (n. s.) 50(153): 31-57. [In Russian.] . 1979. New Heteroptera from the Far East of the USSR. Trudy Zoologischeskogo Instituta Aka- demii Nauk SSSR 81: 14-65. [In Russian.] . 1988a. Fam. Miridae. Jn Ler, P. A., ed., Keys to the Insects of the USSR Far East. Vol. 2: 788- 857. Leningrad, Nauka. [In Russian.] [Issued March 1988]. 1988b. New and Little Known Heteroptera from the USSR Far East. Vladivostok, Akademia Nauk SSSR. 83 pp. (1987). [In Russian.] [Issued April 1988]. Kirkaldy, G. W. 1909. Hemiptera: Old and new. No. 2. Canadian Entomologist 41: 388-392. Knight, H. H. 1923. Family Miridae (Capsidae), pp. 422-658. Jn Britton, W. E., ed., The Hemiptera or Sucking Insects of Connecticut. Connecticut Geological and Natural History Survey Bulletin 34: 1-807. 1925. A list of Miridae and Anthocoridae from Alberta, Canada (Hemiptera). Canadian En- tomologist 57: 181-182. 1927. Notes on the distribution and host plants of some North American Miridae (Hemip- tera). Canadian Entomologist 59: 34-44. 1941. Plant bugs, or Miridae, of Illinois. Il- linois Natural History Survey Bulletin 22: 1-234. 394 1964. Phymatopsallus, new genus and new species of Phylinae from North America (Hemip- tera, Miridae). lowa State Journal of Science 39: 127-152. . 1968. Taxonomic review: Miridae of the Ne- vada Test Site and the western United States. Brig- ham Young University Science Bulletin 9(3): 1- 282. 1969. New species of Pronotocrepis Kngt., Europiella Reut., and Hesperocapsus Kngt., from western United States (Miridae, Hemiptera). Iowa State Journal of Science 44: 79-91. . 1970. Tannerocoris new genus, and new spe- cies of Miridae (Hemiptera) from the western United States. Great Basin Naturalist 30: 227- 238 Kulik, S. A. 1965. New species of capsid bugs (Het- eroptera, Miridae) from East Siberia and the Far East. Izv. Irkutsk. Sel’skokhoz. Inst. 25: 147-156. [In Russian.] 1975. A new species of the genus Plagiog- nathus (Heteroptera, Miridae) from the vicinity of Ussuriysk. Entomologischeskoye Obozreniye 54: 587-588. [In Russian. ] Li, H.-y.and L.-y. Zheng. 1991. Genus Plagiognathus Fieber (Hemiptera: Miridae) from China. Acta Scientiarum Naturalium Univsitatis Nan Kaiensis 3: 88-97. [In Chinese with English summary. ] Lindberg, H. 1934. Inveta entomologica itineris His- panici et Maroccani, quod a. 1926 fecerunt Harald et Hakan Lindberg. Societas Scientiarum Fennica, Commentationes Biologicae 4(12): 23 pp. Matsumura, S. 1917. Oyo Konchu-gaku | [Applied Entomology 1]. Cited from Second Rev. Edition, 1920, 731 + 12 pp., 50 pls. Miyamoto, S. 1969. Notes on the species of the genus Plagiognathus Fieber in Japan and Saghaline. Sie- boldia 4: 85-94. 1977. On the scientific names concerning Japanese Heteroptera (3). Rostria 28: 231-234. [In Japanese. ] Puton, A. 1873. Notes pour servir a l’etude des Hém- iptéres. Annals de la Societe Entomologique de France (5)3: 11-26. Putshkov, V.G. 1971. On the ecology of some little- known species of Heteroptera of the European USSR. Communication IV. Miridae. Vestnik Zoologii 1971(5): 30-35. [In Russian. ] . 1975. New species of Miridae from the Cau- casus (Hepteroptera). Doklady Akademii Nauk Ukrainskoi SSR (B) 1975(3): 281-284. [In Rus- sian.] Qi, B.and Nonnaizab. 1993. New species and a newly recorded species of Plagiognathus Fieber of China (Hemiptera: Miridae). Entomotaxonomia 15: 29- 34. [In Chinese with English summary.] Reuter, O. M. 1875. Genera Cimicidarum Europae. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bihang Kongliga Vetenskaps-Akademien Han- dlingar 3(1): 1-66. 1878. Hemiptera Gymnocerata Europae. Hémiptéres Gymnocérates d’Europe, du bassin de la Méditerranée et de l’Asie russe, 1. Acta Socie- tatis Scientiarum Fennicae 13: 1-188, 8 pls. 1880. Diagnoses Hemipterorum novorum. II. Ofversigt Finska Vetenskaps-Societatens Fér- handlingar 22: 9-24. 1888. Descriptio speciei nova sicilianae ge- neris Plagiognathus (Hemiptera, Capsidae). Na- turalista Siciliana 7: 236. . 1901. Capsidae rossicae descriptae. Ofversigt Finska Vetenskaps-Societatens Férhandlingar B 43: 161-194. 1906. Capsidae in prov. Sz’tschwan Chinae a DD. G. Potanin et M. Beresowski collectae. An- nuaire Musée Zoologique Académie Impériale Sciences St. Petersburg 10: 1-81. 1909. Bemerkungen iiber nearktischen Cap- siden nebst Beschreibung neuer Arten. Acta So- cietatis Scientiarum Fennicae 36(2): 1-86. Sahlberg, J. 1920. Enumeratio Hemipterorum Het- eropterorum faunae Fennicae. Editio secunda auc- ta et emendata. Bidrag Kannedom Finlands Natur Folk 79(2): 1-227. Sienkiewicz, I. 1986. Description de Plagiognathus (Poliopterus) strawinskii n. sp. (Heteroptera, Mir- idae). Nouvelle Revue Entomologique (n.s.) 3: 373- 378. Stichel, W. 1934. Illustrierte Bestimmungstabellen der Deutschen Wanzen, Heft 10: 275-306. W. Sti- chel, Berlin. 1956. Illustrierte Besstmmungstabellen der Wanzen. II. Europa. Vol. 2, Heft 11, pp. 321-352. Privately printed, Berlin-Hermsdorf. Stonedahl, G. M. 1990. Revision and cladistic anal- ysis of the Holarctic genus Atractotomus Fieber (Heteroptera: Miridae: Phylinae). Bulletin of the American Museum of Natural History 198: 1-88. Uhler, P.R. 1893. Summary of the collection of He- miptera secured by Mr. E. A. Schwarz in Utah. Proceeding of the Entomological Society of Wash- ington 2: 366-385. 1894. Observations upon the heteropterous Hemiptera of Lower California, with descriptions of new species. Procceding of the California Acad- emy of Sciences, ser. 2, 4: 223-295. . 1895. Descriptions of new genera and species of Hemiptera, /n Gillette, G. P. and C. F. Baker, A Preliminary List of the Hemiptera of Colorado. Colorado Agricultural Experiment Station Bulle- tin 3)l (lech.“Ser; 1) s13:75pp: Van Duzee, E. P. 1915. New genera and species of North American Hemiptera. Pomona Journal of Entomology and Zoology 7: 109-121. 1916. Checklist of the Hemiptera (excepting VOLUME 97, NUMBER 2 the Aphididae, Aleurodidae and Coccidae) of America, North of Mexico. New York Entomol- ogy Society, New York. xi + 111 pp. . 1917. Catalogue of the Hemiptera of America North of Mexico Excepting the Aphididae, Coc- cidae and Aleurodidae. University of California Publications, Technical Bulletin, Entomology 2: i-xvi + 1-902. . 1923. Expedition of the California Academy of Sciences to the Gulf of California in 1921. The Hemiptera (true bugs, etc.). Proceedings of the California Academy of Sciences, fourth ser. 12: 123-200. Wagner, E. 1941. Zweineue deutsche Plagiognathus- Arten (Hem. Miridae). Stettiner Entomologische Zeitung 102: 248-257. . 1949. Plagiognathus (Poliopterus) litoralis n. sp. (Hem. Heterop. Miridae). Verhandlungen Ver- eins Naturwissenschaftliche Heimatforschung Hamburg 30: 53-58. 1952a. Blindwanzen oder Mindae. Jn Die Tierwelt Deutschlands, 41. 218 pp. . 1952b. Zwei neue Miridenarten aus dem Mit- telmeergebiet (Hem. Heteropt.). Entomologische Berichten 14: 41-44. . 1954. Systematische Ergebnisse einer Durch- sicht der Sammlung O. M. Reuter, I. Notulae En- tomologicae 34: 75-81. 1956a. Die Plagiognathus-Gruppe (Hem. Heteropt. Miridae). Acta Entomologicae Musei Nationalis Pragae 30(1955): 291-304. 1956b. Plagiognathus (Poliopterus) gracilis nov. spec. (Heteropt. Miridae). Deutsche Ento- mologische Zeitschrift (N.F.) 3: 74-75. . 1958. Sur quelques espéces francaises du gen- 395 re Psallus Fieb. (Hem. Het. Miridae). Vie et Milieu 8: 322-325. 1963a. Was ist Psallus argyrotrichus Fieber 1861 (Hem. Het. Miridae)? Miscellanea Zoologica 1(5): 1-4. 1963b. Zur Systematik des Psallus-Kom- plexes (Hem. Het. Miridae). Mitteilungen Miiche- ner Entomologischen Gesellschaft 53: 150-163. 1964. Drei neue Arten aus der Unterfamilie Phylinae Dgl. et Sc. (Hem. Het. Miridae). Bollet- tino Societa Entomologica Italiana 94: 70-75. . 1969a. Einige neue Miriden aus Libyen (Het- eropera). Notulae Entomologicae 49: 11-24. 1969b. Einige Miridae aus Nord- und Vor- derasien (Heteroptera). Notulae Entomologicae 49: 31-36. 1972. Zwei neue Phylinae aus Italien (He- miptera Heteroptera Miridae). Bollettino Societa Entomologica Italiana 104: 111-113. 1975. Die Miridae Hahn, 1831, des Mittel- meerraumes und der Makaronesischen Inseln (He- miptera, Heteroptera). Part 3. Entomologische Abhandlungen 40(suppl.): 1483. Wagner, E. and H. H. Weber. 1964. Hétéroptéres Miridae. Faune de France, 67. 592 pp. Wheeler, A. G., Jr. and E. R. Hoebeke. 1982. Psallus variabilis (Fallén) and P. albipennis (Fallén), two European plant bugs established in North Amer- ica, with notes on taxonomic changes (Hemiptera: Heteroptera: Miridae). Proceedings of the Ento- mological Society of Washington 84: 690-703. Zheng, L.-y. and H.-y. Li. 1991. Two new species of Phylinae from China (Insecta, Heteroptera: Mir- idae). Reichenbachia 28: 113-115. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 396-400 THE FIRST RECORD FOR BOTH YNOTUS PILOSUS (BOHEMAN) (HEMIPTERA; MIRIDAE) IN THE NEARCTIC REGION G. G. E. SCUDDER Department of Zoology, University of British Columbia, Vancouver, British Columbia, Canada, V6T 1Z4. Abstract.—The mirid species Bothynotus pilosus (Boheman) is reported from the Ne- arctic Region for the first time. This species, formerly known only from the Palearctic Region is recorded from Yukon and British Columbia. It is considered a natural Beringian element. Key Words: During a study of the Heteroptera of the Yukon, the deraeocorine mirid Bothynotus pilosus (Boheman) was collected in three lo- calities. It has also been found in one lo- cality in northern British Columbia. The distribution suggests that this species is a natural Beringian element in the Nearctic fauna, and not an introduced species. Male and female of the species are rede- scribed and illustrated, and notes given to distinguish it from other members of the genus in the New World. Bothynotus pilosus (Boheman) Phytocoris pilosus Boheman 1852: 68. Capsus fairmairii Signoret 1852: 542. Syn. by Reuter 1873: 8. Capsus horridus Mulsant and Rey 1852: 132. Syn. by Reuter 1875: 91. Bothynotus minki Fieber 1864: 77. Syn. by Puton 1873: 24. Bothynotus kiritschenkoi Lindberg 1934: 20. Syn. by Josifov and Kerzhner 1972: 152. Redescription (measurements for mean and range in mm; range in parentheses). — Macropterous male (Fig. 1): Oblong, thick- ly clothed with more or less erect, long, brown setae. Head, pronotum (including Heteroptera, Miridae, Bothynotus pilosus, Nearctic Region collar and calli), scutellum and thoracic venter black and shiny; head posteriorly and ventrally sometimes reddish. Antennae brown; rostrum pale brown with tip almost black. Hemelytra with corium uniform brown; cuneus dark brown; membrane dusky brown. Legs pale brown, with femora often yellowish; apex of tibiae and whole of tarsi dark brown. Abdominal venter pale to dark brown. Head smooth and shiny; head width 0.89 (0.83-0.95), vertex width 0.53 (0.50-0.55); first antennal segment about *4 width of ver- tex; second antennal segment cylindrical and not incrassate, distinctly longer than head width; third and fourth antennal segments much thinner than second, combined length of third and fourth segments subequal to or slightly longer than second; all antennal seg- ments with both short and long setae, short- er setae dense, oblique and about as long as width of second antennal segment, longer setae more scattered, more erect and over twice as long as width of second antennal segment; antennal measurements I 0.43 (0.40-0.45): II 1.17 (1.00-1.27): III 0.63 (0.53-0.70): IV 0.48 (0.43-0.57); rostrum reaching to middle coxae, length 1.16 (1.13- 1:20). VOLUME 97, NUMBER 2 397 Fig. 1. Bothynotus pilosus, male. 398 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Bothynotus pilosus, female. Pronotum distinctly punctate; calli smooth and sunken; pronotal length 0.98 (0.87-1.03), pronotal width 1.61 (1.37- 1.77), 1.2-1.6x greater than length of sec- ond antennal segment. Scutellum shiny, transversely wrinkled and appearing some- what swollen. Meso- and metapleura smooth and shiny. Corium and clavus shiny and appearing roughened, but not distinctly punctate; membrane shiny and reticulately wrinkled; hemelytra extending well beyond end of body, apex of cuneus reaching be- VOLUME 97, NUMBER 2 yond end of abdomen. Tibiae with shorter oblique setae equal to width of tibia, and longer, more erect setae much longer than width of tibia. Total length 5.70 (5.30-6.10). Brachypterous female (Fig. 2): Oval, thickly clothed with long, erect, brown se- tae. Body both above and below, including hemelytra, collar and calli, shiny black; head posteriorly and ventrally distinctly red; an- tennae brown, with first segment except ex- treme base and apex pale yellow. Legs brown with femora and tibiae pale yellow. Structured as for male, except as follows: head width 0.99 (0.97-1.00), vertex width 0.59 (0.57-0.63); antennal measurements I 0.44 (0.43-0.45): II 1.05 (1.00—1.13): III 0.67 (0.63-0.70): IV 0.51 (0.50-0.53); rostral length 1.32 (1.30-1.33). Pronotum closely and coarsely punctate; pronotal length 0.86 (0.77-0.92), pronotal width 1.57 (1.50-1.67). Scutellum trans- versely wrinkled, with raised central lon- gitudinal line; meso- and metapleura trans- versely wrinkled. Hemelytra lacking mem- brane, and attaining tergum IV; costal mar- gin of corium strongly convex throughout; surface of corium, clavus and cuneus coarsely roughened, but not distinctly punc- tate. Total length 4.40 (3.80-4.80). North American material examined: | 6, BRITISH COLUMBIA, Summit Lk., 16.vii1.1982 QL. A. Kelton); 1 ¢, YUKON, mi 51, Dempster Hwy., 18-27.vii.1973 (G. and D. M. Wood); | 4, YUKON, Klondike R., Dempster Corner, 1 km W, 63°59'N 138°48'’W, 20.vii.1982 (G. G. E. Scudder); 7652, YUKON, Old Crow, in root mats of Polemonium pulcherrimum Hook., 14.v11.1983 (S. G. Cannings); 1 2, YUKON, Old Crow, fall trap, S-facing Artemisia slope, 4-19.vu1.1983 (S. G. Cannings); 1 4, YU- KON, Old Crow, pitfall trap, boreal forest clearing, 21-26.vii.1984 (S. G. Cannings). In the Canadian National Collection (CNC), Ottawa, ON; Spencer Entomological Mu- seum, University of British Columbia, Van- couver, BC; and Department of Entomol- ogy, Oregon State University, Corvallis, OR. 399 Discussion: The above specimens were compared with material determined as this species by H. Lindberg in 1958 with data: AL. Eckero Torp, 14.v1i.1943 (H. Lindberg) [CNC]. Dissection of the male genitalia by M. D. Schwarz showed no noticeable dif- ferences. Bothynotus pilosus will key to complete 9 in the key to the New World species of Bothynotus Fieber by Henry (1979). Like B. barberi Knight, the calli in B. pilosus appear sunken into the pronotum. However, the pronotal collar and calli are fuscous in B. pilosus and not pale orange as in B. barberi. The male in B. pilosus with an average total length of 5.70 mm 1s also much larger than B. barberi with length 3.44 mm. Bothynotus pilosus with black collar and calli, with black procoxal cleft, and head anteriorly black, is clearly distinct from both B. floridanus Henry and B. mexicanus Henry. Bothynotus pilosus was reported by Car- valho (1957) from Austria, Belgium, Corfu, Finland, France, Great Britain, Germany, Greece, Hungary, Italy, Netherlands, Rus- sia, Scandinavia, and Switzerland. Stichel (1956) also listed Czechoslovakia and Ju- goslavia, and Josifov and Kerzhner (1972) added Korea. Stichel (1956) reported the species from Pinus sylvestris L. and Picea excelsior Lk., but Woodroffe (1970) considered that moss is the primary habitat of this species, at least in Scotland, where he collected both adults and larvae in the moss Hypnum cupressi- forme Hedw. The fact that B. pilosus in the Yukon was collected in root mats of Pole- monium pulcherimum (Polemoniaceae), and also in pitfall traps, confirms that the mem- bers of the genus Bothynotus are largely ground-dwelling as suggested by Henry (1979). However, B. pilosus was also taken by sweeping in the Yukon. Southwood and Leston (1959) also report that this species can often be collected by sweeping in the evenings in the British Isles. Southwood and Leston (1959) observed that the species overwinters in the egg stage, 400 and that adults occur from the last weeks in June until early October in Britian. All Nearctic specimens were collected in July and August, so the life cycle may be the same in the New World. ACKNOWLEDGMENTS Research was supported by grants from the Natural Sciences and Engineering Re- search Council of Canada. Studies in the northern Yukon were possible through lo- gistic support provided through the Polar Continental Shelf program of the Govern- ment of Canada, Department of Indian and Northern Affairs. I am indebted to Drs. A. Asquith (U.S. Fish and Wildlife Service, Honolulu), J. D. Lattin (Oregon State Uni- versity) and M. D. Schwartz (Agriculture Canada, Ottawa) for help with confirmation of the identification of Bothynotus pilosus. Dr. R. Foottit (Agriculture Canada, Ottawa) kindly provided copies of some relevant ref- erences. The drawings are by Launi Lucas (University of British Columbia). LITERATURE CITED Boheman, C. H. 1852. Nya svenska Hemiptera. Ofversigt af Kongliga Svenska Vetenskaps-Aka- demiens Forhandlingar 9(4): 49-60, 65-80. Carvalho, J.C. M. 1957. Catalogue of the Miridae of the World. Part I. Cylapinae, Deraeocorinae, Bryocorinae. Arquivos do Museu Nacional, Rio de Janeiro 44(1): 1-158. Fieber, F. X. 1864. Neuere Entdeckungen in euro- pdischen Hemipteren. Wiener Entomologische Monatschrift 8(3): 65-86. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Henry, T. J. 1979. Review of the New World species of Bothynotus Fieber (Hemiptera: Miridae). Flor- ida Entomologist 62: 232-244. Josifov, M. and I. M. Kerzhner. 1972. Heteroptera aus Korea. I. Teil (Ochteridae, Gerridae, Saldidae, Nabidae, Anthocoridae, Miridae, Tingidae and Reduviidae). Annales Zoologici (Warszawa) 29(6): 147-180. Lindberg, H. 1934. Verzeichnis der von R. Malaise in Jahre 1930 bei Vladivostok gesammelten Het- eropteren. Notulae Entomologicae, Helsingfors 14: 1-23. Mulsant, E. and C. Rey. 1852. Description de quel- ques Hémiptéres Hétéroptroptéres nouveaux ou peu connus. Annales de la Societé Linnéenne de Lyon: 76-141. Puton, A. 1873. Notes pour servir a l’étude des Hé- miptéres. Annales de la Societé Entomologique de France 5(3): 11-26. Reuter, O. M. 1873. Bidrag till nordiska Capsiders synonymi. Notiser ur Férhandlingar Sallskapet pro Fauna et Flora Fennica 14: 25 pp. . 1875. Revisio critica Capsinarum, praecipue Scandinaviae et Fenniae. Akademisk Afhandling, Helsingfors 1(1): 91. Signoret, V.A. 1852. Notice sur quelques Hémiptéres nouveaux ou peu connus. Annales de la Societé Entomologique de France 2(10): 539-544. Southwood, T. R. E. and D. Leston. 1959. Land and Water Bugs of the British Isles. Frederick Warne and Co. Ltd., London and New York. Stichel, W. 1956. Illustrierte Bestimmungstabellen der Wanzen. II. Europa (Hemiptera-Heteroptera Europae). 6. Heft. Cimicomorpha Miridae Cylap1- nae-Deraeocorinae, pp. 170-192. W. Stichel, Ber- lin-Hermsdorf. Woodroffe, G. E. 1970. Notes on some Hemiptera- Heteroptera from Aviemore, Inverness-shire. En- tomologist’s Monthly Magazine 105: 165-166. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 401-408 METASEQUOIAMIRIS CARVALHOI, A NEW GENUS AND SPECIES OF CONIFER-INHABITING MIRINI FROM CHINA (HETEROPTERA: MIRIDAE: MIRINAE) MICHAEL D. SCHWARTZ Biological Resources Division (BRD), Centre for Land and Biological Resources Re- search, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada K1A 0C6. Abstract. — Metasequoiamiris carvalhoi, a new genus and species of mirine Miridae from West Hubei Province, China, is described and illustrated. This bug apparently inhabits the conifers Cephalotaxus fortunei Hook., Metasequoia glyptostroboides Hu and Cheng, and TJorreya sp. The relationships of the new genus to other Mirini is discussed. Key Words: While sorting unidentified Miridae from the California Academy of Sciences, San Francisco (CAS), I uncovered the first plant bug species known to inhabit Metasequoia. This new species of Mirini is reminiscent of Orthops Fieber and Pinalitus Kelton but has several features that require its place- ment in a new genus. Metasequoiamiris car- valhoi, new genus, new species, is described and documented, and its affinities to related Mirini genera are discussed. It is appropriate that the recognition of a new genus, apparently the associate of an ancient and rare host plant, be included in this issue honoring our venerable and pro- ductive colleague, José C. M. Carvalho. Metasequoiamiris Schwartz, NEw GENUS (Figs. 1-19) Type species. —Metasequoiamiris carval- hoi, new species. Diagnosis. — Distinguished from other Mirini by the following combination of at- tributes: wide, pale collar, contrasting in color with dark calli (Fig. 1); campanulate pronotum (Figs. |, 3); rounded, swollen frons and adjoining jugum, imperceptible dorsal Insecta, Heteroptera, Miridae, Mirini, Metasequoiamiris, new genus margin of tylus (Fig. 2); strongly punctate dorsum; nondeflected cuneus; long, sub- erect setae; convergent parempodial apices (Fig. 7); and long, trough-shaped sclerite of vesica (Fig. 14). Description.— Male. Macropterous; gen- eral coloration castanaceous, with pale col- lar and abdominal sternites; dorsal surface strongly punctate, shining; head and scu- tellum smooth; dorsal vestiture consisting of uniformly distributed, long, fine, sub- erect, yellow setae; calli without setae, frons with sparse setae; pronotal and hemelytral setae set in punctures (Fig. 3). Head: tri- angular with tylus obscured by frons and posterior margin straight in dorsal view; frons and temporal area rounded, swollen anterior and dorsal to eyes; dorsal carina obsolete; tylus short with ventral margin not projecting anterior to dorsal junction in lat- eral view; anteocular portion of head short- er than eye height; eye small, gena subequal to half eye height; labium reaching to apex of mesocoxa. Antenna: cylindrical, seg- ments I and II of similar diameter, III and IV slightly narrower; inserted slightly ven- tral to middle of eye, fossa adjacent to an- terior margin of eye; segment I longer than 402 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Metasequoiamiris carvalhoi, n. sp., dorsal habitus of male. VOLUME 97, NUMBER 2 403 17 " ~~, > ceil al ks ae TAS A Figs. 2-7. Scanning electron micrographs of Metasequoiamiris carvalhoi. 2, head, lateral view: 3, pronotum and head, dorsal view; 4, metathoracic scent efferent system, lateral view; 5, apex of protibia, ventral view; 6, mesofemoral trichobothrium; 7, pretarsus, posterior view (pulvillus, arrow). 404 © PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON i SS © 8 cd a 9 ee a AE ra © ©) @ S nN —— ~ os x Figs. 8,9. Metafemoral trichobothria of Metasequoiamiris carvalhoi, ventral view. 8, male; 9, female. width of vertex, segment II just longer than posterior width of pronotum, segment III longer than IV; all segments with reclining dark setae, segments III and IV also with a few longer, suberect setae. Pronotum: cam- panulate; collar wider than width of anten- nal segment I, with dense trichomae (Fig. 2), anterior margin slightly concave medi- ally; calli flat, separated medially, reaching lateral margins of pronotum. Scutellum: weakly elevated. Metathoracic scent effer- ent system: triangular evaporative area; per- itremal disc small, slightly raised; disc and adjacent metapleuron densely pubescent (Fig. 4). Hemelytra: elongate, parallel-sided, surpassing genital segment in lateral view, declivous at cubitus; embolium obsolete; cuneus about two times as long as broad, not bent ventral to corium; cuneal fracture small; primary cell much longer than broad, secondary cell narrow. Legs: with moder- ately distributed, suberect, shining, pale se- tae; femora slightly fusiform with oval cross- section; metafemora with 6 trichobothria (Fig. 8), trichomae strongly developed (Fig. 6); tibiae cylindrical, protibia with concave apex and comb (Fig. 5); tarsi three-seg- mented, segment 3 subequal to combined length of 1 and 2; pretarsus with small, curved claws; lamellate parempodia with convergent apices; pulvilli apparently ab- sent (Fig. 7 cf. arrow). Genitalia: Genital capsule: without tubercles dorsal to para- mere insertions; aperture oval. Left para- mere: C-shaped (Fig. 10). Right paramere: narrow distally (Fig. 12). Phallotheca: horn- shaped, without apical spines or tubercles, aperture almost reaching base (Fig. 13). Ve- VOLUME 97, NUMBER 2 405 13 Figs. 10-15. Male genitalia of Metasequoiamiris carvalhoi. 10, 11, left paramere. 10, lateral view; 11, distal view; 12, right paramere, lateral view; 13, phallotheca, lateral view; 14, 15, vesica. 14, posterior view (BS, basal sclerite; LS, lobal sclerite; TS, trough-shaped distal portion of basal sclerite); 15, anterior view. sica: ductus seminis slightly thickened me- dially; aperture of secondary gonopore oval and complete; basal sclerite trough-shaped and lobal sclerite narrow (Fig. 14). Female.— Macropterous; similar to male in color and structure, except apex of pro- tibia with deeper concavity. Legs: metafe- mora with 6 trichobothria (Fig. 9). Geni- talia: Posterior wall: narrow, with lateral apices strongly recurved (Fig. 16, IS). Lat- eral lobe: absent. Median process: present (Fig. 16, MP). Dorsal structure: oval (Fig. 16). Sclerotized rings: small, flattened oval (Fig. 17, SR). Dorsal labiate plate: contin- uous, narrow and indistinct medially (Fig. 17, DLP). Ventral labiate plate: discontin- uous (Fig. 18, VLP). First valvulae: distal serrations coarse (Fig. 19). Distribution. — All known specimens are from Lichuan district, West Hubei prov- ince, China. Etymology.— Named for one of the host plants, Metasequoia glyptostroboides. Discussion.—The campanulate prono- tum, parallel-sided, elongate hemelytra, nondeflected cuneus, and convergent par- empodia of Metasequoiamiris are similar in appearance to the Neotropical orthotyline genus Falconia Distant. However the new genus is clearly placed in the Mirini and possesses characters which show a relation- ship to some of the genera examined by Schwartz (1994). The addition of the new genus to the data set and analysis in Schwartz (1994, Table 1; coded for characters 0 to 36 as follows: 00001 20000 00010 10033 13301 406 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 16-19. Female genitalia of Metasequoiamiris carvalhoi. 16, posterior wall, anterior view (DS, dorsal structure; IL, inter-ramal lobe; IS, inter-ramal sclerite; MP, median process; SP, shovel-shaped anterior surface of median process); 17, left sclerotized ring, dorsoposterior view (DLP, dorsal labiate plate; SR, sclerotized ring); 18, ventral labiate plate, anterior view (VF, first valvifer; VL, first valvulae; VLP, ventral labiate plate); 19, distal end of first valvulae, lateral view. 35141 21401 00) produces one most par- simonious cladogram (length = 114, ci = 72, ri = 72, after removal of uninformative characters and successive approximations weighting of two resultant equally parsi- monious cladograms) with a structure which places Metasequoiamiris as the sister genus to the monophyletic taxa forming node 5 (Schwartz 1994, Fig. 29). Metasequoiamir- is, Orthops, Pinalitus, Salignus Kelton, and Oreolygus Linnavuori form a group based on the possession of the following charac- ters: white pronotal collar (Fig. 1); left par- amere with tapered shaft (Fig. 11); narrow inter-ramal sclerite with recurved lateral apices (Fig. 16); and small, oval sclerotized rings (Fig. 17) (the left paramere and scler- otized rings are homoplasious in Orthops and the collar and inter-ramal sclerite are homoplasious in Salignus). The vesica of the new genus is most similar to Orthops in the structure of the posterior lobe of the basal sclerite. The strongly developed me- tafemoral trichomae and apparent absence of pretarsal pulvilli, in addition to the suite of characters listed in the Diagnosis of Metasequoiamiris, are unusual for the Mir- ni. The utility of the hypothesized rela- tionships of the ““Lygus complex” genera presented here and in Schwartz (1994) can be properly addressed only with a compre- hensive analysis of the tribe. The presence of lamelliform parempodia with either divergent or convergent apices has been used as a diagnostic feature of the Mirinae and Orthotylinae respectively (Knight 1968). That there is variation in the two subfamilial forms of lamelliform par- empodia is well known (see Figs. 54-67 in Knight 1968). Closterocoris Uhler and Cy- phopelta Van Duzee in the Herdoniini have narrow, straplike lamelliform parempodia which resemble the setiform parempodia of Phylini. Mirini with convergent parempo- VOLUME 97, NUMBER 2 dia, like those in Metasequoiamiris, have not been documented previously. Metasequoiamiris carvalhoi, Schwartz, NEw SPECIES (Figs. 1-19) Description. —Castanaceous with vari- able brown to black areas on head, antennal segments II-IV (except base of II), anterior portion of pronotum (calli always) including disk laterally, proepisternum, propleuron, mesoscutellum, scutellum, near claval com- missure, distal one-third of cubitus, meso and metasternite, anterior portion of evap- orative area of metathoracic scent efferent system, abdominal segment 2, rest of ab- dominal sternite laterally, and genital seg- ment; pale yellow on legs, labium, posterior portion of evaporative area of metathoracic scent efferent system, and abdominal ster- nites 3-8 medially; orange to red on interior and distal lateral margins of cuneus, distal half of metatibia, and most of antennal seg- ment I; hemelytral membrane dusky brown. Genitalia: Left paramere: sensory lobe prominent, without apical spines, with short setae (Fig. 10); arm and angle C-shaped, with short to moderate length setae; shaft short and tapered; apex with rounded dorsal point and sharp ventral point (Fig. 11). Right par- amere: with broadly rounded sensory area, with moderate length setae; distal one-third a narrow shaft; apex with rounded dorsal point and sharp ventral point (Fig. 12). Ve- sica: Basal sclerite: narrowly enveloping base of vesica (Fig. 14, BS); anterior lobe con- tinuing posterior to ductus seminis and sec- ondary gonopore as a narrow, trough-shaped sclerite terminating at edge of membrane (Fig. 14, TS); posterior lobe forming a strongly spinulose patch posterior to sec- ondary gonophore (Fig. 15). Membrane: re- duced dorsal to secondary gonopore, asso- ciated with basal portion of the lobal scler- ite. Lobal sclerite: long, curved, attached at base of vesica projecting dorsally beyond membrane and apex of basal sclerite, with- out spines (Fig. 14, LS). 407 Measurements (in mm; n = 20).—Length from apex of tylus to apices of membrane 3.49-4.10; length from apex of tylus to cu- neal fracture 2.66-2.91:; width across he- melytra at apex of clavus 1.09-1.26; pos- terior width of pronotum 1.13-1.23; width of head across eyes 0.73-0.76; width of ver- tex 0.34-0.38; length of antennal segment I 0.41-0.46; II 1.18-1.28; III 0.59-0.73; IV 0.53-0.58; length of labium 1.08-1.16. Female.—Color similar to male. Geni- talia: Posterior wall: Inter-ramal sclerite: narrow, ventral margin broadly rounded, sclerite deeply incised on both sides of me- dian process (but with membrane); lateral apices strongly recurved (Fig. 16, IS). Inter- ramal lobe: broadly contiguous medially, broadly rounded ventrally (Fig. 16, IL). Me- dian process: well-sclerotized, with broad shovel-shaped anterior surface (Fig. 16, MP, SP). Dorsal structure: edge not extending beyond width of shovel-like median process (Fig. 16, DS). Dorsal labiate plate: width equal to width of rings (Fig. 17, DLP). Measurements (n = 20).—Length from apex of tylus to apices of membrane 4.10- 4.43; length from tylus to cuneal fracture 2.85-3.10; width across hemelytra at apex of clavus 1.28-1.40; posterior width of pro- notum |.18-1.30; width of head across eyes 0.73-0.76; width of vertex 0.38-0.41; length of antennal segment I 0.38-0.43; II 0.95- 1.07; III 0.60-0.66; IV 0.54—0.60; length of labium 1.28-1.40. Distribution.— All known specimens are from Lichuan District, West Hubei Prov- ince, China, a region with a distinctive floral association (Chu and Cooper 1950, Wang 1961). Etymology.—Named to honor the late José C. M. Carvalho in appreciation of his long and productive career devoted to the systematics of the Miridae. Holotype male.—China. West Hupeh: [West Hubei, Lichuan District:] Suisapa [Shui-hua-pa], [1000 m], 25 July 1948, J. L. Gressitt Collector, ex. Torreya sp. (de- posited in the CAS). 408 Paratypes. — All with the same label data as the holotype except as follows—24 July: 1 6, 2 2 [1 2 dissected]; ex. Castanea sp., | 6 [metafemora dissected, Fig. 8]; ex. Ce- phalotaxus fortunei (possibly a Torreya— sterile), 2 3, 4 2 [é used in Fig. 1]. 25 July: 2 9: ex. Metasequoia glyptostroboides Hu and Cheng, 3 6, 1 2; ex. Torreya sp., 1 4, 3 9; ex. C. fortunei (possibly a Torreya—sterile), 14 6, 7 & (2 6 used inthis st] 226 4uly, less 1 Aug., #2401, ex. M. glyptostroboides, Bish- op Museum, | 4, 7 2, 30 Aug.: ex. C. fortunei (possibly a Torreya—sterile), 1 4, 29 2 [6 dissected, Figs. 10-15; 1 2 SEM #189a, Fig. 7]; ex. M. glyptostroboides, 3 2 [metafemora dissected, Fig. 9]. 12 Sept.: ex. C. fortunei (possibly a Torreya—sterile), 20 2; ex. Liq- uidambar formosana Hance, | 2. 17 Sept., 1 2 [dissected, Figs. 16-19]. China. West Hubei: Lichuan District: Leong-Ho-Kow, 9 Sept. 1948, ex. C. fortunei (possibly a Tor- reya—sterile), | 2. Voucher specimens are deposited in the American Museum of Nat- ural History, New York, NY; Bishop Mu- seum, Honolulu, HI; CAS; Canadian Na- tional Collection, Ottawa (CNC); Nankai University, Tianjin; Natural History Mu- seum, London; and U.S. National Museum, Washington, D.C. Other specimens.— All with the same la- bel data as the Suisapa paratypes from the following dates. The specimens are not in- cluded in the type series because they are either damaged or teneral—24 July 1 9; 25 July 3 6 [1 ¢ SEM #288b; Fig. 6], 3 2; 26 July 5 6, 1 9; 1 Aug. 3 2; 30 Aug. 3 6 [1 4 SEM #288a, Figs. 2-5], 4 9; 12 Sept. 15 9 (CASNENG): PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ACKNOWLEDGMENTS I am appreciative of P. H. Arnaud, Jr. and K. Ribardo (Department of Entomol- ogy, California Academy of Sciences, San Francisco, CA) for lending all of the speci- mens of M. carvalhoi: G. M. Stonedahl (CAB International Institute of Entomology) for sorting additional specimens of M. carval- hoi from the CAS and commenting on the manuscript; D. M. Moorehouse (BRD) for the dorsal habitus illustration; K. Bolte (BRD) for the scanning electron micro- graphs; T. J. Henry (Systematic Entomology Laboratory, ARS, USDA, Washington, D:C:); J. E.-O'Hara*(BRD)) Jn] Huber (BRD), and L.-Y. Zheng (Department of Bi- ology, Nankai University, Tianjin, China) for helpful comments on an earlier version of the manuscript. LITERATURE CITED Chu, K. L. and W. S. Cooper. 1950. An ecological reconnaissance in the native home of Metasequoia glyptostroboides. Ecology 31: 260-278. Knight, H. H. 1968. Taxonomic review: Miridae of the Nevada Test Site and western United States. Brigham Young University Science Bulletin 9: 1- 282. Schwartz, M.D. 1994. Review of the genus Salignus Kelton and a character discussion of related genera (Heteroptera: Miridae: Mirinae). Canadian Ento- mologist 126: 971-993. Wang, C. W. 1961. The forests of China with a Survey of Grassland and Desert Vegetation. Maria Moors Cabot Foundation Publication No. 5. Harvard University, Cambridge. 313 pp. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 409-426 NEW GENERA AND SPECIES OF RHYPAROCHROMINAE FROM WEST AFRICA (HEMIPTERA; LYGAEIDAE) DESCRIBED IN HONOR OF J.C. M. CARVALHO JAMES A. SLATER Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06269. Abstract.— Carvalhodrymus n.g.; Carvalhodrymus elegans, Mizaldus lestoni, M. lin- nayuorii, M. carvalhoi, M. tenuis, Fontathanus rostratus, F. ghanaensis, Paromius carvalhoi and Esinerus humidus n. spp. are described as new from West Africa. A key is given for the African species of Mizaldus. Modifications of a key to African Drymini genera and a discussion of zoogeographic relationships of West African Lygaeidae is included. Key Words: It is a special pleasure and honor to ded- icate this paper to Dr. José C. M. Carvalho, my good friend and colleague for half a cen- tury, in recognition of his many contribu- tions to our knowledge of the world fauna of Heteroptera and especially that of the tropics. The present paper has been made possible by the field work and careful study of several of Dr. Carvalho’s friends and colleagues. Without the material that they have brought together, the great West African fauna would still be very poorly understood. In the pres- ent paper I have acknowledged my debt to some of these people by the establishment of patronymics for them. The West African rhyparochromine fau- na is an especially important one for several reasons. Not only is it very rich in species (a great many still undescribed), but it shows a number of close relationships to the Ori- ental fauna, particularly of the forest-living forms. This strongly suggests (as it does in other animal groups) a former closer con- nection faunistically and floristically be- tween West Africa and East Asia. The richness of the West African fauna is Lygaeidae, West Africa, Drymini, Myodochini, zoogeography not only due to isolation within the region but also, presumably, for two additional reasons. First, the occurrence of a savanna corridor of long duration separating the West African forest fauna from that of Central Africa. Second, the distinction between the fauna of the wet tropical forest, both coastal and montane, from the increasingly arid sa- vanna and desert areas to the north. The savanna fauna is for the most part com- posed of species with wide ranges, many occurring in a broad arc around the rain forest, often all the way from Senegal to South Africa (see Mizaldus nidulus discus- sion below). There is also a southern Palearctic ele- ment that reaches the savanna in West Af- rica. When one, in addition, considers the frequent disjunction of the montane and lowland forests, the reasons for the richness and frequent endemicity of parts of the fau- na becomes evident. Students of West Africa have become in- creasingly interested in possible West Af- rican-South American relationships since the acceptance of continental drift. Such re- lationships are not evident in the rhyparo- 410 chromine fauna, at !east, at the present state of our knowledge. For example, the tribes Drymini and Rhyparochromini, which are dominant ciements in West Africa, are complete!y absent in South America and in the Lethaeini and Antillocorini, where such sister groups may exist, our knowledge is totally inadequate to demonstrate any close African-South American relationships. All measurements are in millimeters. Carvalhodrymus, NEw GENUS Body elongate, slender, parallel sided. Eyes set near middle of head, remote from anterior pronotal margin. Ocelli placed far behind posterior margins of eyes. Tylus acu- minate, extending only over basal one-fifth of first antennal segment. Vertex convex. Pronotum with a well-defined anterior col- lar; lateral pronotal margins very narrowly explanate, deeply incised at level of com- plete and deep transverse impression; hu- meral angles evenly rounded; posterior margin shallowly concave. Anterior prono- tal lobe convex, slightly higher than poste- rior lobe. Scutellum not elevated through- out but with a raised Y-shaped carina. Cla- vus with three rows of punctures, but punc- tures of inner row smaller and less closely spaced than those of outer two rows. Lateral corial margins straight, not expanded pos- teriorly. Apical corial margin straight. Metathoracic scent gland auricle short, bent caudad. Evaporative area small, occupying only area immediately around auricle with dorsal margin rounded. Fore femora slen- der, armed below with a single small, in- conspicuous spine near distal end. Body non- pruinose, glabrous or nearly so. Pronotal punctures on anterior and posterior prono- tal lobes equal in size and distribution. An- tennae extremely elongate, very slender, fi- liform, fourth segment not at all fusiform. Type species.—Carvalhodrymus elegans new species. By monotypy. Carvalhodrymus will run to couplet 14 in Slater’s (1993) key to African genera of Dry- mini, but it is not closely related to either PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON genus at that couplet. It is much more sim- ilar to the Palearctic genus Thaumastopus Fieber. Both of these genera are relatively elongate slender taxa with acuminate heads, elongated first antennal segments, coarsely punctate pronota, very narrowly explanate pronotal margins, eyes set well away from the anterior margin of the pronotum, a Y-shaped scutellar carina, and elongate fi- liform antennae. Carvalhodrymus differs from Thaumas- topus in lacking incrassate, multispinose fore femora, possessing straight rather than con- vex lateral corial margins, having com- pound eyes that are elliptical rather than round in dorsal view, by having greatly elongated antennae with both segments two and three much longer than the pronotal length (subequal in Thaumastopus). In the type species of Thaumastopus (marginicol- lis Lucas), the tylus nearly attains the middle of the relatively short first antennal segment that is shorter than or subequal to the pro- notal length (see Kiritshenko’s (1951) ex- cellent figure No. 340), whereas in Carval- hodrymus the tylus attains only the proxi- mal ¥ of the first antennal segment that is appreciably longer than the length of the pronotum. It is a great pleasure to name this striking new genus in honor of Dr. J. C. M. Car- valho. Carvalhodrymus elegans, NEw SPECIES (Fig. 1) Coloration uniformly yellowish-brown with strongly contrasting dark brown punc- tures on pronotum, scutellum and hemel- ytra. Membrane of forewing dark fumose, lacking a pale apical macula. Corium with a narrow, elongate, smooth, pale patch ad- jacent to radial vein at level of apex of scu- tellum. Hemelytra anterior to this macula slightly paler than corium distally (colora- tion difference marked by a straight trans- verse line). Legs and antennae uniformly pale yellow. Length head 1.06 (distance anterior to eye VOLUME 97, NUMBER 2 Fig. 1. Carvalhodrymus elegans n. sp. Dorsal view. 0.46, distance posterior to eye 0.28), width head 0.76, interocular space 0.40. Length pronotum 1.20 (length anterior lobe 0.66, posterior lobe 0.54), width pronotum 1.46. Length scutellum 1.00, width 0.80. Length 411 claval commissure 0.54. Midline distance apex clavus-apex corium 1.30. Midline dis- tance apex corium-apex abdomen 0.60. Length labial segments I 0.66, II 0.80, Il 0.46, IV 0.28 (approx.). Length antennal 412 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON segments I. 1.60, Il 1.76, III 1.44, IV 1.36 (approx). Total body length 5.60. Holotype.—é: Ghana: Sunjami. 12.XII.1965 (Leston) (UV trap). In Amer- ican Museum of Natural History. Paratype.-—1 6: Cameroon: Southwest Prov. 25 km. W. Limbe Bakingil1. 16.V1I.1984 (James Digiulio) (Malaise trap). In Carnegie Museum of Natural History. The paratype, while apparently not dif- fering structurally from the holotype, is very differently colored. The ground color is dark chocolate brown. The pale macula on the hemelytra near the lateral margin at the lev- el of the apex of the scutellum (which is only faintly differentiated in the holotype) is a sharply contrasting white in the paratype and the entire corium anterior to this mac- ula is much paler than the posterior area. The paratype has a small white area on the membrane adjacent to the apex of the co- rium. The third antennal segment is dark chocolate brown and contrasts strikingly from the pale yellow first and second seg- ments and the almost white fourth segment. The fore femora are dark red-brown. De- spite these striking color differences the ho- lotype does not appear to be teneral and there seem to be no significant structural differences. This apparently is a rare species, or at least comes very rarely to lights, as in the many thousands of West African Rhyparo- chrominae that I have examined these are the only two specimens known. Mizaldus Distant Distant 1901: 483. Mizaldus is a genus of small drymine ly- gaeids characterized by the deeply concave apical corial margins, elevated scutellum, lack of fore femoral spines, two rows of cla- val punctures, short head with large eyes and filiform antennae. Only two species previously have been known to occur in Africa, Mizaldus nidulus Slater and Carayon from Guinea in West Africa and Mizaldus sinuaticollis Linna- vuori from the southern Sudan in East Af- rica. Specimens are rare in collections. Appar- ently, species of this genus live in special- ized habitats and come only rarely to lights. In the present paper four new species are described and additional distributional data are given for previously described species. Characters that appear to be most useful in separating the various species are the shape of the scutellum, the nature of the scutellar punctures, the relative thickness of the antennae and, in some cases, the col- oration of these segments, the size and shape of the eyes, and the pronotal proportions. While there appear to be differences in the pruinosity patterns on the pronotum, a more adequate series will be necessary before the utility of this suite of characters can be de- termined. Scudder (1968) established the genus Neomizaldus with Mizaldus lewisi Distant as the type species. His lengthy generic de- scription consists almost entirely of features that are true of both taxa. Scudder notes that the abdominal sterna of species of Neomizaldus are longitudinally striate. This condition is found in all of the African spe- cies of Mizaldus (although obscured by long hairs in Mizaldus tenuis). Scudder’s chief differentiating character was said to be the presence ofa distinct collar in Neomizaldus. He notes that this collar is not “really evi- dent” mid-dorsally, but is distinct laterally. I am very dubious of the desirability of rec- ognizing Neomizaldus as a distinct genus. The collar is very poorly developed in the type species of Scudder’s genus, only slight- ly more so than in some of the African species of Mizaldus. On the other hand, Neomi- zaldus lewisi has a very distinctive scutel- lum with a broad subbasal transverse smooth raised area that is not found in any Mizaldus known to me. However, given the striking differences in the conformation of the scutellum in some of the African species of Mizaldus, this seems questionable as a generic feature. Clearly an investigation of VOLUME 97, NUMBER 2 the Oriental elements of these two taxa is in order. The only published information on the biology, of which I am aware, is that of Sla- ter and Carayon (1963) who reported M. nidulus from the nests of ploceid finches in Guinea, where they occurred with antho- corids and were apparently feeding on small arthropods. This species was reared on Co- leoptera larvae in the laboratory. Such a feeding habit may not be true of other spe- cies of the genus. Mizaldus nidulus is rela- tively weakly sclerotized as compared with the new species described below and the hemelytral punctures, especially those on the clavus, are pale and little differentiated in color from the surrounding surface. By con- trast, the other West African species have hemelytral punctures that are distinctly dif- ferentiated. This suggests that possibly M. nidulus, being associated with cryptic hab- itats (bird nests), has lost some of its scle- rotization, whereas the other species may be living in less protected environments. Distribution.— While previously known on the continent only from West and East Africa the genus will probably be found to be widespread throughout much of tropical and subtropical Africa. As noted below M. nidulus is now known to occur in Nigeria and both on the high veld and in the tropical corridor in South Africa. The unpublished field notes from Ghana of the late Dennis Leston indicate that specimens upon which several of the new species are based were taken in dense forest by pyrethrum knock- down technique, whereas M. nidulus is probably a savanna species. This belief is consistent with the discovery of the latter in South Africa. Such a distribution, as not- ed previously, suggests a range through a wide savanna arc from West to South Africa around the central African rain forest. Mizaldus lestoni, NEw SPECIES (Figs. 4, 5) Body relatively elongate, tapering ante- riorly. Head, pronotum, scutellum, large 413 apical corial macula, basal 34 of first anten- nal segment and femora black. Hemelytra white, central area hyaline. A small, narrow, elongate, dark-brown vitta present on distal 2 to “3 of radial vein. Membrane pale trans- lucent basally, contrastingly fumose from level of dark corial macula to apex. Hemely- tral punctures dark brown, strongly con- trasting with pale ground color. Base and distal end of first antennal segment and all of segments two and three pale yellow (seg- ment four missing). Pleural and sternal sur- faces nearly uniformly dark chocolate brown, with posterior lobe of metapleuron white. Tibiae and tarsi yellowish brown, tib- iae somewhat darker proximally. Head shining. Pronotum and scutellum nearly completely dull pruinose, obscurely and finely punctate. Head and anterior pronotal lobe clothed with numerous decumbent sil- very hairs. Head sharply acuminate, not declivent. Tylus extending to middle of first antennal segment; vertex strongly convex; eyes ses- sile, occupying most of lateral head surface but not strongly globose. Ocelli placed much nearer eyes than meson. Length head 0.60, width 0.65, interocular space 0.40. Prono- tum with lateral margins narrowly but dis- tinctly carinate, strongly sinuate; transverse impression complete; posterior margin straight before base of scutellum, laterally lobately produced. Length pronotum 0.78, width 1.08. Scutellum evenly elevated, lack- ing a median carina, punctures distinctly separated from one another. Length scutel- lum 0.58, width 0.53. Length claval com- missure 0.13. Lateral corial margins strong- ly sinuate. Midline distance apex clavus- apex corium 0.70. Midline distance apex corium-apex abdomen 0.88. Length labial segments I 0.38, II 0.40, III 0.43, IV 0.33. Antennae relatively robust, chiefly terete but with second and third segments slightly en- larged from proximal to distal ends. Length antennal segments I 0.30, II 0.62, III 0.50 (IV missing). Total body length 3.60. Holotype.—?: Ghana: Eastern Region, Atewa Range Forest Reserve. 25.VII.1969 414 (D. Leston) (pyrethrum knockdown). In American Museum of Natural History. This is a relatively large species, generally similar to Mizaldus linnavuorii but with a completely differently shaped scutellum that in this species has small inconspicuous punctures. The antennal segments are much thicker, the eyes less protrudant and more elongate and the femora are completely dark on all legs. The Atewa Range Forest Reserve is ma- ture primary forest, the upper story with a largely complete canopy. This species 1s dedicated to the memory of the late Mr. Dennis Leston for his in- valuable West African collections and for his important contributions to our knowl- edge of the world Hemiptera. Mizaldus linnayuorii, NEw SPECIES (Fig. 2) Body robust. Head, pronotum, scutellum, narrow apex of corium and third and fourth antennal segments black. First antennal seg- ment infuscated with red-brown on basal half. Corium and clavus white, or very pale yellow, lacking a transparent central area. Fore femora and distal 2 of middle and hind femora red-brown; remainder of legs and antennae pale yellow. Upper ' of pos- terior metapleural lobe white. Head shining. Pronotum and scutellum completely prui- nose. Head and pronotum finely and evenly punctate. Scutellar punctures large, coarse and anastomosing. Membrane hyaline, uni- colorous. Body almost completely glabrous above. Head acuminate, tylus reaching to distal ’; of first antennal segment. Eyes large, glo- bose, protruding. Length head 0.56, width 0.64, interocular space 0.36. Pronotum rel- atively broad, posterior lobe elevated above anterior; lateral margins very narrowly car- inate, sinuate; transverse impression com- plete. Length pronotum 0.84 (length ante- rior lobe 0.36, length posterior lobe 0.48), width 1.12. Scutellum with a sharp ovate marginal carina, the margins meeting pos- teriorly in an arc on meson. Scutellar disc PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON convex basally and mesally, somewhat de- pressed distally, with a very weak carina near distal end. Length scutellum 0.56, width 0.60. Lateral corial margins relatively shal- lowly concave. Length claval commissure 0.14. Midline distance apex clavus-apex co- rium 0.72. Midline distance apex corium- apex abdomen 0.62. Labium extending pos- teriorly well between mesocoxae. Length la- bial segments I 0.36, II 0.36, III 0.36, IV 0.30. Antennae elongate, very slender, fourth segment at most narrowly fusiform. Length antennal segments I 0.28, II 0.62, III 0.60, IV 0.54. Total body length 3.12. Holotype.—é: Ivory Coast: Mt. Tonkoui. 15—22.X.1973 (Linnavuori). In American Museum of Natural History. Paratype.—1 ?: Ivory Coast: Adiopo- doume. 29.[X-7.X.1973 (Linnavuor). In J. A. Slater collection. This species is readily recognizable by its large size, robust body form, elongate slen- der antennae with the third and fourth seg- ments black, and especially by the coarsely punctate, elevated and uniquely carinately margined scutellum. The paratype differs from the holotype in having the posterior pronotal lobe and the scutellum somewhat subshining rather than being completely dull pruinose. Mizaldus linnavuorii appears to be most closely related to Mizaldus carvalhoi as dis- cussed under the latter. It is a pleasure to dedicate this species to Dr. Rauno Linnavuori in recognition of his unparalled West African collections and his many contributions to our knowledge of Af- rican Hemiptera. Mizaldus carvalhoi, NEw SPECIES (Fig. 7) Body robust. Head, anterior pronotal lobe, apical corial macula and fourth antennal segment black. Posterior pronotal lobe red- brown. Clavus and corium white, latter with central area subhyaline. Membrane uni- formly translucent hyaline. Fore femora and distal half of middle and hind femora red- brown. Basal half of antennal segment one VOLUME 97, NUMBER 2 415 Ny . Ny ‘ NY Figs. 2-6. 2. Mizaldus linnavuori n. sp. Dorsal view of head and pronotum. 3. Mizaldus tenuis n. sp. Dorsal view of head and pronotum. 4. Mizaldus lestoni n. sp. Dorsal view of head and pronotum. 5. Mizaldus lestoni n. sp. Dorsal view clavus and corium. 6. Mizaldus tenuis n. sp. Dorsal view clavus and corium. 416 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Mizaldus carvalhoi n. sp. Dorsal view. VOLUME 97, NUMBER 2 and suffused proximal half of antennal seg- ment 3 pale yellowish brown. Remainder of legs and antennae pale yellow. Upper por- tion of posterior metapleural lobe white. Head shining. Pronotal calli, posterior pronotal lobe (with exception of a large quadrate pruinose area on either side of midline immediately behind transverse im- pression) and scutellum dull to subshining, but contrasting with pruinosity of anterior pronotal lobe. Head, pronotum, and scu- tellum with distinct well-separated punc- tures, those on scutellum not anastomosing. Dorsal surface almost glabrous, no elongate hairs present. Head broad, sub-acuminate; eyes glo- bose, protruding; vertex only slightly con- vex. Length head 0.50, width 0.60, interoc- ular space 0.34. Pronotum broad, posterior lobe slightly elevated above anterior; lateral margins narrowly carinate, sinuate; trans- verse impression complete, but relatively shallow. Length pronotum 0.74 (length an- terior lobe 0.36, length posterior lobe 0.38), width 0.98. Scutellum with lateral margins carinate, tapering posteriorly to a V-shaped apex; an obtuse median carina present; bas- al half of surface somewhat elevated. Length scutellum 0.43, width 0.46. Lateral corial margins shallowly concave. Length claval commissure 0.14. Midline distance apex clavus-apex corium 0.68. Midline distance apex corium-apex abdomen 0.62. Labium reaching between mesocoxae. (Approxi- mate labial lengths from paratype I 0.35, II 0.38, III 0.35, IV 0.22.) Antennae slender, filiform, third segment very slightly clavate, fourth segment narrowly fusiform. Length antennal segments I 0.30, II 0.54, HI 0.50, IV 0.46. Total body length 2.52. Holotype.—é: Ghana: Nsemre Forest Reserve, Brog Ahafo 13.XII.1965 (D. Les- ton). In American Museum of Natural His- tory. Paratypes.—Ghana: 2 22 same data as holotype. 1 6 Tafo, Eastern Region 23.V.1966 (D. Leston) (pyrethrum knock- 417 down block, R1 cocoa). In J. A. Slater col- lection. This species is closely related to Mizaldus linnavuorii. The type series is considerably smaller with a differently shaped scutellum in which the punctures are small and dis- crete, rather than coarse and anastomosing, and the marginal scutellar carina does not meet in a semi-circular curve near the pos- terior end of the scutellum. The fourth an- tennal segment is also relatively longer (sub- equal in length to segment three rather than being appreciably shorter). The pruinosity patterns of the two species and the pale third antennal segment of the present species all appear to be distinctive features. I have examined an additional female from Nigeria: (SE. St. Obudu Cattle Ranch 16-18.VIII.1973 (Linnavuori), which I be- lieve to be conspecific although it is consid- erably larger. Because this species and M. linnavuorii are more closely related to one another than most of the other species of Mizaldus described here, the measurements of this Nigerian specimen are given to fa- cilitate relationships when more extensive material becomes available. Length head 0.58, width 0.72, interocular space 0.38. Length pronotum 0.90 (length anterior lobe 0.44, posterior lobe 0.46), width 1.18. Length scutellum 0.52, width 0.64. Length claval commissure 0.16. Midline distance apex clavus-apex corium 0.70. Midline dis- tance apex corium-apex abdomen (0.80. Length antennal segments I 0.34, II 0.64, III 0.58, IV 0.60. Total body length 3.40. Mizaldus tenuis, NEw SPECIES (Figs. 3, 6) Body relatively elongate, nearly parallel sided. Head, pronotum, scutellum, a large apical corial macula and pleural and ventral surfaces uniformly black. A brown quadrate macula present on corium at level of pos- terior end of claval commissure extending inward from lateral margin to raised radial 418 vein (Fig. 6). Antennal segments one, two, and three pale yellow, base of segment one weakly infuscated, segment 4 a strongly contrasting dark brown. Fore femora and distal half of middle and hind femora red- dish, remainder of legs pale yellow. Poste- rior metapleural lobe white. Head, prono- tum, and scutellum with numerous shallow, evenly separated punctures. Head and an- terior pronotal lobe subshining. Posterior pronotal lobe and scutellum dull pruinose. Head dorsally with a few elongate hairs. Re- mainder of dorsal surface almost glabrous (minute decumbent hairs arising from punctures). Head strongly declivent. Eyes very large, occupying almost entire lateral surface of head. Tylus extending over basal '% of first antennal segment. Ocelli placed much clos- er to eyes than to meson. Length head 0.60, width 0.70, interocular space 0.36. Prono- tum relatively long and narrow, length and width subequal. Anterior pronotal lobe lon- ger (0.50) than posterior lobe (0.40). Lateral pronotal margins very narrowly carinate and strongly sinuate; transverse impression complete and very deep; anterior lobe even- ly convex. Length pronotum 0.90, width 0.90. Scutellum evenly elevated; without a strong marginal carina. Length scutellum 0.46, width 0.44. Lateral corial margins deeply and evenly concave, maximum width near apex of corium. Length claval com- missure 0.16. Midline distance apex clavus- apex corium 0.60. Midline distance apex corium-apex abdomen 0.54. Labium ob- scured, at most reaching onto mesosternum. Antennae moderately slender, third seg- ment somewhat clavate, fourth segment fu- siform. Length antennal segments I 0.22, II 0.44, III 0.38, IV 0.44. Total body length 2.96. Holotype.—é: Ivory Coast: Adiopo- doume. 29.1X-7.X.1975 (Linnavuori). In American Museum of Natural History. This species is readily recognizable by the relatively elongate, narrow pronotum and other characters as noted in the following key. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mizaldus sinuaticollis Linnavuori Linnavuori 1978: 86. This species was originally described from a female from “‘Equatoria: Yei-Iwatoka.”’ I have examined a male from Kenya: “Li- muru, 3-1V.19355 (D:C.. Thomas) (dead leaves)” in the C. J. Drake collection at the National Museum of Natural History (USNM) that I believe to be conspecific. I have keyed this species out to two places in the following key because of uncertainty as to the condition of the scutellum. Lin- navouri (1978) states that in M. sinuaticollis the scutellum is “densely punctate.” I take it that this means that the scutellar punc- tures are large and anastomosing as they are in M. linnavuorii. In the Kenya specimen before me, the scutellar punctures are large and dense, but they are not anastomosing and the scutellum thus differs in this regard from M. linnavuorii. I have separated the species twice in the following key once on the basis of antennal length in case the scu- tellar punctures of the type specimen are not anastomosing and once where they would be. The Kenya specimen also has the third and fourth antennal segments somewhat darker than the two proximal segments. It is certainly related in general habitus to M. linnavuorii and a series from intervening localities is very desirable. Mizaldus nidulus Slater and Carayon Slater and Carayon 1963: 1-4. This species is readily recognizable by the dark third antennal segment and relatively weak sclerotization in which the hemelytral punctures are nearly concolorous with the surface of the clavus and corium. Mizaldus nidulus was originally described from Guinea but is apparently widespread in Africa, although rare in collections. I have examined the following additional material: Nigeria: 1 6 Zugurma NW state XII.1974 (J. T. Medler). South Africa: Transvaal: 1 2 Lyttelton 29.11.1968 (J. A. & S. Slater) (UV VOLUME 97, NUMBER 2 light). Natal; 1 2 Umtentweni VII.1953 (A. L. Capener). 1 2 Umtentweni VII.1954 (A. L. Capener). In J. A. Slater collection. KEY TO AFRICAN SPECIES OF MIZALDUS 1. Third antennal segment dark red-brown to black, strongly contrasting with pale coloration of second antennal segment ............... 2 — Third antennal segment pale yellowish, con- colorous or nearly so, with coloration of second antennalasegmentemeaemrn tye eae era 3 2. Fourth antennal segment pale yellow, strongly contrasting with dark third segment and con- colorous with antennal segment two; claval punctures obscure, concolorous with pale ground colorof- hemelytra soo... Aen oe MP wis es eer i oe nidulus Slater and Carayon — Fourth antennal segment black, concolorous with dark third antennal segment and strongly contrasting with pale second antennal segment; claval punctures dark brown, strongly contrast- ing with pale hemelytral ground color ...... MOY ete Nee t 9 1 ee linnavuorii n. sp. 3. Scutellar punctures large, irregular and anas- LOMOSING IAA ee sinuaticollis Linnavuori — Scutellar punctures small, distinctly separated from one another and not anastomosing .... 4 4. Head less than 1% times as broad aslong... 5 Head 1'2 or more times as broad as long ... 6 5. Antennal segment two much less than 1.2 times lengthiofsepment threes(5:13)ie5. 5. o> sea fo A EE ee sinuaticollis Linnavuori — Antennal segment two more than 1.2 times lengthtotsezment three: 4s... 2. lestoni n. sp. 6. Pronotal length subequal to width of pronotum across humeral angles; a large quadrate brown macula present at level of distal end of claval commissure laterad of radial vein ... fenuis n. sp. — Pronotum wider than long; hemelytra lacking a dark macula laterad of radial vein ........ nish & Aen A RE ES SERS Sal carvalhoi n. sp. Fontathanus Scudder Scudder 1963: 1233-1234. Scudder (1963) erected the genus Fonta- thanus to include four African species. Three of these were known only from Zaire and the fourth from Zaire, Guinea, and Sierra Leone. Scudder related Fontathanus to Fontejus Stal. However, Sweet (1967) placed the lat- ter genus in the Udeocorini, whereas Fon- tathanus is a member of the Myodochini 419 and the relationship presumably was on the basis of myrmecomorphy. Scudder sepa- rated the two genera by Fontethanus having unarmed fore tibiae, a deeper and more an- teriorly placed transverse pronotal constric- tion and said that “the coloration is differ- entae Harrington (1980) added to this diagnosis of Fontathanus the following: having “‘phal- lic type I,” the posterior edge of the pygo- phore “subsharp,” four or more rows of cla- val punctures, a V-shaped buccular junc- ture, double-ranked fore femoral spines, and an enclosed mesepimeron. Only one of Scudder’s species has a punc- tate anterior pronotal lobe and, even in this species, the punctures are said to be present only mesally and laterally. Both new species described below have completely punctate pronota. They are both strikingly myrme- comorphic with broad ant-shaped heads, as evidenced by the carinate lateral margins of the head before the eyes that seems to sim- ulate the edges of mandibles. The color pat- tern viewed laterally enhances the antlike resemblance by the white of the posterior metapleural lobe being continued onto the proximal portions of the hind femora to ‘break up” the outline and form a pseu- dopedicel appearance. Fontathanus rostratus, NEW SPECIES (Fig. 8) Shining jetblack above on head and pro- notum. A narrow gray pruinose stripe through transverse pronotal impression. Prothorax laterally and ventrally, acetabula, and posterior metapleural lobe gray prui- nose. Silvery pruinosity present as follows: scutellum (with exception of raised apex), a large basal corial patch within radial vein that extends onto adjacent area of clavus, a second patch on corium also within radial vein at level of anterior half of claval com- missure. Corium variegated, a small pale yellow macula basally at lateral margins and a large yellow patch laterad of radial vein in center of corium, mesal portion of which 420 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Fontathanus rostratus n. sp. Dorsal view. becomes bright yellowish orange and is bor- dered both anteriorly and posteriorly at lat- eral ends with a dark brown stripe or vitta. Adjacent to bright yellow area mesad of ra- is TA pees teh dius an additional irregular bright yellow marking, this of irregular outline, almost forming three elongate spots. Remainder of clavus and corium velvety in texture and of VOLUME 97, NUMBER 2 an (almost black) dark red color. Membrane uniformly dark fumose, pruinose proxi- mally within apical corial margins. Extreme postero-dorsal corner of metapleuron and subproximal band on middle and hind fem- ora pale yellow to almost white. Extreme distal end of fore femora and entire foretib- iae pale orange-yellow. Middle tibiae black at proximal ends, becoming orange-yellow distally. Hind tibiae chocolate brown. All tarsal segments and first three antennal seg- ments pale yellow. First antennal segment infuscated with dark brown basally, fourth segment dark chocolate brown. Head and pronotum finely but conspicuously punctate over entire surface. Body surface finely sha- greened, nearly glabrous but with very short fine decumbent hairs arising from punc- tures. Head moderately declivent, tapered an- teriorly. Head anterior to eyes produced and carinate. Eyes sessile. First antennal seg- ment almost attaining apex of tylus. Ocelli placed far laterad, near inner corner of com- pound eyes. Length head 0.98, width 1.32, interocular space 0.86. Pronotum with deeply incised transverse impression. An- terior pronotal lobe rounded, ovate, longer than posterior lobe (length anterior lobe 0.86, length posterior lobe 0.58). Posterior pronotal margin nearly straight; lateral mar- gins extremely deeply sinuate. Length pro- notum 1.48, width 1.68. Scutellum with a very slight median elevation, posterior end sharply carinate. Length scutellum 0.80, width 0.64. Length claval commissure 0.50. Corium with lateral margins strongly sin- uate, narrowest at level of distal portion of scutellum, slightly explanate posterior to this area and very slightly recurved. Midline dis- tance apex clavus-apex corium 0.96. Mid- line distance apex corium-apex membrane 0.98. Abdomen strongly constricted basal- ly. Metathoracic scent gland auricle ellip- tical, short. Evaporative area rugulose, cov- ering almost entire anterior metapleural lobe. Fore femora strongly incrassate, armed below with 2 rows of conspicuous sharp 421 spines, outer row consisting of 2 or 3 major spines with 3 small spines distad of outer large spine; inner row near middle with one extremely large spine and 4 distal small spines. Foretibiae somewhat curved. Labi- um elongate, reaching third abdominal ster- num. First labial segment exceeding base of head. Length labial segments I 0.90, II 0.90, III 0.78, IV 0.62. Antennae slender, seg- ments two and three slightly enlarged near distal ends, fourth segment strongly fusi- form. Length antennal segments I 0.28, II 0.68, III 0.62, IV 0.94. Total body length 5:0: Holotype.—?: Nigeria: Lagos, Olatunde Ayoola Av. 6.III.1975 (Abdul Hamid) (light trap). In American Museum of Natural His- tory. Paratypes.— Nigeria: 1 6: same data as holotype. 1 6 R. St. nr. Mbiama 4- 5.VII.1973 (Linnavuori). 1 2 R. St. Ebubu nr. Bori 2. VII.1973 (Linnavuori). 4 6, 2 9¢ Lagos, Glatunda Ayoola Ave. V.1975 (A. Hamid). Ivory Coast: 1 6 Dunco (sp. ?) For- est 7.X.1973 (Linnavuorli). In National Mu- seum of Natural History (USNM), A. Ha- mid and J. A. Slater collections. I have examined an additional large fe- male from the Cameroon (Abong Mbang- Ayos 13.VI.1973 [Linnavuori]), which is ei- ther conspecific or represents a closely re- lated species. In addition to its larger size this specimen has the pale yellow and or- ange-yellow markings on the hemelytra much more extensively developed so that the corium is predominately pale rather than dark. The membrane by contrast is uni- formly dark and all tibiae, tarsal segments two and three, and antennal segments two, three and four are uniformly dark red- brown. Fontethanus rostratus runs in Scudder’s (1963) key to Fontethanus punctatus Scud- der because of the distinct punctures present on the anterior pronotal lobe. It is appar- ently not closely related to F. punctatus, the latter having slightly ochraceous humeral pronotal angles, only a pale triangular sub- 422 apical corial spot, and has a pale area lat- erally on abdominal sternum five. In F. punctatus the \abium reaches only to the hind coxse and the anterior pronotal lobe is distinctly punctate only mesally and lat- erally, rather than over the entire surface. Fontathanus ghanaensis, NEw SPECIES General form and color similar to F. ros- tratus. Pale areas on lateral portion of co- rium reduced to 3 irregular spots mesad of radial vein and 2 elongate white lateral streaks. Middle and hind femora lacking pale proximal annulations, completely black. First antennal segment black above, with red-brown coloration laterally. Body col- oration, punctures and pubescence as in F. rostratus. Corium lacking shining silvery pruinose bars adjacent to claval suture. Head strongly declivent, very broad, margins of head anterior to eyes strongly produced laterad and very sharply carinate. Length head 1.02, width 1.55, interocular space 1.0. Anterior pronotal lobe broadly obovate, maximum width equaling width across humeri; transverse impression ex- tremely deep. Length pronotum 1.55 (length anterior lobe 0.78, length posterior lobe 0.75), width 1.42. Length scutellum 0.88, width 0.72. Length claval commissure 0.40. Clavus with a pruinose area occurring di- agonally entirely across clavus near middle. Midline distance apex clavus-apex corium 1.02. Midline distance apex corium-apex membrane 1.12. Metathoracic scent gland auricle relatively slender, elongately ellip- tical, slightly bent posteriorly. Fore femora similar to rostratus. Labium relatively short, extending only slightly beyond mesocoxae, first segment not attaining base of head. Length labial segments I 0.68, II 0.62, III 0.60, IV 0.50. Length antennal segments I O:25.11-0.70; 11 0.58; DV. 10; Total body length 5.50. Holotype.—¢: Ghana: Tafo 9.X.1965 (D. Leston). In American Museum of Natural History. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON This species is very similar to F. rostratus in both form and color. It is most readily recognizable by the relatively short labium that does not reach the metacoxae, in con- trast to F. rostratus where the labium ex- tends well back onto the anterior portion of the abdomen. Also in F. ghanaensis the head is much broader and the produced lateral margins anterior to the eye are much more produced and prominent. Both of these species are strikingly myr- mecomorphic. The corial color pattern, striking constriction between the pronotal lobes, constricted base of the abdomen, and the pruinose patches are all presumably ant mimetic adaptations. Paromius carvalhoi, NEw SPECIES Elongate, slender, parallel sided. Head black. Anterior pronotal lobe dark gray to nearly black; posterior lobe red-brown, be- coming pale testaceous yellow on humeral angles and near posterior margin on either side of midline. Scutellum nearly uniformly red-brown, apex white. Hemelytra nearly uniformly pale testaceous yellow, with punctures strongly contrastingly dark brown; explanate margins dull white. Membrane pale fumose with veins strongly contrasting white. Abdomen red-brown. Femora vary- ing from dark to bright red-brown. Tibiae and first two tarsal segments yellow, third tarsal segment black. Punctuation typical for genus, very fine and obscure on anterior pronotal lobe, except collar where more prominent. Head moderately acuminate, eyes set far from posterior margin, prominently swol- len, tylus extending midway to distal end of first antennal segment, very slightly decli- vent. Length head 0.92, width 1.05, inter- ocular space 0.58. Pronotum elongate, ta- pering, anterior lobe not strongly convex dorsally, not appearing globose; lateral mar- gins sinuate; transverse impression com- plete; posterior lobe with a raised median elevation on anterior half. Length prono- tum 1.58, width 1.50. Scutellum elevated VOLUME 97, NUMBER 2 mesally, strongly sloping laterad. Length scutellum 1.15, width 0.72. Midline dis- tance clavus-apex corium 1.80. Midline dis- tance apex corium-apex membrane 1.45. Fore femora moderately incrassate, armed below with two rows of white, dark tipped spines with four to five spines and spinules present in each row. Labium elongate ex- tending between mesocoxae. First segment not quite attaining base of head. Length la- bial segments I 1.05, II 1.05, III 0.80, IV 0.38. Antennae conventionally terete, slen- der. Length antennal segments I 0.52, II 1.42, III 1.18, IV 1.48. Total body length 8.00. Holotype.—é: Ghana: Tafo 1.IV.1967 (D. Leston). In American Museum of Nat- ural History. Paratypes. — Ghana: lieeercheary Lato 12.XI1I.1965 (Leston) (UV trap). 1 ¢ same Exceptr2 5511966, 1 Bo ysame) (except 15.XII.1965. 1 2 same except 16.XI.1965. 1 6 Accra 1.XII.1969 (C. W. Campbell) (blacklight trap). In National Museum of Natural History (USNM) and J. A. Slater collections. Many of the species of Paromius are very similar in habitus to one another. The genus is certainly in need of a revisional study. However, only three species of Paromius have the labium elongate 1.e. extending to, or nearly to, the mesocoxae. In all other species the labium is appreciably shorter, barely extending onto the anterior portion of the mesosternum. Of the three species in which the labium 1s elongate, the Ethiopian species Paromius apicatus Stal differs from P. carvalhoi by being much larger, having a more elongate head, a strongly tapered pro- notum, and a very much longer labium that in P. apicatus extends well onto the abdo- men. Paromius carvalhoiis much more sim- ilar in general habitus to the Oriental species Paromius piratoides (Costa), which it close- ly resembles. However, the two are certainly distinct in that P. piratoides has a relatively shorter more strongly declivent head, a more prominently, convexly rounded, anterior 423 pronotal lobe, lacks the median elevation of the posterior pronotal lobe, and has many less claval punctures mesad of the cubital vein. Also in most specimens of P. pira- toides, only the apex of the third tarsal seg- ment is black, whereas in P. carvalhoi the entire third tarsal segment of all legs is black. Actually P. carvalhoi may prove to be more closely related to P. apicatus, despite the more readily apparent morphological differences. Both of these species have an elevated mesal posterior line on the pro- notum and elongate attenuated pronota and heads. Both Paromius paraclypeatus Scudder (1969a) and P. carvalhoi have been con- fused in the literature with Paromius gracilis (Rambur). I have not seen authentic spec- imens of P. gracilis from Ghana, although it is probable that it occurs in the northern savanna. Paromius gracilis is an abundant species in eastern and southern Africa. Esinerus Scudder Scudder 1969b: 88-90. Scudder (1969b) separated Esinerus from Sinierus Distant by the former lacking a pronotal collar, having a relatively short first antennal segment, a dull tylus, a more flat- tened scutellum, and with less prominently erect body hairs. The new species described below neces- sitates some modifications of the generic di- agnosis, because it has considerably longer hairs than does the type species of Esinerus (refractarius Scudder) and has a pronotal collar that is nearly as well developed as it is in species of Sinierus. The relatively flat- tened scutellum, short first antennal seg- ment, and dull tylus will still separate the two genera but are questionable features for generic status. However, I believe that Fs- inerus and Sinierus should be retained as distinct genera, as there are three morpho- logical characters not mentioned by Scud- der that seem to be important. In species of Esinerus the bucculae are well developed 424 along the entire veniral surface of the head with a groove present between them so that the first labial segment lies immersed be- tween the bucculae and scarcely extends out of the b lar groove. In Sinierus the buc- culae are !ow, obsolete posteriorly and the first labial segment is not enclosed by the bucculae for most of its length. In Esinerus the compound eyes do not have a series of large conspicuous hairs protruding from them as do the species of Sinierus. The metathoracic scent gland auricle of species of Sinierus is produced well above the sur- rounding evaporative area and has an oval buttonlike shape, whereas in Esinerus the auricle is not raised above the evaporative surface and curves evenly posteriorly as a narrow lunate arc. Esinerus species have a longitudinal row of silvery hairs running through the center of the clavus. They are not present in species of Sinierus. These sil- very hairs are rather obscure in E. refrac- tarius, but are very conspicuous in FE. humi- dus. My recent (Slater 1993) key to the African genera of Drymini is inaccurate. Couplet 1 of that key separates the genera on the basis of elongate hairs on the dorsal surface. Both Esinerus and Sinierus have numerous up- standing hairs present but are keyed to the half of the couplet where such hairs are lack- ing. It is true that the vestiture of these gen- era 1s much less elongate and upstanding than in the genera keyed that way, but any- one using the key would surely be misled by this couplet. Also couplet 3 states that the apical corial margin of Esinerus must be deeply concave. It is true that Esinerus species do have a weakly concave area near the base of the apical corial margin, but this is easily overlooked and certainly is not “deeply concave” in the sense that it is used in Mizaldus. The Slater (1993) key will work much better if Sinierus Distant (1901) and Esinerus are carried to couplet 2 where they may readily be separated from Parastilbo- coris Carayon (1964) and Psilomydrus Scudder (1969b), the other two genera with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON elongate dorsal hairs, by having the dorsal body surface pruinose rather than shining or subshining. Esinerus humidus, NEw SPECIES Head uniformly gray, shading to dark gray-brown on anterior pronotal lobe and basal third of scutellum. Posterior pronotal lobe, posterior two-thirds of scutellum and distal half of corium bright red-brown. Cla- vus, anterior half of corium, a large macula on membrane adjacent to apex of corium and an elongate macula distally in middle of membrane yellow. Remainder of mem- brane black. Legs pale testaceous. Antennae uniformly dark brown. Head and anterior pronotal lobe clothed with conspicuous vel- vety pruinosity, remainder of body surface also pruinose but less conspicuously so. Dorsal surface thickly clothed with upright, or semidecumbent, yellowish hairs. Ante- rior and posterior pronotal lobes, scutellum and hemelytra densely and finely punctate. Head slightly declivent, tylus greatly ex- ceeding juga, reaching midway to distal end of first antennal segment. Eyes set slightly away from antero-lateral pronotal angles. Length head 0.32, width 0.60, interocular space 0.40. Pronotum with a distinct an- terior collar, lateral margins deeply sinuate and obtusely carinate; posterior margin straight or very slightly concave before scu- tellum; transverse pronotal impression ob- solete mesally; calli low and faintly differ- entiated from remainder of pronotal disc. Length pronotum 0.52, width 0.98. Scutel- lum somewhat impressed basally, flat over most of surface, lacking a median elevation. Length scutellum 0.52, width 0.55. Length claval commissure 0.20. Corium with lat- eral margins somewhat sinuate, narrowest at level of distal third of scutellum; apical corial margin slightly concave near inner end. Midline distance apex clavus-apex co- rium 0.58. Midline distance apex corium- apex membrane 0.52. Metathoracic scent gland auricle appearing to be bilobed (a con- ventional posteriorly curving lobe present, VOLUME 97, NUMBER 2 but with also an ovoid raised anterior but- tonlike lobe nearly reaching anterior margin of metapleuron). Evaporative area small, not extending dorsally to middle of metapleu- ron, its dorsal margin truncate but rounded posteriorly. Metapleuron completely dull, covered with short pruinose hairs in addi- tion to the elongate upright hairs. Labium extending to mesocoxae. Length labial seg- ments I 0.30, II 0.28, II 0.25, IV obscured. Antennae relatively slender, second seg- ment terete, segments three and four fusi- form. Length antennal segments I 0.25, II 0.50, HI 0.38, IV 0.35. Total body length 250: Holotype.—é: Ghana: Eastern region forest reserve, Nkwanda 28.X.1967 (D. Leston). In American Museum of Natural History. Paratype.—[no abdomen]. Ghana: Tafo 17.X.1965 (D. Leston). In J. A. Slater col- lection. Esinerus humidus is readily separable from E. refractorius in being considerably more elongate and slender and having a dis- tinct anterior pronotal collar, with much more elongate upstanding hairs, lacking a white fourth antennal segment, with obtuse lateral pronotal margins (in contrast to E. refractorius where the margins are narrowly but sharply carinate), and in having a much longer, and relatively more slender, second antennal segment. The presence of three large pale spots on the membrane of the forewing in this species also occurs on many species of forest rhy- parochromines from West Africa. Whether this is a deflective color marking for species living on the forest floor should certainly be investigated. The biology of E. humidus is unknown. Our collecting party (Schuh, J. & S. Slater, Sweet) took many specimens of E. refrac- torius in South Africa feeding on the fallen seeds of Ficus sycamorus L. I have examined specimens from the Cameroon, which probably represent an un- described species. These specimens have a 425 pale fourth antennal segment, lack the large pale spots on the membrane of the forewing, have the metapleuron subshining with elon- gate hairs present, and the middle of the abdominal sternum shining. ACKNOWLEDGMENTS I am especially indebted to the following individuals who through their assiduous collecting have made serious study of the West African lygaeid fauna possible: Dr. Abdul Hamid who brought together a large amount of Nigerian material from the coastal area around Lagos and from the in- terior savanna. The late Dennis Leston who collected carefully in central Ghana and who first stimulated my serious interest in this fauna; Dr. Rauno Linnavuori whose metic- ulous collecting through West Africa re- sulted in the discovery of a rich fauna much of which was previously unknown. Dr. John Medler who collected for years in Nigeria and made his material available to me. I am also indebted to Mr. A. L. Capener for the gift of many species from South Af- rica; to Mr. Steven Thurston (formerly U. of Connecticut) for producing the dorsal view illustrations; to Dr. Roy Danielsson (Lund University) for making material from the Cameroons and Gambia available for study; to Dr. R. C. Froeschner of the Na- tional Museum of Natural History (USNM) and Dr. Randall T. Schuh (American Mu- seum of Natural History) for the loan of specimens; to the Plant Protection Institute Pretoria and especially the late Dr. W. G. H. Coaton for making facilities available, and to Dr. Schuh and my son Samuel Slater for much assistance in the field in South Africa. LITERATURE CITED Carayon, J. 1964. La spermathéque et les voies géni- tales femelles des Lygaeides Oxycareninae (Het- eroptera). Revue Francaise d’Entomologique 31: 196-218. Costa, A. 1864. Acquisti fatti durante l’Anno 1862. Annuario del Museo Zoologico dell Regia Univer- sita di Napoli 2: 8-94. 426 Distant, W. L. 1901. Rhynchotal notes. XI. Heter- optera. Fam. Lygaeidae. Annals and Magazine of Natural History Series 7:8: 464-486, 497-510. Fieber, F. X. 1870. Dodecas neuer Gattungen und neuer Arten europaischer Hemiptera. Verhand- lungen der Kaiserlich-Koniglischen Zoologisch- Botanischen Gesellschaft in Wien 20: 243-264. Harrington, J. 1980. A generic level revision and cladistic analysis of the Myodochini of the World (Hemiptera, Lygaeidae, Rhyparochrominae). Bul- letin American Museum of Natural History 167: 49-116. Kiritshenko, A. N. 1951. True Hemiptera of Euro- pean USSR. Key and Bibliography. Opred. Faune SSSR 42: 1-423. Linnavuori, R. 1978. Hemiptera of the Sudan, with remarks on some species of the adjacent countries 6. Aradidae, Meziridae, Aneuridae, Pyrrhocori- dae, Stenocephalidae, Coreidae, Alydidae, Rho- palidae, Lygaeidae. Acta Zoologica Fennica 153: 1-108. Lucas, P. H. 1849. Exploration scientifique de |’Al- gerie. Zoologie, Paris: Imprimerie Nationale 3: 1- S276 Rambur, J. P. 1839. Faune entomologique d I’An- dalousie. Tome II. Arthus Bertrand. Libraire, Par- is. 336 pp. Scudder, G. G. E. 1963. The World Rhyparochromi- nae (Hem.: Lyg.) III. New Rhyparochrominae from the Ethiopian Region. Canadian Entomologist 95: 1233-1253. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1968. The World Rhyparochrominae (Hem. Lyg.). VI. Further new genera for previously de- scribed species and some additional new species contained therein. Journal of Natural History 2: 577-592. . 1969a. The Lygaeidae (Hemiptera) collected by Dr. M. Mrazek and Mrs. I. Korecka in the republic of Guinea. Acta Musei Moraviae 54: 169- 182. . 1969b. The World Rhyparochrominae (Hem. Lyg.) IX. Four new genera collected by Dr. R. Linnavuori in the Sudan. Annales Entomologici Fennici 35: 85-94. Slater, J. A. 1993. A new genus and two new species of Drymini from Africa (Hemiptera: Lygaeidae). Revue de Zoologie Africaine 107: 373-381. Slater, J. A. and J. Carayon. 1963. Ethiopian Ly- gaeidae IV. A new predatory lygaeid from Africa with a discussion of its biology and morphology. (Hemiptera: Heteroptera). Proceedings of the Royal Entomological Society of London (A) 38: I-11. Stal, C. 1862. Hemiptera Mexicana enumeravit spe- cies-que novas descripsit. Stettin Entomologische Zeitung 23: 289-325. Sweet, M. H. 1967. The tribal classification of the Rhyparochrominae (Heteroptera: Lygaeidae). An- nals of the Entomological Society of America 60: 208-226. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 427-434 CARVALHOFULVIUS GIGANTOCHLOAE, A NEW GENUS AND SPECIES OF BAMBOO-INHABITING FULVIINI FROM WEST MALAYSIA (HETEROPTERA: MIRIDAE: CYLAPINAE) GARY M. STONEDAHL AND DAMIR KOVAC (GMS) International Institute of Entomology, 56 Queen’s Gate, London SW7 5JR, UK; (DK) Forschunsinstitut Senckenberg, Senckenberganlage 25, D-60325 Frankfurt a.M., Germany. Abstract. — Carvalhofulvius gigantochloae, a new genus and species of fulviine Miridae from West Malaysia is described and illustrated. This bug lives inside the shoot internodes of Gigantochloa scortechinii (Poaceae: Bambusoideae), where it apparently feeds on de- veloping fungal hyphae. The tribal placement of Carvalhofulvius and its relationship to other fulviine mirids are discussed. Biological information is provided, including a sum- mary of a field study of colonization densities on different internode types (e.g. shoots, older culms). Key Words: Heteroptera, Miridae, Fulviini, Carvalhofulvius, new genus, West Malaysia, bamboo-inhabiting, phytotelmata, biology An ongoing study of the arthropod spe- cies inhabiting internodes of the giant bam- boo, Gigantochloa scortechinii Gamble, in West Malaysia (coordinated by DK) has re- vealed an interesting species of fulviine Mir- idae that cannot be placed in any known genus of the tribe. Carvalhofulvius gigan- tochloae, new genus and new species, is here described and illustrated; its biology, place- ment in the Fulviini, and relationship to other fulviine genera are discussed. Institute abbreviations used to record the depositories of types and other specimens are as follows: American Museum of Nat- ural History, New York (AMNH); For- schunsinstitut Senckenberg, Frankfurt (FS); National Museum of Natural History, Washington, D.C. (USNM); Natural His- tory Museum, London (NHM). All measurements are given in millime- ters. Carvalhofulvius Stonedahl and Kovac, NEw GENUS (Figs. 1-14) Type species.—Carvalhofulvius giganto- chloae, new species, here designated. Diagnosis.— Distinguished from other genera of Fulviini by the long head, with eyes well removed from anterior margin of pronotum (Figs. 1, 2, 9); enlarged profe- mora, with two irregular rows of socketed spines ventrally (Figs. 6, 8); and by the structure of the male genitalia, especially the various sclerotized appendages of the vesica (Fig. 14). Description. — Male. Macropterous; dark brown to nearly black with pale markings at base of hemelytra and apex of corium; dorsal surface finely granular, slightly shin- ing; posterolateral regions of pronotum fine- ly wrinkled; dorsal vestiture of short, stout, 428 Fig. 1. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Carvalhofulvius gigantochloae, n. sp., dorsal habitus of male. VOLUME 97, NUMBER 2 429 Figs. 2-7. Scanning electron micrographs of Carvalhofulvius gigantochloae. 2, lateral view of head. 3, me- tathoracic scent efferent system. 4, distal third of antennal segment II. 5, detail of woolly setae on antennal segment II. 6, lateral view of profemora showing socketed spines on ventral surface. 7, pretarsus, posterior view. 430 suberect, dark setae, usually slightly longer and more densely distributed along lateral margins of pronotum, sparsely distributed on heme 20°) that are characterized by high in- solation temperatures and low moisture availability at the surface; they are often lee of mountains and within rain shadows, receive an average rainfall of 70 to 100 cm per year (Platt 1951, Brooks 1965, Keener 1983, DeSelm and Murdock 1993). Despite the hot, dry microclimate and sparse veg- etation—treeless openings are common— these shale slopes retain sufficient soil mois- ture to sustain the growth of herbaceous en- demic plants (Platt 1951, Keener 1983). Occurring mainly on Upper Devonian 436 shales of the Brallier formation, shale bar- rens may also be associated with Ordovi- cian Martinsburg shales or even Silurian strata (Ar 937, Platt 1951, Keener 1983, Morse !|983). Like other unusual soil types, these edaphic islands in the eastern decid- uous forest of the Valley and Ridge phy- togeographic province (Appalachian Val- ley) support a distinctive flora (e.g., Wherry 1930, Core 1940, 1952, Allard and Leonard 1946, Platt 1951, Henry 1954, Keener 1970, 1983, Morse 1993). The presence of several endemic plants, disjuncts, and various rare or endangered plant taxa that are absent or seldom present in the surrounding wood- lands suggests the occurrence of a diverse fauna of phytophagous insects. Such is in- deed the case (A.G.W. unpublished data), although entomologists have paid relatively little attention to insect-plant relationships in these communities. The butterflies and skippers characteristic of shale barrens, however, are well known (Clench and Opler 1983, Opler and Krizek 1984, Pague and Schweitzer 1991), and Pennsylvania shale barrens contain some of that state’s rarest butterfly and moth species (Smith 1989). Species in a few other insect groups have also been studied, e.g. the grasshopper Me- lanoplus impudicus Scudder (Gurney 1941) and the psyllid Craspedolepta eas (McAtee) (Wheeler 1994). Moss phlox, or moss or mountain pink (Phlox subulata L.), is a prostrate or matlike suffruticose (i.e. becoming somewhat woody) perennial that characterizes shale barrens. It harbors an unusually rich insect fauna, including the recently described whitefly Trialeurodes phlogis Russell (Russell 1993), and several other poorly known species (Henry 1979, Wheeler et al. 1983, Wheeler 1994, unpublished data). In the present pa- per, I provide biological information on the mirids or plant bugs that develop on P. sub- ulata growing on various types of bedrock; mirid-moss phlox relationships in mid-Ap- palachian shale barrens are emphasized. Data from an inventory of the mirids as- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sociated with other eastern subulate or nar- row-leaved phloxes are also presented. This paper commemorates the remarkably pro- ductive scientific career of José C. M. Car- valho and his devotion to the Miridae. His research and numerous publications, in- cluding a world catalog of the group, stim- ulated interest in mirid systematics and bi- ology. Host PLANTS The genus Phlox belongs to the Pole- moniaceae, a mostly New World family. It contains numerous endemic eastern species but is most diverse in western North Amer- ica (Grant 1959). Mirids were collected principally from P. subulata, although col- lections were also made from P. bifida Beck and P. nivalis Lodd. ex Sweet. These three apparently closely related species (particu- larly P. bifida and P. subulata [Gleason and Cronquist 1964]) are the only narrow-leaved members of the genus (Section or Subsec- tion Subulatae) occurring in eastern North America (Wherry 1935b, 1951, Flory 1970). Phlox nivalis is sometimes relegated to the Section Protophlox (Wherry 1955) but is actually similar in both morphology and cy- tology to P. bifida and P. subulata (Smith and Levin 1967). Phlox subulata. —Moss phlox (Figs. 1, 2) is a suffruticose perennial that has many- branched trailing or creeping stems, is woody at the base, and has persistent awl-shaped or needlelike leaves. The main blooming period is April to May, or in cooler locali- ties, into June; colonies vary in flower color from rose-purple to pink or white. Forming dense, low, evergreen mats on sandy or gravelly soil and rocky ledges (Fig. 1), moss phlox occurs not only in shale barrens but also in serpentine barrens, on limestone cliffs, and on other types of bedrock (Stout 1917, Wherry 1929a, 1930, 1936, 1955, 1964, Core 1966, Everett 1981, Keener 1983, Bartgis 1985). Phlox subulata typi- cally grows in well-drained, slightly alkaline to strongly acidic soils. This pioneer, xero- VOLUME 97, NUMBER 2 437 Figs. 1-2. Phlox subulata. 1, Colony on shale outcrop in West Virginia. 2, Close-up of inflorescence. (Photographs courtesy of T. J. Allen.) 438 seldom found in nearby ws, where it is outcompeted phytic plant | woods or mea by faster-growing plants (Wherry 1929b, 1935b, Allard and Leonard 1946, Morse 1988) Moss phlox is native from southern On- tario and New York to southern Michigan and south to the mountains of North Car- olina and Tennessee (Wherry 1929b, 1935b, 1955, Flory 1970, Everett 1981, Gleason and Cronquist 1991). Its dispersal center may be the Appalachians of eastern West Virginia (Wherry 1935b). Phlox subulata is used extensively in rock gardens and bor- ders, and as a ground cover on slopes; it often escapes from cultivation, establishing colonies in cemeteries and on artificial bar- rens such as road and railroad cuts (Wherry 1929b, c, Everett 1981, Cooperrider 1986, Hudak 1993). Phlox bifida.—Cleft or sand phlox is a tuft-forming species superficially similar to P. subulata. It differs, however, in its looser, often more erect, growth habit; longer leaves, which are fewer in number: and deeply cleft (bifid) petal blades. Flowers vary from pale blue or violet to white, but are mostly lav- ender. This spring-blooming perennial shows a disjunct distribution from southern Michigan to Tennessee, northern Arkansas, and eastern Kansas. A plant mainly of the Interior Highland and Low Plateaus with its range centering in southern Illinois, this pioneer species colonizes sandy banks and dunes, rocky slopes, and bare cliffs (Wherry 1929b, 1935b, 1955, Winterringer and Ves- tal 1956, Everett 1981, Gleason and Cron- quist 1991), Phlox nivalis.—Piney-woods or trailing phlox has foliage similar to that of P. sub- ulata, but some of its leaves are usually broader. Although these species have been confused botanically, they differ markedly in floral structure: P. nivalis has much short- er stamens and style and somewhat larger flowers. The specific epithet, nivalis, refers to the often snowy white flowers, but the corolla may be pink or light purple. More southern in distribution than the other east- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ern narrow-leaved phloxes, P. nivalis ranges from southern Virginia to Florida and Al- abama. Its dispersal center lies in the Pied- mont of South Carolina. Piney-woods phlox grows mainly in acid soils on dry sandy slopes of open pine and oak woods. It blooms from March to April in the southern part of its range, April and May toward the northern end (Wherry 1929b, 1935b,d, 1955, Everett 1981, Gleason and Cronquist 1991). Stupy SITES AND METHODS Muirids were observed and collected from 1989 to 1994, chiefly on P. subulata in mid- Appalachian shale barrens of Upper De- vonian strata and Ordovician Martinsburg outcrops (Fig. 3). Included were such well- known botanical sites as Green Ridge in Maryland; Eagle Rock, Head Waters, Mill- boro, and Short Mountain in Virginia; and Kates Mountain in West Virginia (Wherry 1935a, Butts 1940, Core 1940, 1952, Allard and Leonard 1946, Platt 1951, Keener 1983, Morse 1983). Numerous shale outcrops that cannot be characterized as true barrens were also sampled in the Valley and Ridge Prov- ince, as were a few ornamental plantings. In addition, collection sites included eastern serpentine barrens such as Nottingham, Pink Hill, and Unionville in Pennsylvania, and Soldiers Delight in Maryland (Wheeler 1988 and references therein), and sites in the Ap- palachian Highlands where P. subulata grows on dolomite, greenstone, limestone, and other substrates. Limited sampling was conducted to determine the mirid fauna of P. subulata outside this plant’s center of dis- tribution: in Kentucky, New Jersey, New York, North Carolina, and Ohio. Infor- mation on seasonality and habits of mirids associated with P.subulata was obtained largely from the following sites; the number of visits to each site is given in parentheses (see also Table 1). MARYLAND: Allegany Co., Country Club shale barren above Evitts Cr., NE. of Cumberland (3) and shale barrens, Green Ridge State Forest, Fifteen Mile Creek Rd. VOLUME 97, NUMBER 2 439 Table 1. Miridae observed on Phlox subulata during 1989-1994; asterisks denote ornamental plantings. idea heidemanni [P.wheeleri_ L. minor oc ye Oe et. eee | Country Club shale barren, NE. of Cumberland | Fort Hill, south tip ridgetop Green Ridge State Forest, Fifteen Mile Creek Rd. at Piclic Rd. Romney-Oldtown shale barren, E. of Oldtown Soldiers Delight serpentine barren, SW. of Reisterstown Great Falls Boy Scout shale barren, Sideling Hill Wildlife Management Area North Carolina Union *N. of Fairview, Rt. 601 eee Guernse nnsylvania OBODOBOOOEO ., Twp. Rd. ; N. of Winterset, Twp. Rd. 871 O O SW. of Tarentum, Bull Creek Rd. '@| E. of Fishertown, Rt. 56 @| NE. of Ryot, Rt. 96 '@| Silver Mills shale barren, E. of Inglesmith |@| Fern Hill serpentine barren, West Goshen Twp. oO Nottingham Park serpentine barren |@| Sugartown serpentine barren, Willistown Twp. O Unionville serpentine barren, NE. of Unionville |@| Pink Hill serpentine barren, Tyler Arboretum, nr. Lima NE. of Shawnee, Mosier Knob Rd. *2 mi. E. of Milton, Rt. 642 O |@| ea 7.6 mi. S. of Covington, Rt. 18 |@| O O > IS l= g |e | 5 5 Pe Alleghen Bedford Chester Delaware Monroe Northumberland Alleghany 10.6 mi. S. of Covington, Rt. 18 N. of Sweet Chalybeate, Rt. 311 Fort Lewis shale barren, Rt. 678 at Cowpasture River 3 mi. SW. of Millboro Springs, Rt. 42 *Millboro 3.5 mi. SW. of Bland, Rt. 42 1.2 mi. E. of Crandon, Rt. 42 Eagle Rock shale barren, 2.5 mi. NW. of Eagle Rock SE. of Dugspur, Rt. 638 *SE. of Willis, Rt. 799 Head Waters shale barren E. of Frost, W. Va., Rt. 600 S. of Rt. 64 Madison Shenandoah Nat. Park, Skyline Drive, Franklin Cliffs @ @| el O O O O O Montgomery Coffee Valley, S. of Ironto @| O fe O a O a O O Bland Botetourt Carroll Floyd Highland N. of Ironto, Rt. 713 Page Shenandoah Nat. Park, Skyline Drive, Little Stony Man Cliffs Dixie Cliff, SW. of Glenvar Roanoke . of Catawba, Rt. 311 Rockbridge Goshen, Rts. 39-42 Rockingham George Washington Nat. Forest, For. Rd. 87, W. of Fulks Run Shenandoah 2.5 mi. N.o inburg, Rt Short Mountain shale barren, 3 mi. SE. of Mount Jackson }@/ je] | jeje! [ele jeeee | OCECCOsCECESSaaaEOs OBS fs Hig. " ) | jee +s at Piclic Rd. (5); Baltimore Co., serpentine Sideling Hill Cr., E. of Little Orleans (4). barrens, Soldiers Delight Natural Environ- OHIO: Athens Co., Buffalo Beats prairie, mental Area, W. of Owings Mills (3); Wash- SE. of Buchtel (2); Guernsey Co., shale banks ington Co., Boy Scout shale barren above along county road, NW. of Winterset (3). 440 a Ne fable 1. Continued. Coun Greenbrier ESIRONSROKEL) DAF BY 3 i=" ~A Hampshire S. of Romne Mineral Morgan Pendleton PENNSYLVANIA: Bedford Co., road- side slope, NE. of Ryot (4); Chester Co., Nottingham(3),Sugartown(4),andUnionville (2) serpentine barrens; Delaware Co., Pink Hill serpentine barren, Tyler Arboretum, NE. of Lima (4); Monroe Co., roadside bank along Mosier Knob Rd., NE. of Shawnee (2). VIRGINA: Alleghany Co., shale barren above Potts Creek, Rt. 18, 10.6 mi. SW. of Covington (8); Bath Co., shale barren above Cowpasture River, Rt. 678, Ft. Lewis (5); Highland Co., Head Waters shale barren ite phur Springs E. of Alvon, Whites Draft Rd. Cave Mountain, 1 mi. N. of Landes N. of Arthur, Patterson Creek Rd. 4 mi. N. of Landes, Rt. 220 2.3 mi. S. of Landes, Rt. 220 3 mi. S. of Landes, Rt. 220 3.5 mi. S. of Petersburg, Rt. 220 , River Rd. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON above Shaws Fork, nr. junc. Rts. 250 and 616 (19); Montgomery Co., shaly slope, Rt. 622 nr. junc. Rt. 603 N. of Ironto (5); Ro- anoke Co., Dixie Cliff, SW. of Glenvar (2); Rockbridge Co., shale slopes along Rts. 39- 42, Goshen (9); Rockingham Co., shaly banks along For. Rd. 87, George Washing- ton National Forest, W. of Fulks Run (4); Shenandoah Co., shale barren, south end of South Mountain, SE. of Mount Jackson (8). WEST VIRGINIA: Greenbrier Co., shale barren, Kates Mountain, S. of White Sul- phur Springs (9) and shale banks along VOLUME 97, NUMBER 2 441 New York —— Pennsylvania a Ue A i \West Virginia / é A Fige3: | a q “New Nersey Fe =Maryland’ a? \D ay ; fs Posie Virginia Occurrence of mid-Appalachian shale barrens based on Keener (1983); larger stippled area = Upper Devonian shale strata, smaller area = Ordovician Martinsburg outcrops in the Massanutten Mountains. Whites Draft Rd., E. of Alvon (3); Hamp- shire Co., roadside slopes, Rt. 29, 3.5 mi. N. of Slanesville (6) and Rt. 50, Shanks (6); Hardy Co., shale banks along Lost River Rd., W. of Mathias (2); Mineral Co., shale banks, Rt. 50 nr. Burlington (2), Pendleton Co., shale banks along Schmucker Rd., SE. of Upper Tract (3). Mirids were sampled on P. bifida in Ili- nois along bluff top woods, LaRue-Pine Hills Ecological Area, Shawnee National Forest, Union Co.; on limestone bluffs near Prairie du Rocher, Randolph Co.; and in Mason Co. in loess hill prairie, Revis Nature Pre- serve SW. of Mason City, and in sand prai- rie, Sand Ridge State Forest NE. of Havana. Collections were made in Indiana on lime- stone bluffs above the Ohio River at Porters Point, SW. of Laconia, Harrison Co.; in Kentucky on limestone ledges of ““Boones Knoll,’ Camp Nelson, Jessamine Co.; and in Tennessee in cedar glades near Cedars of Lebanon State Park and at Long Hunter State Recreation Area, Wilson Co. Limited sampling of mirids associated with P. nivalis was conducted. In North Car- olina a collection was made from a roadside bank along Rt. 601 W. of Stanfield, Cabar- rus Co.; no mirids were found on this plant at Bluff Mountain, N.C., or at several sites in South Carolina (Lexington, Pickens, and York counties). The main collecting sites were visited at irregular intervals from early or mid-April through June; in some years, sampling of P. subulata began in late March and extended into early October. At each site, mirids were sampled by shaking mats of phlox over a shallow white tray and recording any mirid species present and their relative abun- dance. Sampling times varied from 10-15 minutes for small host patches to 45-60 minutes or more for sites having extensive phlox colonies. In the first season of study 442 (1989), different-appearing nymphs were reared on sprigs of the host plant to asso- ciate them with adults of the various mirids. Nymphal stages of each species present were recorded in the field or, when necessary, collected and sorted to instar in the labo- ratory. [niormation on feeding habits was obtained in the field and supplemented by laboratory observations. Voucher speci- mens of the four phlox-associated mirids have been deposited in the collections of Cornell University, Ithaca, N.Y.; the Na- tional Museum of Natural History, Wash- ington, D.C.; and the Pennsylvania De- partment of Agriculture, Harrisburg. RESULTS Information is presented on four mirid species that occur consistently on P. subu- lata: the generalists Lopidea heidemanni Knight and L. minor Knight (Orthotylinae: Orthotylini) and specialists Polymerus tinc- tipes Knight and P. wheeleri Henry (Miri- nae: Mirini). For each species, the known distribution is summarized and any new state records obtained are noted; literature on host plants, including those used for re- production as well as adult feeding, is re- viewed; and the seasonal history and habits on P. subulata are summarized. Associa- tions with other plants found in shale bar- rens and shale outcrops, and on other nar- row-leaved eastern phloxes, are included. Nymphs of the phyline Plagiognathus poli- tus Uhler occasionally developed on P. sub- ulata in shale barrens, but this polyphagous species will not be mentioned further. Lopidea heidemanni Knight This polyphagous plant bug, common on weedy vegetation along roads and in other disturbed habitats, is more likely to be en- countered by the general collector than the other phlox-associated miurids that were studied. Despite its wide range and fre- quency of collection, the habits of L. hei- demanni are not well known. Distribution. —Described by Knight PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (1917) from New York, L. heidemanni has since been recorded from 30 additional states and the District of Columbia, ranging from Vermont, south to Georgia, and west to Minnesota, Wyoming, and Texas (Henry and Wheeler 1988, Asquith 1991). AI- though L. heidemanni was present on P. subulata at 62 of 79 sites that yielded plant bugs (Table 1), no new state records were obtained during this study. Asquith (1991) mapped the known distribution of this spe- Cies: Host plants. —Nymphs have been record- ed from yarrow (Achillea millefolium L.), ninebark (Physocarpus opulifolius (L.) Max- im.), and terminal growth of young elm (U/- mus sp.) (Knight 1923, 1941, Wheeler and Hoebeke 1985). This species has also been collected on numerous other herbs, shrubs, and trees (e.g. Messina 1978, Snodgrass et al. 1984, Blinn and Yonke 1985, Asquith 1991). Because adults readily disperse to various plants (see “Seasonal history and habits’), some of the recorded “hosts”? may be used only for adult feeding or shelter. Although this species is said to develop mainly on deciduous trees (Asquith 1993), herbs would appear to be used more fre- quently as host plants than shrubs or trees. In the present study, nymphs developed on Phlox bifida and P. nivalis, and on P. subulata in native colonies and landscape plantings. In shale barrens and outcrops, nymphs were also observed on Ceanothus americanus L., Centaurea biebersteinii DC. (= C. maculosa of authors), Clematis albi- coma Wherry, Crataegus sp., Draba ra- mosissima Desv., Eriogonum allenii S. Wats., Oxalis stricta L., Penstemon canes- cens (Britt.) Britt., Rhus aromatica Ait., Se- necio antennariifolius (Britt.) Britt., Silene caroliniana ssp. pensylvanica (Michx.) Clausen, Thlaspi perfoliatum L., and Vicia villosa Roth. Adults were found on Hyper- icum prolificum L., Quercus ilicifolia Wan- genh., Rhus copallinum L., and Senecio anonymus Wood in shale barrens. A com- mon breeding host in serpentine barrens was VOLUME 97, NUMBER 2 Egg i . Nymph Adult Lopidea heidemanni , Egg Lopidea ‘ Nymph minor Adult Polymerus Egg tincti, Nymph “is Adult Polymerus — h wheeleri aie Fig. 4. Cerastium arvense L.; adults were observed on Quercus stellata Wangenh. Seasonal history and habits. —Overwin- tered eggs usually began to hatch from mid- or late March to early April in shale barrens of Virginia (Fig. 4) and mid- to late April in Pennsylvania serpentine barrens. In early season, the bright red nymphs were often shaken only from plants that were in bloom. Early instars fed on parts of the corolla, both in the field and laboratory, but probably also feed on the foliage. Although the needlelike leaves of P. subulata did not show feeding injury, chlorotic blotches were observed on the foliage of V. villosa fed on by L. hei- demanni nymphs. In Virginia, fourth and fifth instars were present on P. subulata as early as mid- or late April (rarely in early April), with adults appearing during the first week of May (Fig. 4). Late-instar nymphs, however, were typically present in shale barrens until mid- or late May, often feeding on the fruits. Development of populations at the Kates Mountain shale barren (ele- vation about 760 m) lagged behind that in other barrens of Virginia and West Virginia (elev. generally 300-700 m). During 5-6 May 1990, for example, late instars and ten- eral adults were observed in several shale Mari spaces [¥AN [ip [MAR|APR [MAY|JUN [sux [AUG] Sep [OCT Nov] DEC Ns 443 gman Generalized mirid seasonal histories based on sampling of Phlox subulata in eastern West Virginia— southwestern Virginia, 1989-1994; see text for seasonality of populations occurring at higher altitudes and latitudes. barrens of southwestern Virginia, but only third instars were seen on Kates Mountain. First-generation adults of L. heidemanni often dispersed to nearby shrubs and trees soon after developing on P. subulata; adults were never aS common on moss phlox as the nymphs. A short-distance dispersal from breeding hosts is known in several other Lopidea species (Asquith 1991). At several sites, a small second generation developed on P. subulata. The first through third in- stars observed in late June 1990 and fourth and fifth instars in late June 1994 apparently represented another generation rather than a late hatching of overwintered eggs. The collection of adults over a three-month pe- riod (19 May—18 August) in North Carolina (McPherson et al. 1983), and the appear- ance of adults in that state during April (A.G.W. unpublished data) is further evi- dence for bivoltinism in L. heidemanni. The latest records of adults on P. subulata were 10 July at the Head Waters shale barren in Virginia and 11 July at the West Virginia site near Slanesville. Adults, however, have been collected in West Virginia on other plants as late as 19 August (Wheeler et al. 1983). On P. nivalis in North Carolina (Cabarrus 4qad Co.), fourth instars were observed in mid- April 1990. During 19-23 April 1991, sec- ond and third instars were found on P. bifida in sand prairies of Illinois and on Boones Bluffat Camp Nelson, Kentucky, and fourth and fifth instars were present on P. bifida in Tennessee cedar glades. Lopidea minor Knight This widespread generalist feeder is found east of the Rocky Mountains to New York and south to Florida. A characteristic plant bug of the semiarid plains (Knight 1965), it occurs mainly in relict prairies, serpentine barrens, and shale barrens in the mid-At- lantic states. This plant bug was usually found on moss phlox growing in the most open, sunlit portions of shale barrens. Some of the records of L. minor have been pub- lished under names recently proposed as synonyms: L. petalostemi Knight, L. john- stoni Knight, and L. phlogis Knight (As- quith 1991). Distribution.—This orthotyline was de- scribed from Colorado and North Dakota (Knight 1918) and has since been recorded from Alberta, British Columbia, Florida, II- linois, lowa, Kansas, Louisiana, Manitoba, Minnesota, Mississippi, Missouri, New York, North Carolina, Oklahoma, Penn- sylvania, Saskatchewan, South Dakota, Texas, Virginia, West Virginia, and Wyo- ming (Henry and Wheeler 1988, Asquith 1991). Maryland is a new state record. Lopi- dea minor was present in 32 of 79 sites (Ta- ble 1). Host plants. —In western North America, prairie clover (Dalea purpurea Vent var. purpurea (= Pelalostemon purpureus) serves as acommon host (Knight 1927, 1941, 1965, Blinn and Yonke 1985); L. minor has also been recorded from the legume Hedysarum sp. in the Canadian Prairie Provinces (Kel- ton 1980). Phlox subulata is used as a host in the eastern United States (Knight 1965, Wheeler et al. 1983); records from this plant in a Pennsylvania serpentine barren and shale barrens in Virginia and West Virginia PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Asquith 1991) were based on material col- lected during the present study. In this study, L. minor was found only on P. subulata among the eastern narrow-leaved phloxes, occurring occasionally in orna- mental plantings. Nymphs were also col- lected on Eriogonum allenii, Paronychia montana (Small) Pax & K. Hoffmann, and Senecio anonymus in shale barrens, and they were found once on and under a fern (Chei- lanthes lanosa (Michx.) D.C. Eat.) in the Virginia shale barren near Eagle Rock in Botetourt Co. Adult males sometimes dis- persed to flowers of Chrysanthemum leu- canthemum L., Sedum spp., and grasses, and to fruits of Staphylea trifolia L. Seasonal history and habits. —Hatching of overwintered eggs began in early or mid- April in shale barrens of southwestern Vir- ginia and in late April in Pennsylvania ser- pentine barrens (Fig. 4). Egg hatch typically was spread over several weeks so that four or even all five nymphal stages overlapped in populations sampled during May. Nymphs are gray rather than bright red as in L. heidemanni. Adults first appeared in mid-May in more southern shale barrens, although late instars were often observed until early June; females persisted until mid- July. Each season, a clear-cut second gen- eration was produced (Fig. 4), which con- firms the bivoltine life history that had been suggested for L. minor (Asquith 1991). Egg hatch began in late June in southern shale barrens, and second-generation adults be- gan to appear in late July. Late instars, how- ever, were usually common through August and were sometimes present into Septem- ber. The latest records of adults were 1-2 October in three Virginia shale barrens. Nymphs of L. minor, while occasionally seen on flower buds of Erigonum allenii in shale barrens, were not observed on flower buds or inflorescences of P. subulata. They live within or under mats of moss phlox and apparently feed on leaves or stems. Second- generation nymphs develop in midsummer when moss phlox is essentially dormant, VOLUME 97, NUMBER 2 \ Ohio Illinois Indiana | es e e (West Virginia ie. o Pe ee wc Virginia . \ ae) is Q Map 1. e i 445 \ } ee if } . *\ | New Hampshire reer New York a ~ _ Massachusetts. * an 1 \ Connectic \ a , \ ~~ ray Rhode Island £ il at Jersey Pennsylvania me } ; -WaryianAl\' =F aal\ rate x af IDeeeware ye * : e (i \ t L _) ple "4 \ / 4 S LA M is Known distribution of Polymerus tinctipes. Star = type locality; filled circles = new records (see also Table 1); shading = counties in which specimens were collected. having finished blooming and produced seeds, and are restricted to feeding on fo- liage that often appears brownish rather than lush green as in April and May. Polymerus tinctipes Knight This phlox specialist has not been men- tioned in the literature since its original de- scription (Knight 1923), except in catalogs (Carvalho 1959, Henry and Wheeler 1988). It is, however, a characteristic species of moss phlox, the dearth of biological infor- mation merely reflecting entomologists’ lack of attention to the insect fauna associated with this prostrate plant. Distribution. —This mirine is known his- torically only from the type locality, Great Falls, Md., where four specimens were col- lected on 3 May 1915 (Knight 1923). In the present study, P. tinctipes was found on P. subulata at 32 of 79 sites sampled. New state records are Ohio, Pennsylvania, Virginia, and West Virginia (Map 1, Table 1). Host plants.—No host has been previ- ously recorded for P. tinctipes, the type spec- imens having been taken at Great Falls, Md., on a “lichen covered rock” (Knight 1923). This plant bug was found in the present study only on P. subulata, mainly in mid-Appa- lachian shale barrens and other shale out- crops, and in Pennsylvania serpentine bar- rens; it was also collected on moss phlox in a relict prairie in Ohio, a greenstone glade in the Blue Ridge of Virginia, and rarely in ornamental plantings. During visits to the type locality, where P. subulata grows on rocks above the Potomac River (Hitchcock and Standley 1919), this bug was found to be common. Adults mate on P. subulata and tend to remain on their host rather than disperse to nearby vegetation, unlike the two Lopidea species. Seasonal history and habits. —Eggs of this univoltine mirid may hatch in late March, as evidenced by the presence of third instars during 8-12 April 1990. In most years, 446 Wisconsin Illinois Indiana ) West Virginia Missouri Map 2. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON { { 1 l | Vermont } by ) \ New Hampshire ) New York | Massachusetts, 1 i UP Connecticut \” 4 “7 Rhode Island ( ib Pennsylvania | e { ( New Jersey i A — 7G Maryland?) \ \/ oS) Tare it Delaware ; Je N } pve Cae pr? < t } (es t Tal Virginia North Carolina Known distribution of Polymerus wheeleri. Star = type locality; filled circles = new records (see also Table 1); shading = counties in which specimens were collected. hatching probably does not begin in south- western Virginia shale barrens until early or mid-April (Fig. 4). Eggs did not hatch until late April or early May in higher-elevation shale barrens (e.g. Kates Mountain) or sites at even higher latitudes, in serpentine bar- rens in Pennsylvania, and at sites in Ohio. Fourth and fifth instars were usually com- mon in more southern shale barrens by ear- ly May. The green nymphs were observed to feed on flowers and fruits of P. subulata. The earliest record of adults was 7 May, but typically they did not appear until mid- to late May and were present only until late June or early July. The latest records for adults were 17-18 July at Millboro, Va., and Kates Mountain, W. Va. Polymerus wheeleri Henry The collection of this plant bug, at the time undescribed, from a West Virginia shale barren in 1978 was responsible for stimu- lating my interest in mirids and other in- sects associated with P. subulata. No ad- ditional collections of this phlox specialist have been reported since the original de- scription (Henry 1979, Henry and Wheeler 1988). Distribution. —This mirine has been known only from the type locality, a shale barren west of Petersburg, W. Va. (Henry 1979), which has nearly been destroyed by construction. New state records obtained during the present study are Illinois, Mary- land, Pennsylvania, and Virginia (Table 1). Polymerus wheeleri was found at fewer sites (12 on P. subulata) than any of the other plant bugs associated with eastern narrow- leaved phloxes (Map 2, Table 1). Host plants. — Described from specimens collected on P. subulata, this species is char- acteristic of the fauna of mid-Appalachian shale barrens and outcrops, as well as mats of P. bifida ssp. bifida that festoon limestone ledges and bluffs in the Ozark Division (Mohlenbrock 1986) of southern Illinois. It was not found on P. subulata in eastern ser- pentine barrens. Polymerus wheeleri adults VOLUME 97, NUMBER 2 show little tendency to disperse to nearby plants, although at the Short Mountain shale barren in Virginia an adult was observed on flowers of the crucifer Draba ramosissima. Seasonal history and habits. —Eggs of this univoltine, early-season species began to hatch at about the same time as those of L. heidemanni: generally late March to early April (Fig. 4). In 1991 the presence of a third-instar nymph (with firsts and seconds) at the Head Waters shale barren on 27 March suggests that hatching had begun by mid- March. Nymphs are dark red rather than green as in P. tinctipes; they fed on petals, sepals, fruits, and pedicels of P. subulata, but they may also feed on the foliage. Fourth instars were observed as early as 8 April but were not usually present until mid- to late April. Adults began to appear from late April to early or mid-May at most sites in Virginia and West Virginia, although in 1991 a fifth instar collected on 14 April at the Short Mountain shale barren molted just two days later when held at room temperature. Adults remained on mats of moss phlox and gen- erally disappeared by late June, males usu- ally dying 7-10 days before females. The latest records were 10-11 July at Head Wa- ters, Va., and near Slanesville, W. Va. DISCUSSION Phlox subulata supports an unexpectedly rich mirid fauna: two generalist and two specialist species. Many shrubs and trees of eastern North America do not serve as hosts for as many as four mirids (personal obser- vation). Absent from the fauna of P. sub- ulata and other eastern narrow-leaved phloxes of prostrate growth habit are mirids that develop on native and cultivated phloxes of erect growth; Lopidea davisi Knight (Cory and McConnell 1927) and L. confluenta (Say) (Asquith 1991, personal observation) were not encountered during this study. Lopidea heidemanni, L. minor, Polyme- rus tinctipes, and P. wheeleri develop on the glandular P. subulata ssp. brittonii in the 447 region sometimes referred to as the Southern Appalachian Highlands (Blauch 1975); they appear to be most common in the southern part of this region. These bugs were often present on P. subulata that colonized the most xeric sites. The importance of slope was evident when colonies subject to dif- ferent exposures were sampled along a winding mountain road; mirids typically were found on moss phlox in the drier sites rather than in more mesic sites less than a kilometer away. Except for occasional in- dividuals observed feeding on reproductive structures, the bugs generally were not seen on exposed portions of P. subulata. They perhaps remain mostly within or under mats of the host plant (at least by day) to reduce the risk of desiccation in shale barrens and other slopes of high insolation tempera- tures. Even when bugs were not apparent on host plants, several individuals (some- times 5-10) could be found when a plant was shaken over a pan or tray. Plant bugs that develop on P. subulata are characteristic members of a shale barren fauna, occurring in the Valley and Ridge Province on Upper Devonian shales of the Brallier formation (elev. 300-700 m) and on Ordovician Martinsburg outcrops in Virginia’s Massanutten Mountains (elev. about 700 m at sample site). Only at the southern end of Short Mountain in the Mas- sanutten complex, however, were all four mirid species observed at one site. Poly- merus tinctipes was found at altitudes rang- ing from near sea level along the Potomac River at Great Falls, Md., to about 1040 m in Madison Co., Va., in the Blue Ridge. This species and the other mirids of P. subulata were generally scarce in the Blue Ridge (Po- lymerus wheeleri was absent). Lopidea heidemanni, which shows the greatest diet breadth among the four spe- cies, occurred on P. subulata at more sites (62) than any of the other plant bugs and was the sole mirid recorded from 19 sites. It was also found in several ornamental plantings of P. subulata and developed on 448 P. bifida and P. nivalis, the other eastern narrow-leaved phioxes. It was the only one of the four | species that developed on P. divaric mong the phloxes of upright growth habit that were sampled in the Ap- palachians during this study. The generalist L. minor and phlox spe- cialist P. tinctipes were both found at 32 sites and showed spotty distributions in shale and in serpentine barrens; they coexisted at 12 sites. Both were found occasionally in landscape colonies of P. subulata. Their presence perhaps was the result of being moved with native plants collected for or- namental use, or possibly should be attrib- uted to dispersal to ornamental plants from nearby native colonies of P. subulata. The surprising record of P. tinctipes from an or- namental planting in Northumberland Co., Pa. (E. of Milton), might be explained by this site’s proximity to the historical occur- rence of the host 1.5 mi. ESE of Milton (1961 record in herbarium of Academy of Natural Sciences, Philadelphia). The specialist P. wheeleri occurred at 12 sites in shale barrens and outcrops, coex- isting with P. tinctipes at only 2 sites; it was absent from serpentine barrens. It also de- veloped on P. bifida in the Ozark region of southern Illinois. Another phlox specialist, the psyllid Craspedolepta eas, shows a sim- ilar disjunct distribution (Wheeler 1994). Appalachian-Ozarkian distributions are also known in other insect groups (e.g. Ross 1944, Stannard 1968, Pratt et al. 1994), and the two regions have long been considered “‘fau- nally related” (e.g. Holt 1969). Mirids were nearly always present and, often, were abundant in colonies of P. sub- ulata in eastern West Virginia and western Virginia, an area that likely is the host plant’s dispersal center (Wherry 1935b). Plant bugs typically were scarce or absent from P. sub- ulata colonies in northern New Jersey and Pennsylvania and in New York, but more sampling is needed to determine if the Po- lymerus spp. might be present in these for- merly glaciated areas. Additional sampling PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON is also necessary to determine if mirids are associated with P. subulata on Bluff Moun- tain (Tucker 1972), which is near the fringe of this plant’s range in northwestern North Carolina. The Southern Appalachian Highlands is a center of distribution for P. tinctipes and P. wheeleri, species that can be considered monophagous (i.e. restricted to developing on Phlox spp.). How these specialists on mat-forming phloxes came to occupy this region and their history during the Pleis- tocene remain unknown. Polymerus tinc- tipes may prove to be a southern Appala- chian endemic. The two Polymerus species represent the most significant discoveries among the mir- ids that occur on eastern narrow-leaved phloxes. Polymerus tinctipes had been known previously only from the type lo- cality and its host had remained unrecord- ed; P. wheeleri was an undescribed species whose collection prompted a closer look at mirid-phlox associations in shale barrens. The distribution of both specialist herbi- vores is now better known, and information on their host plant range and seasonality is available. Much, however, remains to be learned about their distribution and habits; a phylogenetic study of the genus would elu- cidate the relationship of these apparently closely related species and, perhaps, suggest modes of speciation. Lopidea minor and Polymerus tinctipes can be considered characteristic insects of serpentine barrens. In addition, these spe- cies, plus P. wheeleri, are as characteristic of shale barrens as the much better known endemic plants occurring in these com- munities. The plant bugs are subject to the same types of anthropogenic disturbances (road construction, quarrying, etc.) that threaten the shale (and serpentine) barren flora. Protection of Appalachian shale bar- rens will not only preserve the endemic plants of these communities but also the phytophagous insects that depend on them. This specialized biota can be used in studies VOLUME 97, NUMBER 2 of island biogeography (see Tepedino and Stanton 1976) and eventually may be useful in helping to interpret the complex history of the Appalachians. ACKNOWLEDGMENTS Numerous individuals contributed their expertise to this study. T. J. Henry kindly confirmed my mirid identifications and W. L. Mountain identified plants. For the pho- tographs used in Figs. | and 2, I thank T. J. Allen; J. F. Stimmel and R. K. Tressler helped with the preparation of the figures and maps. I am also grateful to all those who provided directions to sites with Phlox subulata, P. bifida, or P. nivalis; accompa- nied me in the field; facilitated the process of obtaining collecting permits; or granted permission to collect mirids: T. J. Allen, B. D. Anderson, B. K. Andreas, R. L. Bartgis, M. A. Berdine, C. W. Bier, R. Boyle, J. J. N. Campbell, F. Catchpole, A. W. Cusick, BoA. Cypher R.. Dino. GP. Fleming. F. P. Griffin, T. Hallowell, P. J. Harmon, R. L. Hoffman, R. Jensen, C. S. Keener, R. Latham, W. E. McClain, J. E. McPherson, R. Mancke, C. H. Mayes, R. Milewski, R. H. Mohlenbrock, M. S. Mumford, J. B. Nel- son, M. Ortt, T. J. Rawinski, A. F. Rhoads, J. Schwegman, T. L. Smith, P. Somers, and T. E. Wieboldt. I also thank T. J. Henry, E. R. Hoebeke, T. J. Rawinski, and G. L. Mil- ler for reviewing the manuscript and pro- viding useful suggestions. LITERATURE CITED Allard, H. A. and E. C. Leonard. 1946. Shale barren associations on Massanutten Mountain, Virginia. 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Epilogue, pp. 293-295. In Holt, P. C., ed., The Distributional History of the Biota of the Southern Appalachians. Pt. II: Flora. Research Division Monograph 2. Virginia Polytechnic In- stitute and State University, Blacksburg. Hudak, J. 1993. Gardening with Perennials Month by Month, 2nd ed. rev. Timber Press, Portland, Ore. 327 pp. Keener, C. S. 1970. The natural history of the mid- Appalachian share barren flora, pp. 215-248. In Holt, P. C., ed., Distributional History of the Biota of the Southern Appalachians. Pt. II: Flora. Re- search Division Monograph 2. Virginia Polytech- nic Institute and State University, Blacksburg. . 1983. Distribution and biohistory of the en- demic flora of the mid-Appalachian shale barrens. Botanical Review 49: 65-115. Kelton, L. A. 1980. The Insects and Arachnids of Can- ada. Part 8. The Plant Bugs of the Prairie Prov- inces of Canada (Heteroptera: Miridae). Agricul- ture Canada Research Publication 1703, Ottawa. 408 pp. Knight, H. H. 1917. New species of Lopidea (Miridae, Hemip.). Entomological News 28: 455-461. . 1918. Old and new species of Lopidea from the United States (Hemip., Miridae). Entomolog- ical News 29: 210-216. . 1923. Family Miridae (Capsidae), pp. 422- 658. In Britton, W. E., ed., The Hemiptera or Sucking Insects of Connecticut. Connecticut Geo- logical and Natural History Survey Bulletin 34. . 1927. Notes on the distribution and host plants of some North American Miridae (Hemiptera). Canadian Entomologist 59: 34-44. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1941. The plant bugs, or Miridae, of Illinois. Illinois Natural History Survey Buletin 22: 1-234. . 1965. Old and new species of Lopidea Uhler and Lopidella Knight (Hemiptera, Miridae). lowa State Journal of Science 40: 1-26. McPherson, J. E., B. C. Weber, and T. J. Henry. 1983. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 7. Miridae. Great Lakes Entomologist 16: 35-42. Messina, F. J. 1978. Mirid fauna associated with old- field goldenrods (Solidago: Compositae) in Ithaca, N.Y. Journal of the New York Entomological So- ciety 86: 137-143. Mohlenbrock, R. H. 1986. Guide to the Vascular Flora of Illinois. Southern Illinois University Press, Car- bondale. 507 pp. Morse, L. E. 1983. A shale barren on Silurian strata in Maryland. Castanea 48: 206-208. . 1988. Rare plants of Appalachian bedrock. Nature Conservancy Magazine 38(2): 28-30. . 1993. Mid-Appalachian shale barrens: an overview. American Journal of Botany 80(6 suppl.): oy Opler, P. A. and G. O. Krizek. 1984. Butterflies East of the Great Plains. Johns Hopkins University Press, Baltimore. 294 pp. Pague, C. A. and D. F. Schweitzer. 1991. Butterflies and moths, pp. 237-246. Jn Hoffman, R. L., Ar- thropods, pp. 173-249. Jn Terwilliger, K., coor- dinator, Virginia’s Endangered Species. Mc- Donald and Woodward Publishing Co., Blacks- burg, Va. Platt, R. B. 1951. An ecological study of the mid-Ap- palachian shale barrens and of the plants endemic to them. Ecological Monographs 21: 269-300. Pratt, G. F., D. M. Wright, and H. Pavulaan. 1994. The various taxa and hosts of the North American Celastrina (Lepidoptera: Lycaenidae). Proceed- ings of the Entomological Society of Washington 96: 566-578. Ross, H. H. 1944. The caddis flies, or Trichoptera, of Illinois. Illinois Natural History Survey Bulletin 23: 1-326. Russell, L. M. 1993. A new species of Trialeurodes (Homoptera: Aleyrodidae) from Phlox. Proceed- ings of the Entomological Society of Washington 95: 583-586. Smith, D. M. and D. A. Levin. 1967. Karyotypes of eastern North American phlox. American Journal of Botany 54: 324-334. Smith, T. L. 1989. Pennsylvania habitats, pp. 52-56. In Cuff, D. J. et al., eds., The Atlas of Pennsyl- vania. Temple University Press, Philadelphia. Snodgrass, G. L., T. J., Henry, and W. P. Scott. 1984. An annotated list of the Miridae (Heteroptera) found in the Yazoo-Mississippi Delta and asso- ciated areas in Arkansas and Louisiana. Proceed- VOLUME 97, NUMBER 2 ings of the Entomological Society of Washington 86: 845-860. Stannard, L. J. 1968. The thrips, or Thysanoptera, of Illinois. Illinois Natural History Survey Bulletin 29(4): 215-552. Stout, A. B. 1917. Variation in the moss pink, Phlox subulata. Journal of the New York Botanical Gar- den 18: 75-83. Tepedino, V. J. and N. L. Stanton. 1976. Cushion plants as islands. Oecologia (Berl.) 25: 243-256. Tucker, G. E. 1972. The vascular flora of Bluff Moun- tain, Ashe County, North Carolina. Castanea 37: 2-26. Wheeler, A. G., Jr. 1988. Diabrotica cristata, a chrys- omelid (Coleoptera) of relict midwestern prairies discovered in eastern serpentine barrens. Ento- mological News 99: 134-142. . 1994. Craspedolepta eas: Distribution, hosts, and habits of a phlox specialist (Homoptera: Psyl- loidea: Aphalaridae). Proceedings of the Ento- mological Society of Washington 96: 91-97. Wheeler, A. G., Jr., T. J. Henry, and T. L. Mason, Jr. 1983. An annotated list of the Miridae of West Virginia (Hemiptera-Heteroptera). Transactions of the American Entomological Society 109: 127- 159. Wheeler, A. G., Jr. and E. R. Hoebeke. 1985. The insect fauna of ninebark, Physocarpus opulifolius (Ro- saceae). Proceedings of the Entomological Society of Washington 87: 356-370. 451 Wherry, E. T. 1929a. Picking out the Polemoniaceae. Bartonia 11: 1-4. . 1929b. The eastern subulate-leaved phloxes. Bartonia 11: 5-35. . 1930. Plants of the Appalachian shale-barrens. Journal of the Washington Academy of Sciences 20: 43-52. . 1935a. Fifteen notable shale-barren plants. Claytonia 2: 19-22. . 1935b. Our native phloxes and their horticul- tural derivatives. National Horticultural Maga- zine 14: 209-231. . 1935c. The Polemoniaceae of Pennsylvania. Proceedings of the Pennsylvania Academy of Sci- ence 9: 150-155. . 1935d. The Polemoniaceae of Virginia. Clay- tonia 2(1): 3-4, 2(2): 11-14. . 1936. Polemoniaceae of the middle Appala- chian region. Castanea 1: 13-15, 32-35. . 1951. Subspecies of three eastern phloxes. Cas- tanea 16: 97-100. . 1955. The Genus Phlox. Morris Arboretum Monographs 3. Philadelphia. 174 pp. . 1964. Some Pennsylvania barrens and their flora. II. Bartonia 34: 8-11. Winterringer, G. S. and A. G. Vestal. 1956. Rock- ledge vegetation in southern Illinois. Ecological Monographs 26: 105-130. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 452-457 A NEW GENUS OF MIRINE PLANT BUG, CARVALHOPANTILIUS, WITH TWO NEW SPECIES FROM TAIWAN (HETEROPTERA, MIRIDAE) TOMOHIDE YASUNAGA Biological Laboratory, Hokkaido University of Education Ainosato 5-3-1, Sapporo 002, Japan. Abstract.—A new genus of the Miridae, Carvalhopantilius, and two new species, C. purus and C. rufescens, are described from Taiwan. The genus is characterized by the elongate body, long antenna and rather delicate hemelytra. The vertical head and laterally carinate pronotum are similar to those found in Pantilius Curtis, to which the new genus seems to have the closest relationship. Key Words: The mirid fauna of Taiwan is poorly known. Only a few authors have described new taxa and/or presented faunal lists (e.g. Poppius 1914, Esaki 1926, Miyamoto and Yasunaga 1989, 1992, Yasunaga 1994). Needless to say, the Taiwanese fauna is very rich and in great need of investigation. Recently, through the courtesy of Mr. M. Tomokuni, of the National Science Muse- um, Tokyo, Japan, I had an opportunity to examine three specimens of two peculiar mirine plant bugs collected in Taiwan. These specimens were found to be two unde- scribed species that represent a new genus in the tribe Mirini of the subfamily Mirinae. Externally, they are similar to certain spe- cies of the genus Pantilius Curtis in having the vertical head and laterally carinate pro- notum, or to those of Megacoelum Fieber in the elongate, subparallel-sided dorsal habitus. But the male genital structures and several external characters sufficiently differ from those of the latter two genera, and, therefore, I describe them as two new spe- cies in a new genus. All measurements in the text are given in millimeters. Type specimens are all depos- Heteroptera, Miridae, new genus, new species, Taiwan ited in the collection of Department of Zo- ology, National Science Museum, Tokyo, Japan. Carvalhopantilius Yasunaga, New GENUS Body elongate, subparallel-sided, more than 8 mm in length; dorsal surface sparsely clothed with uniform pale setae. Head ver- tical; eyes in dorsal view rather small, re- moved from pronotal collar; vertex with a shallow, short, longitudinal, mesal sulca- tion, not carinate basally; frons projected anteriorly. Antenna long; segment I broad, provided with dark setae inwardly; segment II almost linear; segment III slightly shorter than II, longer than basal width of prono- tum. Rostrum relatively long, extending be- yond middle coxa. Pronotum weakly shining, minutely, shallowly, and sparsely punctate, weakly carinate laterally, sparsely set with pale erect or suberect setae; calli undeveloped; collar shagreened, narrow, with several pale, erect setae. Scutellum rather flat, sparsely set with pale, erect setae. Hemelytra semitranspar- ent, delicate, sparsely and minutely punc- VOLUME 97, NUMBER 2 tate, sparsely clothed with pale, suberect se- tae. Legs long; tibiae with brownish spines and erect pale setae; tarsomere III longer than I or II. Male genitalia (Fig. 2): Parameres lacking setae; left paramere semicircularly curved, triangularly projected basally (A); right par- amere short and straight, terminated in small apical claw (B). Vesica bilobed, with a re- markable elongate, apical spiculum termi- nating in apical hook and a thin ventral sclerite; gonopore opening rather large, with U-shaped sclerite above gonoporal rim (C & D). Type species.—Carvalhopantilius purus Yasunaga, new species. Etymology.— Named after the late Dr. J. C. M. Carvalho, in combination with ge- neric name Pantilius Curtis, to which this new genus seems to be related; gender mas- culine. Distribution. — Central Taiwan. Discussion. — This new genus is similar in some external characters to the Palearctic genus Pantilius Curtis, especially in having the vertical head and projected frons. How- ever, the following characters differ from those of the latter: the dorsal surface uni- formly clothed only with suberect setae and lacking silvery pubescence; antennal seg- ment II almost linear and not strongly in- crassate; antennal segments III and IV much longer; lateral carina of pronotum weak; he- melytra composed of rather delicate integ- ument; vesica with a remarkable elongate spiculum but lacking a pair of rounded api- cal spinose sclerites, which are always found in species of Pantilius (see Yasunaga 1992). Carvalhopantilius also resembles certain species of the genus Megacoelum Fieber in having the elongate, subparallel-sided dor- sal habitus, but the structures of head (es- pecially, the anteriorly projected frons), pronotum (presence of lateral carination), and genitalia are different. The new genus Carvalhopantilius is known by two conspicuous species endemic to the central mountain area of Taiwan. 453 Carvalhopantilius purus Yasunaga, NEw SPECIES (Figs. 1 & 2) Description.— Male: Body generally whitish, large; dorsal surface sparsely clothed with suberect pale setae. Head pale brown, shining; vertex 0.41 times as wide as head, glabrous, with a short, longitudinal, mesal sulcation; margin of antennal tubercle dark- ened; tylus, jugum, lorum, gena, and buc- cula bearing silky erect hairs. Antennal seg- ment I pale brown, with a dark brown stripe dorsally and several dark spots, bearing blackish setae; segment II pale brown, with dark narrow bands on basal *%% and apex; segment III pale brown, somewhat tinged with red at apical half, with narrow, dark reddish brown bands on basal 74 and apex; segment IV pale reddish brown, with pale brown base; length of segments I-IV: 1.60, 3.93, 3.70, 1.58. Rostrum pale brown, ex- cept dark apical part of segment IV, reach- ing hind coxae; length of segments I-IV: 09370290) OF Sales: Pronotum yellowish white, sparsely closed with pale, suberect, short setae, lateral ca- rina darkened; propleuron with a dark me- dian stripe. Mesoscutum and scutellum with a dark mesal stripe. Hemelytra yellowish white, semitransparent, with a dark mesal stripe along each inner margin of clavus; each corium with a small dark submarginal spots before midlength and one at cuneal fracture; lateral margin of embolium nar- rowly darkened; membrane pale brown, with 3 somber spots along posterior margin of vein. Legs pale brown; femora with small dark spots, bearing pale erect setae and some dark trichobothria; tibiae with pale erect se- tae and brownish spines; hind tibia with a dark spot at base and a narrow stripe from base to middle; apical parts of tarsomeres III somewhat darkened; length of hind fe- mur, tibia and tarsus: 4.10, 6.60, 0.90; that of hind tarsomeres I-III: 0.31, 0.38, 0.44. Abdomen pale brown; male genital seg- 454 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. I. ment with a mesal keel-like process poste- riorly. Male genitalia as mentioned in generic description (Fig. 2). Dimensions: Body length 11.10, head width 1.38, total rostral length 3.55, mesal pronotal length 1.38, basal pronotal width 2.73 and width across hemelytra 3.20. Female: Unknown. HOLOTYPE: 6, Wuling-Chika Shan- chuang, 1900-2400 m alt., Mt. Hsucshan, Hoping, Taichung Hsien, 13. Aug, 1990, M. Tomokuni. Etymology. — From the Latin, purus (pure Carvalhopantilius purus, holotype male. A, dorsal habitus. B, left lateral view of head and pronotum. or untainted), referring to the pure whitish general coloration. Distribution. —Central mountain area of Taiwan. Remarks. — This new species is easily rec- ognized by the large, elongate, whitish body, and such whitish coloration that at first ap- pears teneral is unusual within the Miridae. Yasunaga and Takai (1994) recently de- scribed Eocalocoris albicerus from Japan, which also exhibits yellowish-white general coloration. The latter species is associated with the white flowers of Clethra barbinervis (Clethraceae). Its cryptic whitish coloration VOLUME 97, NUMBER 2 455 Fig. 2. Male genitalia of Carvalhopantilius purus, holotype. A, left paramere. B, right paramere. C, vesica in dorsal view. D, vesica in ventral view. undoubtedly affords considerable protec- tion against natural enemies. No other in- formation is available on its ecology. Carvalhopantilius rufescens Yasunaga, NEw SPECIES (Fig. 3) Description.—Female: Body generally pale reddish brown, somewhat sanguine- Ous; setae on dorsum indistinct. Head red- dish brown, subshining, somewhat sha- greened, sparsely set with short erect hairs; vertex 0.38—0.39 times as wide as head, with a weak mesal longitudinal sulcation. Anten- nae almost unicolorously reddish brown: apical part of segment II and median and apical parts of III slightly darker; extreme base of segment IV yellowish; length of seg- ments I-IV: 1.23, 3.05-3.13, 2.90-2.98, 1.33-1.45. Rostrum pale reddish brown, 456 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. reaching middle coxa; apical half of segment IV darkened; length of segments I-IV: 0.73- 0.75, 0.73-0.75, 0.50-0.55, 0.80-0.88. Pronotum reddish brown, somewhat darkened medially and laterally, sparsely with shallow minute punctures, lateral mar- gin weakly carinate; setae on pronotum very short and sparse; ventral margin of pro- pleuron darkened. Mesoscutum and ante- rior margin of scutellum somewhat dark- ened medially. Hemelytra reddish brown, sparsely set with pale suberect short setae; lateral margin of corium narrowly darkened medially; apex of embolium darkened; cu- neus deep red or sanguineous; membrane somber pale brown, with partly reddish veins. Legs pale reddish brown; bases of femora paler; tibiae lacking erect setae; tarsi pale brown, except apical halves of tarso- meres III dark brown; length of hind femur, tibia and tarsus: 3.18-3.33, 5.08—5.30, 0.70- 0.73; that of hind tarsomeres I-III: 0.25, 0.31-0.36, 0.34—0.39. Abdomen reddish brown, widely dark- ened dorsally. Female genitalia (Fig. 3): Sclerotized ring thin-rimed, with a lateral pointed projec- tion, each ring contiguous to one another medially (A); posterior wall of bursa cop- ulatrix with rather wide interramal lobes and a distinct interramal sclerite (B). Dimensions: Body length 8.15-8.40, head width 1.28-1.30, total rostral length 2.75- 2.78, mesal pronotal length 1.20-1.23, basal Female genitalia of Carvalhopantilius rufescens, paratype. A, sclerotized ring in ventral view. B, posterior wall of bursa copulatrix in anterior view. pronotal width 2.33-2.48 and width across hemelytra 2.93. Male: Unknown. HOLOTYPE: 92, Wuling, 1900 m alt., Hoping, Taichung Hsien, 11. Aug. 1990, M. Tomokuni. PARATYPE: | 2, same data as for holotype. Etymology.—From the Latin, rufescens (becoming red), referring to the reddish gen- eral coloration. Distribution.—Central mountain area of Taiwan. Remarks.—This new species resembles C. purus, from which it is easily distin- guished by the reddish coloration of the body, the shorter and more sparse setae on dorsum, the long antennal segment IV that is longer than segment I, the shorter rostrum that is not reaching the hind coxa, and the hind tibiae lacking the erect setae, in ad- dition to being significantly smaller in size. There is no information on its ecology. ACKNOWLEDGMENTS I express my sincere gratitude to Dr. S. Miyamoto, my constant advisor of Fuku- oka City, Japan and to Mr. M. Tomokuni (National Science Museum, Tokyo, Japan) for the loan of specimens. I also thank Dr. T. J. Henry (Systematic Entomology Lab- oratory, USDA, Washington, D.C.) for in- viting me to participate in this volume hon- oring the late Dr. J. C. M. Carvalho and kindly reviewing the manuscript. VOLUME 97, NUMBER 2 LITERATURE CITED Esaki, T. 1926. Verzeichnis der Hemiptera-Heter- optera der Insel Formosa. Annales Musei Nation- alis Hungarici 14: 136-189. Miyamoto, S. and T. Yasunaga. 1989. Two new spe- cies of the Miridae (Heteroptera) from Japan and Taiwan. Japanese Journal of Entomology 57: 257- 263. 1992. A new mirid bug (Heteroptera: Miri- dae) from Taiwan. Esakia, Fukuoka (32): 93-96. Poppius, B. 1914. H. Sauter’s Formosa=Ausbeute: Nabidae, Anthocoridae, Termatophylidae, Miri- dae, Isometopidae und Ceratocombidae (Hemip- tera). Archiv fiir Naturgeshichte 8(A): 1-80. 457 Yasunaga, T. 1992. On the Palearctic genus Pantilius Curtis (Heteroptera: Miridae). Bulletin of the Bio- geographical Society of Japan 47: 109-116. . 1994. Review of the Sino-Japanese plant bug genus Parapantilius Reuter (Heteroptera, Miri- dae), with description of a new species from Tai- wan. Japanese Journal of Entomology 62: 683- 691. Yasunaga, T. and M. Takai. 1994. Review of the genus Eocalocoris Miyamoto et Yasunaga (Het- eroptera, Miridae) of Japan, with description of a new species from the mountains of Shikoku and southwestern Honshu. Proceedings of the Japa- nese Society of Systematic Zoology (52): 75-80. PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 458-473 A LIST OF THE MIRIDAE (HETEROPTERA) RECORDED FROM CHINA SINCE J. C. M. CARVALHO’S “WORLD CATALOGUE” LE-Y1 ZHENG Department of Biology, Nankai University, Tianjin 300071, P.R. China. Abstract.—This paper provides a list of the Miridae recorded from China since the publication of the J. C. M. Carvalho (1952-1960) “World Catalogue of Miridae.” A total of 423 species are listed, bringing the total number of Miridae known from China to 554 species. Bibliographical sources and provincial (or territorial) distribution data are given for each species. Key Words: Miridae, list, China This paper is dedicated to Dr. J. C. M. Carvalho in honor of his contribution to the systematics of Miridae, especially for his monumental “World Catalogue of Miri- dae,” which tremendously facilitated the studies of this large family all over the world and in China as well. Records of Chinese mirids since the pub- lication of the ““World Catalogue”’ are doc- umented herein, including a few unpub- lished records by the author. A total of 423 species are listed, bringing the total number of Miridae known from China to 554 spe- cies. Chinese records already found in Dr. Carvalho’s World Catalogue are marked with an asterisk. Original references are in- cluded for genera having the type species from China and type localities are given for species described from China. Distribution data (not including those given in the lit- erature), listed by province or region are given in brackets at the end of each species. Only literature since the Carvalho Catalog is cited in the bibliography. The subfamilial system used in this paper follows that of Henry and Wheeler (1988). Iam greatly indebted to Thomas J. Henry of (Systematic Entomology Lab., USDA, Washington, D.C.) for his encouragement, comments, English corrections, and kind assistance in publishing this paper. Abbreviations used in this paper are as follows: A = Anhui, C = China, CEN = Central China, F = Fujian, GD = Guang- dong, GS = Gansu, GX = Guangxi, GZ = Guizou, H = Hongkong, HA = Hainan, HEB = Hebei, HLJ = Heilongjian, NH = Henan, HUB = Hubei, HUN = Hunan, IM = Inner Mongolia, JL = Jilin, JS = Jiangsu, JX = Jiangxi, LN = Liaoning, N = Ningxia, NC = North China, NE = Northeast China, O.D. = original description, PT(S) = paratype(s), Q = Qinghai, SAX = Shaanxi, SC = Si- chuan, SD = Shandong, SX = Shanxi, T = Taiwan, unpub. = unpublished data, XJ = Xinjiang, XZ = Xizang (= Tibet), Y = Yun- nan, Z = Zhejiang, (!) = written in Chinese with English summary. Subfamily Bryocorinae Abibalus regulus Distant 1909. Zheng [un- pub.] [F] *Bryocoris convexicollis Hsiao 1941. Zheng [unpub.] [GS] Bryocoris flaviceps Zheng and Liu 1992 [OFD.- IWIN] Bryocoris gracilis Linnavuori 1962. Miya- moto and Yasunaga 1989b [T] VOLUME 97, NUMBER 2 Bryocoris hsiaoi Zheng and Liu 1992 [O.D.: HUN] Bryocoris pallidipes Zheng and Liu 1992 [O.D.: HUN] *Cyrtopeltis tenuis Reuter 1895. Hsiao and Meng 1963 [C] *Ernestinus pallidiscutum (Poppius) 1915. Miyamoto 1965 [T] *Harpedona fulvigenis (Poppius) 1915. Sto- nedahl 1988 [T] Harpedona marginata (Distant) 1904. Sto- nedahl 1988 [F] Helopeltis bradyi Waterhouse 1886. Zheng [unpub.] [HA (det. by Stonedahl)] *Helopeltis cinchonae Mann 1907 (= H. brevicornis Poppius 1915). Stonedahl 1991b [T] *Helopeltis fasciaticollis Poppius 1915. Sto- nedahl 1991b; Zheng [unpub. (det. by G. M. Stonedahl)] [GX, T, Y] Helopeltis theivora Waterhouse 1886. Sto- nedahl 1991b [C]; Zheng [unpub. (det. by G. M. Stonedahl)] [HA] Pachypeltis cinnamomi Zheng and Liu 1992 [O.D.: HUN] Pachypeltis politus (Walker) 1873. Zheng and Liu 1992 [HUN] Pachypeltis sassafri Zheng and Liu 1992 [O.D.: HUN] Pachypeltis wangi Zheng and X.-Z. Li 1992 [O.D.: HUN] Prodromus clypeatus Distant 1904. Stone- dahl 1988 [F, T, GD] Pseudodoniella chinensis Zheng 1992b [O.D:: GX] Subfamily Deraeocorinae *Alloeotomus chinensis Reuter 1903. Lin- navuori 1963; Kerzhner 1988; Zheng and X.-Z. Li 1992 [GX, JS, HEB] Alloeotomus simplus (Uhler) 1896. Miya- moto and Yasunaga 1989b; Zheng [un- pub.] [GS, HEB] Angerianus maurus Distant 1904. Stone- dahl 1991a [T] Genus Apilophorus Hsiao and Ren 1983 [O.D., type sp.: Apilophorus fasciatus Hsiao and Ren 1983] 459 Apilophorus fasciatus Hsiao and Ren 1983 [(O-D:3-Y¥] Bothynotus pilosus (Boheman) 1852. Zheng [unpub.] [LN] Genus Cimidaeorus Hsiao and Ren 1983 [O.D., type sp.: Cimidaeorus nigrorufus Hsiao and Ren 1983] Cimidaeorus nigrorufus Hsiao and Ren 1983 [O.D.: F] *Deraeocoris annulipes (Herrich-Schaeffer) 1842. Qi and Nonnaizab 1993a [IM] *Deraeocoris annulus Hsiao and Ren 1983 [O:D SE ¥] *Deraeocoris aphidicidus Ballard 1927. Zheng and Liu 1992 [HUN, SC] *Deraeocoris ater Jakovlev 1889. Qi and Nonnaizab 1993a; Kerzhner 1988; Zheng and Gao 1990 [GS, IM] Deraeocoris kerzhneri Josifov 1983. Ker- zhner 1988; Qi and Nonaizab 1993a [IM, NE] Deraeocoris koreanus Linnavuori 1963. Zheng and Liang 1991 [HEB] Deraeocoris lutescens (Schilling) 1837. Qi and Nonnaizab 1993a [IM] *Deraeocoris montanus Hsiao 1941. Zheng and Liu 1992 [HUN, HEB, SC] Deraeocoris morio (Boheman) 1852. Qi and Nonnaizab 1993a [IM] Deraeocoris olivaceus (Fabricius) 1777. Kerzhner 1988; Zheng and Gao 1990; Qi and Nonaizab 1993a [IM, N, NE] Deraeocoris omeiensis Hsiao and Ren 1983 [O:D:7 SE] Deraeocoris pallidicornis Josifov 1983. Qi and Nonnaizab 1993a [IM] *Deraeocoris punctulatus Fallen 1807. Hsiao and Meng 1962; Zheng and Gao 1990; Qi and Nonnaizab 1993a [IM, N, NC] Deraeocoris salicis Josifov 1983. Zheng and Gao 1990 [N] Deraeocoris scutellaris (Fabricius) 1794. Zheng and Gao 1990 [N] Deraeocoris serenus Douglas and Scott 1868. Zheng and Gao 1990; Qi and Nonnaizab 1993a [IM, N] Deraeocoris ventralis (Reuter) 1904. Qi and Nonaizab 1993a [IM] 460 Fingulus brevirostris Ren 1983 [O.D.: Y] Fingulus inflatus Stonedahl and Cassis 1991 [T] Fingulus /ongicornis Miyamoto 1965. Sto- nedah! and Cassis 1991 [T] Fingulus porrectus (Bergroth) 1916. Stone- dahl and Cassis 1991 [H] Fingulus ruficeps Hsiao and Ren 1983 [O.D.: SC] Nicostratus sinicus Hsiao and Ren 1983 [O.D.: HA] Genus Paranix Hsiao and Ren 1983 [O.D., type sp.: Paranix bicolor Hsiao and Ren 1983] Paranix bicolor Hsiao and Ren 1983 [O.D.: ¥] Stethoconus japonicus Schumacher 1910. Kerzhner 1988: [C] Termatophylum montanum Ren 1983 (ODA Termatophylum yunnanum Ren 1983 [O.D.: Y] Subfamily Isometopinae Isometopus beijingensis Ren and Yang 1988 [O.D.: HEB] Isometopus citri Ren 1987 [O.D.: SAX] Isometopus marginatus Ren and Yang 1988 [O:D.: Y] Isometopus nigrisignatus Ren 1987 [O.D.: Y] Isometopus puberus Ren 1991 [O.D.: Y] Isometopus shaowuensis Ren 1987 [O.D.: F] Letaba xizangana Ren 1988 [O.D.: XZ] Myiomma altica Ren 1987 [O.D. SC]; Ren 1992 [SC] Genus Paraletaba Ren and Yang 1988: [O.D., type sp.: Paraletaba montana Ren and Yang 1988] Paraletaba annulata Ren and Huang 1987 [O.D7"F] Paraletaba montana Ren and Yang 1988 [O°D;=GX] Sophianus lamellatus Ren and Yang 1988 [O.D.: GX] Subfamily Mirinae * Adelphocoris albonotatus (Jakovlev) 1881. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hsiao and Meng 1963 (Trichophoron- chus), Kerzhner 1988; Zheng and X.-Z. Lit 1989 [AGS HEBSHE J ie iss aoe SAX, SC] * Adelphocoris annulicornis (Sahlberg) 1848. Hsiao 1962 (as A. quadripunctatus (Fabr.)); Zheng and X.-Z. Li 1989; Qi et al. 1992 [GS, HEB, HLJ, IM, LN, N, SC, SJ, SX] * Adelphocoris apicalis Reuter 1906. Zheng and X.-Z. Li 1989; Qi et al. 1992 [GS, GZ, HEB ALS; HUB. IMIS Ie Ser7) * Adelphocoris divergens Reuter 1906. Zheng and X.-Z. Li 1989 [SC] *Adelphocoris fasciaticollis Reuter 1903. Hsiao 1962; Hsiao and Meng 1963; Zheng and X.-Z. Li 1989; Qi et al. 1992 [A, HA, HEB, HE), HN; HUB. IMe JE IS be LN, SAX, SC, SD, SX] * Adelphocoris fasciiger Reuter 1906. Zheng and X.-Z. Li 1989 [SC] Adelphocoris ferrugineus Hsiao 1962 [O.D.: HEB, SX]; Zheng and X.-Z. Li 1989; Qi et al. 1992 [HEB, IM, SX] * Adelphocoris funestus Reuter 1903. Zheng and X.-Z. Li 1989; Zheng and Liu 1992; Zheng and X. Z. Li 1992; Zheng [unpub.] [GS, HEB, HUB, SAX, SC] Adelphocoris fuscicornis Hsiao 1962 [O.D.: HEB, HN]; Zheng and X. Z. Li 1989 (HEB, HN, SX] Adelphocoris laeviusculus Vinokurov 1975. Zheng and X. Z. Li 1989; Qi et al. 1992 [HLJ, IM, SC] *Adelphocoris lineolatus (Goeze) 1778. Hsiao 1962; Hsiao and Meng 1963; Zheng and X.Z. Li 1989; Zheng and Gao 1990; Qi et.al: 1992: [GS,-GXe HEB? HEI. HIN: HUB, IMe Jie IS; XEN N: ORSAXS S@; SD: SX.Z| *4delphocoris luridus Reuter 1906. Hsiao 1962; Zheng and X. Z. Li 1989; Qi et al. 1992 [GS, IM, SC] * 4delphocoris melanocephalus Reuter 1903 (= A. transversus Lindberg 1934) Hsiao 1962; Zheng and X. Z. Li 1989 (synon- ymy with A. transversus Lindb.); Zheng VOLUME 97, NUMBER 2 and Gao 1990; Qi et al. 1992 [HEB, IM, LN, N, SX] Adelphocoris nigritylus Hsiao 1962 [O.D.: HEB, HN, SD]; Hsiao and Meng 1963; Zheng and X.Z. Li 1989; Zheng and Gao 1990; Zheng and X. Z. Li 1992; Qi et al. 1992 [A, GS, GZ, HA, HEB, HLJ, HN, Vie Sex IN, IN, SAX] SCY SD, SXG;Z] Adelphocoris obliquefasciatus Lindberg 1934. Kerzhner 1988; Zheng and X. Z. Li 1989; Qi et al. 1992; Zheng [unpub.] [HEB, HLJ, HUB, IM, LN] Adelphocoris piceosetosus Kulik 1965. Qi et al. 1992 [IM] Adelphocoris ponghvariensis Josifov 1977. Kerzhner 1988; Zheng and X. Z. Li 1989; Qi et al. 1992 [HEB, HLJ, IM, JL, JX, SAX, SD] Adelphocoris reicheli (Fieber) 1837 (= A. flavicornis Hsiao 1962). Hsiao 1962 [O.D. of A. flavicornis]; Zheng and X.-Z. Li 1989 (synonymy with 4. flavicornis Hsiao); Zheng and Gao 1990; Qiet al. 1992 [HEB, HLJ, IM, N, SD] Adelphocoris rufescens Hsiao 1962 [O.D.: HEB, HN, SD]; Kerzhner 1988; Zheng and X.-Z. Li 1989; Qi et al. 1992 [F, GZ, HEB, HLJ, HN, HUB, IM, SD, SX, Z] *Adelphocoris seticornis (Fabricius) 1775. Hsiao 1962; Zheng and X.-Z. Li 1989 [A, GS HEB HE) SC xd] *Adelphocoris suturalis Jakovlev 1882. Hsiao 1962; Hsiao and Meng 1963; Ker- zhner 1988; Zheng and X.-Z. Li 1989; Zheng and Liu 1992; Zheng and X.-Z. Li 1992-4 @i-et al: 19924[A, GS, ‘GX, "GZ, HEB; HE, HN; HUB, HUN, IM, JL, JS, TXSENESC, SD;Z] *Adelphocoris taeniophorus Reuter 1906. Zheng and X.-Z. Li 1989 [SC] Adelphocoris tenebrosus Reuter 1875. Hsiao 1962; Zheng and X.-Z. Li 1986; Kerzhner 1988; Qi et al. 1992; Zheng [unpub.] (HEB; HEJ,.IMs JL, LN; SX] Adelphocoris tibetanus Zheng and X.-Z. Li 1990 [O.D.: XZ] 461 * 4delphocoris torquatus Reuter 1906. Zheng and X.-Z. Li 1989 [SC] *Adelphocoris triannulatus (Stal) 1858. Hsiao 1962; Kerzhner 1988; Zheng and XZ; La 19892" Oiret aly 1992-(GS: HL; IM, JL] Adelphocoris yunnanensis Zheng and X.-Z. Li 1990 [O.D.: Y] Adelphocoris zoui Zheng and X.-Z. Li 1990 [OLDE SE] Allorhinocoris chinensis Lu 1994 (in Lu and Zhang 1994) [O.D.: GS, HEB, SX] Allorhinocoris flavus J. Sahlberg 1878. Lu and Zhang 1994 [XJ] Calocoris striatellus (Fabricius) (= C. och- romelas (Gmelin) 1788). Zheng and Gao 1990 [N] Capsus cinctus (Kolenati) 1845. Vinokurov 1977 [XJ] Capsus pilifer (Remane) 1950. Vinokurov 1977 [IM, HLJ] Capsus wagneri (Remane) 1950. Vinokurov 1977 [IM, HLJ] Charagochilus angusticollis Linnavuori 1961. Zheng 1990; Zheng and Liu 1992: Zheng and X.-Z. Li 1992; Zheng and Liu 1993 [A, F, GD, GX, GZ, HA, HEB, HN, HUB; HUN, x SAX {SC SXo YZ] *Charagochilus longicornis (Reuter) 1884. Zheng 1990; Zheng and X.-Z. Li 1992; Zheng and Liu 1993 [A, F, GD, GX, GZ, HUB SC, lg XZ.nYal Charagochilus pallidicollis Zheng 1990 [O:D::-F: GD; GX¢.GZ. HUBLI. SG Y]; Zheng and X.-Z. Li 1992 [same as O.D.] Charagochilus spiralifer Kerzhner 1987 (= Charagochilus similis Zheng 1990, syn. nov.). Zheng 1990 [O.D. of C. similis: HLJ] *Charagochilus taivanus (Poppius) 1915. Zheng 1990 (transfer from Proboscido- coris) [T] Charagochilus yulongensis Zheng 1990 [OID] Chrysorrhanis lineatus Carvalho 1979 [O.D.: HA (PTS)] 462 Creontiades bipunctatus Poppius 1915, Mi- yatomo and Yasunaga 1989b [T] Creontiades coloripes Hsiao 1963. Hsiao and Meng 1963 [O.D.: HN] Creontiades gossypii Hsiao 1963. Hsiao and Meng 1963 [O.D.: JX, Y] *Creontiades pallidifer (Walker) 1873. Mi- yamoto and Lee 1966 [T] Cyphodema inexpectata Zheng and Liu 1992 [O.D.: HUN] Dagbertus kirkaldyi (Poppius) 1915. Mi- yamoto and Yasunaga 1989b: [T] Dolichomiris antennatis (Distant) 1904. Zheng 1986, 1987; Zheng and Liu 1992; Zheng and X.-Z. Li 1992 [F, GD, HUB, HUN EX N-SAX= SG) YY. x7] Dolichomiris hirticornis Zheng 1986 [O.D.: Y]; Zheng 1988 [Y, XZ] Dolichomiris planiceps Zheng 1986 [O.D.: Y]; Zheng 1988 [Y, XZ] Genus E/themidea Zheng 1992a [O.D., type sp.: Elthemidea picea Zheng 1992] Elthemidea picea Zheng 1992a [O.D.: SC] Elthemidea sichuaense Zheng 1992a [O.D.: SC] Eurystylopsis chinensis Zheng and Chen 1991 [O.D.: SC, Y]; Zheng and Gao 1990 [INSSGey] *Eurystylopsis clavicornis (Jakovlev) 1890. Zheng and Chen 1991 (transfer from Cal- ocoris), Zheng and X.-Z. Li 1992 [F, GS, GX7GZ; HUB: SC, Y, Z] Eurystylus burmanicus (Distant) 1904. Zheng and Chen 1991 [Y] *Eurystylus coelestialium (Kirkaldy) 1902. Kerzhner 1988; Zheng and Chen 1991; Zheng and Liu 1992; Zheng and X.-Z. Li 1992 [F, GD, GX, HEB, HLJ, HUB, HUN, ISS EXssGSG.Z] *FEurystylus costalis Stal 1870. Zheng and Chen 1991; Zheng and Liu 1993 [A, F, Heb; HIN, JS; SCusDyY] *Eurystylus luteus Hsiao 1941. Zheng and Chen 1991. [A, F, GD, HA, JX, SC, Y] *Eurystylus sauteri Poppius 1915. Miya- moto and Yasunaga 1989b; Zheng and Chen 1991 [T] PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Genus Heterolygus Zheng and Yu 1990 [O.D., type sp.: Lygus trivittulatus Reuter 1906] *Heterolygus clavicornis (Reuter) 1906. Zheng and Yu 1990 (transfer from Lygus) [SC, SAX] *Heterolygus duplicatus (Reuter) 1903. Zheng and Yu 1990 (transfer from Char- agochilus), Zheng [unpub.] [GS, HUB, SGRY] Heterolygus flavoventris Zheng and Yu 1990 (OD! SSC] Heterolygus fusconiger Zheng and Yu 1990 [O:DSE] Heterolygus longus Zheng and Yu 1990 [Os ss :¥4] Heterolygus tenuicornis Zheng and Yu 1990 [O.D.: SC] *Heterolygus trivittulatus (Reuter) 1906. Zheng and Yu 1990 (transfer from Lygus) [SC, XZ] Heterolygus univittatus Zheng and Yu 1990 [O.Di: XZ] *Heterolygus validicornis (Reuter) 1906. Zheng and Yu 1990 (transfer from Ly- gus); Zheng [unpub.] [GS, SC] Heterolygus yadongensis Zheng and Yu 1990 [O.D.: XZ] Genus Heteropantilius Zheng and Liu 1992 [O.D., type sp.: Heteropantilius rhopali- morphus Zheng and Liu 1992] Heteropantilius rhopalimorphus Zheng and Liu 1992 [O.D.: HUN] Hyalopeplus spinosus Distant 1904. Zheng [unpub.] [GD, GX] Hyalopeplus vitripennis (Stal) 1855. Zheng [unpub.] [GX, HA] * Isabel ravana (Kirby) 1891. Zheng and Liu 1992; Zheng [unpub.] [F, GD, GZ, JX, SC] Lasiomiris picturatus Zheng 1986 [O.D.: F, Ser T] Lasiaomiris purpurascens Zheng 1986 (OFDe Yi] Leptopterna albescens (Reuter) 1891. Vi- nokurov 1981 [IM] VOLUME 97, NUMBER 2 *T eptopterna ferrugata (Fallén) 1807. Non- naizab and Jorigtoo 1993 [IM] Leptopterna griesheimae Wagner 1952. Nonnaizab and Jorigtoo 1993 [IM] Leptopterna kerzhneri Vinokurov 1981. Zheng 1986; Nonnaizab and Jorigtoo 1993 [HLJ, IM] Leptopterna magnospicula Lu and Tang 1987 [O.D.: N] Leptopterna xilingolana Nonnaizab and Jo- rigtoo 1993 [O.D.: IM] *Ziocoridae mutabilis Reuter 1904. Zheng and Liu 1992 [HUB, HUN, GS, GZ, SC] Loristes decoratus (Reuter) 1908. Kerzhner 1988; Zheng [unpub.] [LN] Lygidea illota Stal 1858. Kerzhner 1988 [NE] Lygocoris (Apolygus) angustus Zheng and Wang 1983 (Lygus) [O.D.: SC]; Lu and Zheng (in press) (transfer from Lygus) [SC] Lygocoris (Apolygus) badius Lu and Zheng (in press) (= Lygocoris (Apolygus) pubes- cens (Zheng and Wang) 1982, nom. preoccup.). Zheng and Wang 1982 (Ly- gus) [O.D. of Lygus pubescens: GD, SC}; Lu and Zheng (in press) [new name for Lygus pubescens Zheng and Wang] [GD, SC] Lygocoris (Apolygus) castaneus Zheng and Wang 1983 (Lygus) [O.D.: SC]; Lu and Zheng (in press) (transfer from Lygus) [SC] Lygocoris (Apolygus) concinnus Wang and Zheng 1982 (Lygus) [O.D.: F]; Lu and Zheng (in press) (transfer from Lygus) [F] Lygocoris (Apolygus) curvipes Zheng and Wang 1982 (Lygus) [O.D.: HUB]; Lu and Zheng (in press) (transfer from Lygus) [HUB] Lygocoris (Apolygus) elegans Zheng and Wang 1982 (Lygus) [O.D.: HA]; Lu and Zheng (in press) (transfer from Lygus) [HA] Lygocoris (Apolygus) emeia Zheng and Wang, 1982 (Lygus) [O.D.: SC]; Lu and Zheng (in press) (transfer from Lygus) [SC] Lygocoris (Apolygus) evonymi Zheng and Wang 1983 (Lygus) [O.D.: SAX]; Lu and 463 Zheng (in press) (transfer from Lygus) [SAX] Lygocoris (Apolygus) fujianensis Wang and Zheng 1982 (Lygus) [O.D.: F]; Lu and Zheng (in press) (transfer from Lygus) [F] Lygocoris (Apolygus) hainanensis Zheng and Wang 1983 (Lygus) [O.D.: HA]; Lu and Zheng (in press) (transfer from Lygus) [HA] Lygocoris (Apolygus) hilaris (Horvath) 1905. Zheng and Wang 1983 (Lygus) [Z] *T ygocoris (Apolygus) lucorum (Meyer-Diir) 1845. Hsiao and Meng 1963 (Lygus) [NC—possibly a mixture of allied spe- cies]; Zheng and Liu 1992; Zheng [un- pub.] [GS, HUN, LN, LN] Lygocoris (Apolygus) major Zheng and Wang 1983 (Lygus) [O.D.: SC]; Lu and Zheng 1992 (in press) (transfer from Lygus) [SC] Lygocoris (Apolygus) marginatus Zheng and Wang 1982 (Lygus) [O.D.: SC]; Lu and Zheng (in press) (transfer from Lygus) [SC] Lygocoris (Apolygus) mosaicus Zheng and Wang 1982 (Lygus) [O.D.: GD]; Lu and Zheng (in press) (transfer from Lygus) [GD] Lygocoris (Apolygus) nigricans Wang and Zheng 1982 (Lygus) [O.D.: F]; Lu and Zheng (in press) (transfer from Lygus) [F] Lygocoris (Apolygus) nigritulus (Linnavuo- ri) 1961. Zheng and Wang 1983 (Lygus); Zheng and X.-Z. Li 1992 [F, GD, GX, HUN, SC] *T ygocoris (Apolygus) nigrocinctus (Reuter) 1906. Kerzhner 1972 (sp. distinct. = L. pulchellus var. nigrocinctus Reuter) [SC] Lygocoris (Apolygus) nigrovirens Kerzhner 1987. Kerzhner 1988 [CEN] Lygocoris (Apolygus) ornatus Zheng and Wang 1983 (Lygus) [O.D.: HUB]; Lu and Zheng (in press) (transfer from Lygus) [HUB] Lygocoris (Apolygus) picturatus Zheng and Wang 1982 (Lygus) [O.D.: GX]; Lu and Zheng (in press) (transfer from /ygus) [GX] *Tygocoris (Apolygus) pulchellus Reuter 1906. Linnavuori 1961 (Lygus) (sp. dis- 464 tinct., in Hsiao 1942 as synonym of L. adustus); Linnavuori 1963 (Lygus); Ker- zhner 1972 (transfer from Lygus) [SC] Lygocoris (Apolygus) signatus Zheng and Wang 1982 (Lygus) [O.D.: SC]; Lu and Zheng (in press) (transfer from Lygus) [SC] Lygocoris (Apolygus) triangulus Zheng and Wang 1983 (Lygus) [O.D.: HUB]; Lu and Zheng (in press) (transfer from Lygus) [HUB] Lygocoris (Apolygus) ulmicolus Lu and Zheng (in press) (= Lygocoris (Apolygus) ulmi (Zheng and Wang) 1983, nom. preoccup.). Zheng and Wang 1983 (Ly- gus) [O.D.: HEB, HN, HUB]; Lu and Zheng (in press) (new name for /ygus ulmi Zheng and Wang) [HEB, HN, HUB] Lygocoris (Apolygus) yunnananus Zheng and Wang 1982 (Lygus) [O.D.: Y]; Lu and Zheng (in press) (transfer from Lygus) [Y] *L ygocoris (Arbolygus) dasypterus (Reuter) 1906. Kerzhner 1979 (transfer from Ly- gus); Zheng [unpub.] [HUN, SC] *Tygocoris (Arbolygus) potanini (Reuter) 1906. Kerzhner 1979 (transfer from Ly- gus); Kerzhner 1988; Zheng and Gao 1990; Zheng [unpub.] [GS, LN, N, SC] Lygocoris (Arbolygus) pronotalis Zheng and Liu 1992 [O.D.: HUN] Lygocoris (Arbolygus) rubripes (Jakovlev) 1876. Zheng and Gao 1990 [N] Lygocoris (Arbolygus) ulmi Kerzhner 1979 [O.D.: HLJ (PT)]; Kerzhner 1988 [HLJ] *T ygocoris (Lygocoris) longipennis (Reuter) 1906. Kerzhner 1972 (transfer from Ly- gus) [SC] *L ygocoris (Lygocoris) rugosicollis (Reuter) 1906. Kerzhner 1972 (transfer from Ly- gus) [SC] *Tygocoris (Lygocoris) striicornis (Reuter) 1906. Kerzhner 1972 (transfer from Ly- gus) [SC] Lygus adspersus (Schilling) 1836. Qi 1993 [IM] *Lygus discrepans Reuter 1906. Zheng 1990: 16; Zheng and Yu 1992; Qi 1993; Zheng [unpub.] [GS, HEB, IM, N, SC, Y] PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON *Tygus gemellatus (Herrich-Schaeffer) 1835. Zheng and Gao 1990; Zheng and Yu 1992; Qi 1993 [GS, IM, N, SAX, SX, XJ] Lygus hsiaoi Zheng and Yu 1992 [O.D.: XZ] Lygus paradiscrepans Zheng and Yu 1992 [O:D:> GS; SG} Y-xXxZ] Lygus poluensis (Wagner) 1967. Zheng and Yu 1992 (transfer from Exolygus) [IM, SAX, XJ, XZ] *T_ ygus pratensis (Linnaeus) 1758. Hsiao and Meng 1963 [NC—possibly a mixture of allied species]; Zheng and Yu 1992 [SC, XJ—The HN & IM records in Hsiao 1942 and GS record in Lindberg 1934 need verification] *Lygus punctatus (Zetterstedt) 1840. Zheng and Gao 1990; Zheng and Yu 1992; Qi 1993 [GS, HEB, HLJ, IM, JL, N, SC, XZ] *Tygus rugulipennis (Poppius) 1807. Josi- fov and Kerzhner 1972 (as L. disponsi Linnavuori); Zheng and Gao 1990; Zheng and Yu 1992; Qi 1993 [HEB, HLJ, HN, IM, LN, N, SJ] Lygus saundersi Reuter 1906. Kerzhner 1988; Zheng and Gao 1990; Zheng [un- pub.] [GS, N] Lygus sibiricus (Aglyamzyanov) 1989; Qi 1993; Zheng [unpub.] [IM, HLJ, GS, SC, XJ] Lygus tibetanus Zheng and Yu 1992 [O.D.: XZ] Lygus wagneri (Remane) 1955. Zheng and Yu 1992; Qi 1993 [HLJ, IM, SC, XJ] *Macrolonius schenklingi (Poppius) 1915. Carvalho 1979 [T] *Mecistoscelis scirtetoides Reuter 1891. Zheng [unpub.] [Y] Megacoelum chinensis X.-Z. Li and Zheng 1991 [O.D.: F, GD, Z]; Zheng and X.-Z: Li 1992 [F, GD, HUB, HUN, Z] *Megacoelum clypeale Poppius 1915. X.-Z. Li and Zheng 1991 [T] Megacoelum fuscescens Hsiao 1963, Hsiao and Meng 1963 [O.D.: Y], X.-Z. Li & Zheng 1991 [GD, GX, HA, Y] *Megacoelum minutum Poppius 1915. X.- Z. Li and Zheng 1991 [T] VOLUME 97, NUMBER 2 *Megacoelum picea (Reuter) 1906. Zheng and X.-Z. Li 1992; Zheng [unpub.] [GS, HUB, SC] Megacoelum pronotalis X.-Z. Li and Zheng 1991 [O.D.: JX, SAX, Z] Megacoelum pseudoprontalis X.-Z. Li and Zheng 1991 [O.D.: F] Megacoelum rubripedum X.-Z. Li and Zheng ISOS TOsDiA FI Megacoelum tenuicorne X.-Z. Li and Zheng 1991 [O.D.: GD, GX, SC]; Zheng and Liu 1992 [GD, GX, HUN, SC] Megacoelum yunnananum X.-Z. Li and Zheng 1991 [O.D.: Y] Megacoelum zoui X.-Z. Li and Zheng 1991 [O.D= ¥] *Mermitelocerus annulipes Reuter 1908. Kerzhner 1972, 1988; Zheng [unpub.] [GS, HLJ] Mystilus priamus Distant 1904. Zheng [un- pub.: Y] *Notostira poppiusi Reuter 1911. Zheng 1987 [XZ] Notostira sibirica Golub 1978. Kerzhner 1988 [NE] Onomaus coloratus Zheng and Liu 1992 [O.D.: HUN] Onomaus lautus (Uhler) 1896. Zheng and Liu 1992 [HUN, HUB] *Orthops kalmi (Linnaeus) 1758. Hsiao and Meng 1963 (Lygus) [HN —identification doubtful] *Orthops lindbergi (Hsiao) 1941. Kerzhner 1987 (transfer from Lygus) [SC] *Orthops minutus (Hsiao) 1941. Kerzhner 1987 (transfer from Lygus) [SC] *Orthops mutans (Stal) 1858. Zheng and Gao 1990 [N] Orthops sachalinus (Carvalho) 1959. Ker- zhner 1988; Zheng and Gao 1990 [N, NE] Genus Paramiridius Miyamoto and Yasun- aga 1992 [O.D., type sp.: Paramiridius tigrinus Miyamoto and Yasunaga 1992] Paramiridius trigrinus Miyamoto and Ya- sunaga 1992 [O.D.: T] Parapantilius flavomarginatus Miyamoto and Yasunaga 1989a [O.D.: T] 465 *Parapantilius flavomarginatus Miyamoto and Yasunaga 1989a [O.D.: T] *Parapantilius thibetanus Reuter 1903. Zheng and Gao 1990 [N]; Zheng and Liu 1992, 1993 [GS, HUB, HUN, N, S, SC] Phytocoris (Ktenocoris) alashanensis Non- naizab and Jorigtoo 1992 [O.D.: IM] Phytocoris (Ktenocoris) caraganae Nonnai- zab and Jorigtoo 1992 [O.D.: IM] Phytocoris (Ktenocoris) desertorum Non- naizab and Jorigtoo 1992 [O.D.: IM] Phytocoris (Ktenocoris) insignis Reuter 1876. Nonnaizab and Jorigtoo 1992 [IM] Phytocoris (Ktenocoris) issykensis Poppius 1912. Nonnaizab and Jorigtoo 1992 [O.D.: IM] Phytocoris (Ktenocoris) mongolicus Non- naizab and Jorigtoo 1992 [O.D.: IM] Phytocoris (Ktenocoris) nigritus Nonnaizab and Jorigtoo 1992 Jorigtoo 1992 [IM] Phytocoris (Ktenocoris) rubiginosus Non- naizab and Jorigtoo 1992 [O.D.: IM] *Phytocoris (Phytocoris) intricatus Flor 1860. Nonnaizab and Jorigtoo 1992 [IM, SC] Phytocoris (Phytocoris) longipennis Flor 1860. Nonnaizab and Jorigtoo 1992 [IM] Phytocoris (Phytocoris) populi (Linnaeus) 1758. Nonnaizab and Jorigtoo 1992 [IM] Phytocoris (Phytocoris) procerus Nonnaizab and Jorigtoo 1992 [O.D.: IM] Phytocoris (phytocoris) zhengi Nonnaizab and Jorigtoo 1992 [O.D.: IM] *Phytocoris knighti Hsiao 1941. Zheng [un- pub.: GS] Polymerus brevicornis (Reuter) 1879. Zheng and X.-Z. Li 1986; Zheng [unpub.] [IM, HEJZEN] Polymerus carpathicus Horvath 1861. Zheng and X.-Z. Li 1986 [HLJ, IM] *Polymerus cognatus (Fieber) 1858. Hsiao and Meng 1963; Zheng and X.-Z. Li 1986; Kerzhner 1988 [GS, HEB, HLJ, HN, IM, SAX, SC, SD, SX, XJ] *Polymerus funestus (Reuter) 1906. Zheng and X.-Z. Li 1986 [SC, XZ] Polymerus nigritus (Fallen) 1807. Zheng and X.-Z. Li 1986 [XJ] 466 Polymerus palustris (Reuter) 1905. Zheng and <-2, Li 1986 (HUI; JE) * Polymerus pekinensis Horvath 1900. Zheng and X.-Z. Li 1986; Kerzhner 1988; Zheng and Gao 1990 [A, F, HEB, HLJ, IM, JX, MESAX, SC, SD; SKeZ] *Polymerus unifasciatus (Fabricius) 1794. Zheng and X.-Z. Li 1986 [GS, HEB, IM, SC, XJ] Polymerus vulneratus (Wolff) 1801. Zheng and X.-Z. Li 1986 [XJ] * Proboscidocoris malayus Reuter 1907. Mi- yamoto and Lee 1966 [T]; Zheng 1990; Zheng and Liu 1992; Zheng and Liu 1993 [AL FE) GDyGXsHAsHUB YM HUING 1x. S@y da * Salignus distinguendus Reuter 1875. Ker- zhner 1979, 1988; Zheng [unpub.] [GS, SC] *Salignus duplicatus (Reuter) 1906. Ker- zhner 1979 (sp. distinct., = Lygus distin- guendus var. duplicatus Rt.); Kerzhner 1988 [GS, SC] *Stenodema alpestre Reuter 1904. Zheng 1981b; Zheng and Gao 1990; Zheng 1992a [FP HUB MeN: SAX, SGV] Stenodema alticola Zheng 198 1a [O.D.: SC]; Zheng 1992a [SC, Y] Stenodema angustatum Zheng 198 1a [O.D.: XZ]; Zheng 1981b, 1987, 1988 [XZ] Stenodema antennatum Zheng 198 1a [O.D.: XZ]; Zheng 1981b [XZ] Stenodema calcaratum (Fallén) 1807. Zheng 198la, b; Nonnaizab & Jorigtoo 1994 (RET. IM, JL) * Stenodema chinense Reuter 1904. Zheng 1981b, 1992a [SC] Stenodema crassipes Kiritschenko 1931. Nonnaizab and Jorigtoo 1994 [IM] Stenodema daliensis Zheng 1992a [O.D.: Y] Stenodema deserta Nonnaizab and Jorigtoo 1994 [O.D.: IM] *Stenodema elegans Reuter 1904. Zheng 1981b, 1992a; Zheng and Gao 1990; Zheng and X.-Z. Li 1992; Zheng and Liu 1993; Zheng [unpub.] [F, GS, GZ, HUB, HUN, JX,.N, SAX, SCT, Y,.Z] PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Stenodema holsatum (Fabricius) 1787. Zheng 198la, b; Muminov 1989; Non- naizab and Jorigtoo 1994 [IM, SC, XJ] Stenodema hsiaoi Zheng 1981b [O.D.: SC] *Stenodema laevigatum (Linnaeus) 1758. Zheng 1981b; Nonnaizab and Jorigtoo 1994 [GS, IM] *Stenodema longicolle Poppius 1915. Zheng 1981b; Zheng [unpub.] [GD, T] Stenodema longulum Zheng 1981la [O.D.: JX, SC]; Zheng 1988; Zheng and X.-Z. Li 1992. (GZ; JXeSE.XZ) Stenodema mongolica Nonnaizab and Jo- rigtoo 1994 [O.D.: IM] Stenodema nigricallum Zheng 198 1a [O.D.: SC]; Zheng 1981b, 1992a; Zheng and Gao 1990; Zheng [unpub.] [GS, N, Y] Stenodema parvulum Zheng 1981la [O.D.: SC, XZ]; Zheng 1981b, 1987, 1992a [SC, XZ] Stenodema pilosa (Jakovlev) 1889. Mumi- nov 1989; Nonnaizab and Jorigtoo 1994 [IM, NE, XJ] Stenodema qinlingensis Tang 1994 [O.D.: SAX] *Stenodema plebejum Reuter 1904. Zheng 1981b; Zheng [unpub.] [GS, SC] Stenodema sericans Fieber 1861. Nonnai- zab and Jorigtoo 1994 [IM] Stenodema sibiricum Bergroth 1878. Zheng 198la, 1981b; Kerzhner 1988; Muminov 1989; Nonnaizab and Jorigtoo 1994, [IM, JL] Stenodema tibetum Zheng 198la [O.D.: XZ]; Zheng 1981b, 1987 [XZ] Stenodema trispinosum Reuter 1904. Zheng 1981la, b; Nonnaizab and Jorigtoo 1994 [IM, N, SC, XJ] Stenodema turanicum Reuter 1904. Zheng 1981b [as S. virens (L.), misidentified]; Muminov 1989; Zheng and Gao 1990; Nonnaizab and Jorigtoo 1994 [IM, N, Q, XJ] Stenodema virens (Linnaeus) 1767. Non- naizab and Jorigtoo 1994 [IM] Stenotus rubrovittatus (Matsumura) 1913. Hsiao and Meng 1963 [HN, HUB] VOLUME 97, NUMBER 2 *Tailorilygus apicalis (Leston) 1952. Mi- yamoto and Lee 1966; Zheng [unpub.] [GX, T] Teratocoris saundersi Douglas and Scott 1869. Jorigtoo and Nonnaizab 1991 [IM] *Trigonotylus caelestialium (Kirkaldy) 1902. Carvalho and Wagner 1957; Zheng 1985; Zheng and Gao 1990 [HEB, HLJ, HN, HUB wIM SIL JS. JX, LN; N,, SC,. SD; See Yi Trigonotylus cremeus Golub 1989 [O.D.: IM (PTS)] Trigonotylus fuscitarsis Lammes 1987. Jo- rigtoo and Nonnaizab 1993 [IM] Trigonotylus longitarsis Golub 1989 [O.D.: IM, HLJ] Trigonotylus major Zheng 1985 [O.D.: XJ]; Golub 1989 [Q, XJ] Trigonotylus pallescens Golub 1989. Jorig- too and Nonnaizab 1993 [IM] Trigonotylus pilipes Golub 1989. Jorigtoo and Nonnaizab 1993 [IM] Trigonotylus procerus Jorigtoo and Non- naizab 1993 [O.D.: IM] Trigonotylus pulchellus (Hahn) 1834. Jorig- too and Nonnaizab 1993 [IM] *Trigonotylus ruficornis (Geoffroy) 1785. Hsiao and Meng 1963 [NC—misidenti- fication of T. caelestialium]; Zheng 1985 [IM] Trigonotylus tenuis Reuter 1893. Zheng 1985 (as T. doddi (Distant)) [F, GD, GX, HUBIIXG SG. 1..Y.Z) Trigonotylus viridis (Provencher) 1827. Zheng 1985 (as T. bianchi Kiritschenko 1926); Zheng 1992a (as T. bianchi) [SC] Trigonotylus yangi Tang 1994 [O.D.: GX] Genus Zhengiella Yasunaga and Lu 1994 [O.D., type sp.: Zhengiella scutellata Ya- sunaga and Lu 1994] Zhengiella scutellata Yasunaga and Lu 1994 (OLD? Gx] Subfamily Orthotylinae Campylotropis jakovlevi Reuter 1904. Ker- zhner 1988 [IM, NE] Cyllecoris equestris Stal 1858. Kerzhner 1988; Zheng and Gao 1990 [N] 467 *Cyrtorrhinus lividipennis Reuter 1884. Hsiao and Meng 1963; Zheng and Liu 1992 [A, F, GD, HA, HN, HUN, JX, Z] Dryophilocoris alni Zou 1986 [O.D.: SC] Dryophilocoris limbatus Zou 1986 [O.D.: SC] Dryophilocoris longus Zou 1986 [O.D.: SC] *Ectmetopterus micantulus (Horvath) 1905 (= E. angusticeps Reuter). Josifov and Kerzhner 1972; Zheng and Liu 1992; Zheng and Liu 1993 [F, HEB, HUN, JS, SC, Z] Halticus apterus (Linnaeus) 1758. Zheng and Gao 1990 [N] Halticus comitans Josifov and Kerzhner 1972 [O.D.: HEB]; Kerzhner 1988 [HEB] Halticus fuscous Zou 1985 [O.D.: JX] Halticus maculipes Zou 1985 [O.D.: HUB, SE] *Halticus minutus Reuter 1884. Hsiao and Meng 1963; Zheng [unpub.] [A, HUN, JS, SAK. SCGe7Z] Halticus niger Zou 1985 [O.D.: GZ] Mecomma ambulans (Fallén) 1807. Liu and Zheng 1993 [Y, SC] *Mecomma chinensis Reuter 1906. Liu and Zheng 1993 [SC] Mecomma capitata Liu and Zheng 1993 [O;D:: SG] Mecomma gansuana Liu and Zheng 1993 [O.D.: GS] Mecomma opaca Liu and Zheng 1993 [O.D.: SC] *Orthocephalus funestus Jakovlev 1881. Kerzhner 1988; Zheng and Gao 1990; Zheng [unpub.] [GS, N] *Orthotylus (Melanotrichus) flavosparsus Sahlberg 1842. Hsiao and Meng 1963 [HEB, HN, SD] Orthotylus interpositus Schmidt 1938. Zheng and Gao 1990 [N] Pseudoloxops guttatus Zou 1987b [O.D.: HN, SD]; Zheng and Liang 1991 [HN, SD] Pseudoloxops marginatus Zou 1987b [O.D.: GD] 468 Genus Reuterio/a Hsiao 1963 [O.D., type sp.: Reuteriola annulicornis Hsiao 1963] Reuteriola annulicornis Hsiao 1963. Hsiao and Meng 1963 [O.D.: Y] * Strongylocoris leucocephalus (Linnaeus) 1758. Zheng and Gao 1990 [N] Strongylocoris niger (Herrich-Schaeffer) 1835. Zheng [unpub.] [XJ] Ulmica baicalica (Kulik) 1965. (= Mala- cocoris baicalica). Kerzhner 1987; Ker- zhner 1988 [HLJ] Genus Ulmocyllus Seidenstucker 1964 [O.D., type sp.: Ul/mocyllus virens Seiden- stucker 1964] Ulmocyllus virens Seidenstucker 1964. Sei- denstucker 1964 [O.D.: HLJ]; Kerzhner 1987 [C] Zanchius marmoratus Zou 1987a [O.D.: Y] Zanchius mosaicus Zheng and Liang 1991 [O.D.: HEB] Zanchius quinquemaculatus Zou 1987a [O:D Y] Zanchius tarasovi Kerzhner 1987. Zheng and Liang 1991 [HEB] Zanchius zoui Zheng and Liu 1993 [O.D.: Y] Subfamily Phylinae Acrorrhinium hongkong Schuh 1984 [O.D.: H] Acrotelus pilosicornis (Reuter) 1901. Ker- zhner 1962 [HLJ, IM] Atomophora flavidus Nonnaizab and Yang 1994 [O.D.: IM] Atomophora punctulatus Nonnaizab and Yang 1994 [O.D.: IM] Atomoscelis asiaticus Josifov 1979. Zheng and Gao 1990 [N]; Nonnaizab 1992 [IM] Atomoscelis onustus (Fieber) 1961. Non- naizab 1992 [IM] Atomoscelis pubescens Nonnaizab 1992 [O.D.: IM] Camptotylus reuteri Jakovlev 1881. Hsiao and Meng 1963 [HEB, SD] Campylomma chinensis Schuh 1984 [O.D.: F. H] *Campylomma diversicornis Reuter 1878. Hsiao and Meng 1963; Zheng and Gao 1990 [HEB, HN, N, SAX, SC, XJ] PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON *Campylomma livida Reuter 1884. Miya- moto and Yasunaga 1989b [T, C] Campylomma nicolasi Puton and Reuter 1883. Hsiao and Meng 1963 [HN, HUB, JXeSECPY] *Chlamydatus pullus Reuter 1870. Hsiao and Meng 1963; Zheng and Gao 1990 [GS, HEB, HN, JL, N, SAX] Cleotomiris chinensis Schuh 1984 [O.D.: F] Compsidolon gobicus Nonnaizab and Yang 1994 [O.D.: IM] Compsidolon pumilus (Jakovlev) 1876. Nonnaizab and Yang 1994 [IM] *Decomia cephalotes Poppius 1915. Schuh 1984 [T] Decomioides schneirlai Schuh 1984 [O.D.: T, H] *Druthmarus coxalis (Reuter) 1891. Schuh 1984 (transfer from Atractotomus) [T] *Ellenia obscuricornis (Poppius) 1914. Schuh 1984 [T] Eumecotarsus breviceps (Reuter) 1878. Ker- zhner 1962; Zheng [unpub.] [XJ] Eumecotarsus chinensis Kerzhner 1962 [O.D:: SG] Excentricoris pictipes Reuter 1897. Ker- zhner 1988 [HLJ] Glaucopterum gobicum Kerzhner 1984 [O.D.: IM]; Zheng [unpub.] [GS, IM] Hallodapus albofasciatus (Motschulsky) 1863. Schuh 1984 [H] *Hallodapus brunneus Poppius 1915. Schuh 1984 [T] Hallodapus centrimaculatus (Poppius) 1914. Schuh 1984 [H] Hallodapus fasciatus (Poppius) 1909. Schuh 1984 [H] Hallodapus fenestratus Linnavuori 1961. Miyamoto and Lee 1966; Schuh 1984 [T] *Hallodapus persimilis (Poppius) 1915. Schuh 1984 [T] Hallodapus ravenar (Poppius) 1914. Schuh 1984 [H] Opuna annulatus (Knight) 1935. Schuh 1984 evel 18) Pherolepis aenescens (Reuter) 1901. Ker- zhner 1970, 1988 [LN] VOLUME 97, NUMBER 2 Phylus coryloides Josifov and Kerzhner 1972 [O.D.: HLJ]; Kerzhner 1988 [HLJ] Pilophorus alstoni Schuh 1984. Zou 1989 (GDASC. Y] Pilophorus aureus Zou 1983 [O.D.: HEB] Pilophorus bistriatus Zou 1987c [O.D.: Y] Pilophorus castaneus (Zou) 1983 (Stricto- tergum) [O.D.: Y]; Schuh 1989 (Synon- ymy) [Y] Pilophorus dailahn Schuh 1984. Zou 1989 [Y] Pilophorus decimaculatus Zou 1983 [O.D.: yal *Pilophorus formosanus Poppius 1914. Schuh 1984 [T] Pilophorus gallicus Remane 1954. Zou 1989 [SC] Pilophorus koreanus Josifov 1977. Zou 1989 [HN] Pilophorus latus Zou 1989 [O.D.: Y] Pilophorus lucidus Linnavuori 1962. Zou 1989 [HN, HUB] Pilophorus niger Poppius 1914. Schuh 1984 [HU] Pilophorus setulosus Horvath 1905. Zou 1989; Zheng and Gao 1990 [IM, N] Pilophorus typicus (Distant) 1909. Schuh 1984 [F, H, T] Pilophorus yunganensis Schuh 1984 [O.D.: F, H] Pilophorus vitellinus Zou 1989 [O.D.: Y] Plagiognathus (Plagiognathus) alashanen- sis Qi and Nonnaizab 1993b [O.D.: IM] Plagiognathus (Plagiognathus) amurensis Reuter 1883. Hsiao and Meng 1963 [O.D. of P. nigricornis]; Kerzhner 1988; H.-Y. Li and Zheng 1991b (Synonymy, = P. nigricornis Hsiao and Meng 1963) [GZ, HA HEB, HES; HN; 3x; SD; SX] *Plagiognathus (Plagiognathus) arbusto- rum (Fabricius) 1794. H.-Y. Liand Zheng 1991b [SC] *Plagiognathus (Plagiognathus) chrysan- themi (Wolff) 1804. Zheng and Gao 1990; H.-Y. Li and Zheng 1991b [HJL, IM, N, SC, XJ] Plagiognathus (Plagiognathus) collaris (Matsumura) 1912. Kerzhner 1988; Zheng and Gao 1990; H.-Y. Li & Zheng 1991b; 469 Zheng [unpub.] [GS, HEB, HLJ, IM, N, XJ] Plagiognathus (Plagiognathus) kiritschen- koi Kulik 1975. H.-Y. Liand Zheng 1991b [HLJ] Plagiognathus (Plagiognathus) leucopus Kerzhner 1979. H.-Y. Liand Zheng 1991b [HLJ] Plagiognathus (Plagiognathus) lividellus Kerzhner 1979. Zheng and Gao 1990; H.- Y. Li and Zheng 1991b [F, HLJ, N] *Plagiognathus (Plagiognathus) lividus Reuter 1906. Kerzhner 1979; Kerzhner 1988; H.-Y. Li and Zheng 1991b [HEB, HLJ, SC] Plagiognathus (Plagiognathus) obscuriceps (Stal) 1858. Qi and Nonnaizab 1993b [IM] Plagiognathus (Plagiognathus) pallescens Zheng and H.-Y. Li 1991 [O.D.: SC]; H.- Y. Li and Zheng 1991b [SC] Plagiognathus (Plagiognathus) yomogi (Mi- yamoto) 1969. H.-Y. Liand Zheng 1991b [GZ, HEB, HLJ, SC, Y] *Plagiognathus (Poliopterus) albipennis (Fallen) 1929. Zheng and Gao 1990; H.- Y. Li and Zheng 1991b. [F, HA; HEB, HL, NSC, SDeSxXe Yq Plagiognathus (Poliopterus) canoflavidus Qi and Nonnaizab 1993b [O.D.: IM] *Plagiognathus (Poliopterus) moestus Reu- ter 1906. H.-Y. Li and Zheng 1991b [SC] Psallopsis halostachydis Putshkov 1975. Nonnaizab and Yang 1994 [IM] Psallopsis kirgisicus (Becker) 1864. Non- naizab and Yang 1994 [IM] *Psallus alpestris Reuter 1906. H.-Y. Li and Zheng 1991a [SC] Psallus ater Josifov 1983. H.-Y. Li and Zheng 1991a [SC] Psallus betuleti (Fallén) 1776. H.-Y. Li and Zheng 1991a [IM] Psallus castanae Josifov 1983. H.-Y. Li and Zheng 1991a [SC] Psallus clarus Kerzhner 1988. H.-Y. Li and Zheng 1991a [SC] Psallus falleni Reuter 1883. Qi and Non- naizab 1994 [IM] Psallus flavescens Kerzhner 1987. Qi and Nonnaizab 1994 [IM] 470 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Psallus fukienanus Zheng and H.-Y. Li 1990 [O.D.: F]; H.-Y. Li and Zheng 1991a [F] Psallus guttatus Zheng and H.-Y. Li 1990 [O.D.: SC]; H.-Y. Li and Zheng 1991a [SC] Psallus hani Zheng and H.-Y. Li 1990 [O.D.: HN]; H.-Y. Li and Zheng 1991a [HN] *Psallus holomelas Reuter 1906. H.-Y. Li and Zheng 1991a [SC, Y] Psallus kerzhneri Qi and Nonnaizab 1994 [O.D.: IM] Psallus luridus (Reuter) 1878. H.-Y. Li and Zheng 1991a [HLJ] Psallus mali Zheng and H.-Y. Li 1990 [O.D.: GS, SAX]; H.-Y. Liand Zheng 1991a[GS, SAX] Psallus tonnaichanus Muramoto 1973. H.- Y. Li and Zheng 1991la [HUB] Psallus ulmi Kerzhner and Josifov 1966 [O:D:; HE (PTS): H.-Y Lisand Zheng 1991a [HEB, HLJ, IM, N, SAX] Psallus vittatus (Fieber) 1861. H.-Y. Li and Zheng 1991la [HEB] Salicarus bimaculatus Zheng and H.-Y. Li 1991) [(O:Di2 SE] Sejanus hongkong Schuh 1984 [O.D.: H] Sejanus neofunereus Schuh 1984 [O.D.: H] Sejanus potanini (Reuter) 1906. Kerzhner 1987 (transfer from Sthenarus),; Kerzhner 1988 [SC] *Sthenaridea rufescens (Poppius) 1915. Schuh 1984 [T] Sthenaridea piceoniger (Motschulsky) 1863. Schuh 1984 [F, H, T] Sthenaropsis gobicus Putshkov 1977. Zheng [unpub.] [GS, IM, N] Genus Strictotergum Zou 1983: 283, 287 [O.D., type sp.: Strictotergum castaneus Zou 1983] Gunior synonym of Pilophorus Hahn; synonymized by Schuh 1989) Tuponia (Chlorotuponia) albescens Zheng and H.-Y. Li 1992 [O.D.: XJ] Tuponia (Chlorotuponia) chinensis Zheng and H.-Y. Li 1992 [O.D.: HEB, SD] Tuponia (Tuponia) elegans (Jakovlev) 1867. Zheng and H.-Y. Li 1992 [N] Tuponia (Tuponia) elegantulus Zheng and HEY 111992 [O:D:: XJ] Tuponia (Tuponia) hsiaoi Zheng and H.-Y. Li 1992 (= Tuponia tamaricicola Hsiao 1963 (preocc.)). Hsiao and Meng 1963 [O.D. of T. tamaricicola: HEB, SD]; Dra- polyuk 1980 (doubtful synonymy with T. arcufera Reuter 1879); Zheng and H.-Y. Li 1992 (new name for Tuponia tamari- cicola Hsiao; sp. distinct.) [SD, HEB] Tuponia (Tuponia) roseipennis Reuter 1879. Zheng and H.-Y. Li 1992 [XJ] Tuponia unicolor (Scott) 1872. 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Zheng, L.-Y. and H.-Y. Li. 1990. Four new species of Psallus Fieb. from China (Insecta, Hemiptera, Heteroptera: Miridae). Reichenbachia 28: 15-19. . 1991. Two new species of Phylinae from Chi- na (Insecta, Heteroptera: Miridae). Reichenbachia 28: 113-115. 1992. Genus Tuponia Reuter from China (Insecta, Heteroptera: Miridae). Reichenbachia 29: 9-13. Zheng, L.-Y. and X.-Z. Li. 1986. Notes on genus Polymerus from China (Hemiptera: Miridae). Acta Scientiarum Naturalium Universitatis Nankaien- sis 1986 (2): 45-52. (!) 1989. An annotated check-list of the genus Adelphocoris Reuter from China (Hemiptera: Mir- idae). Acta Scientiarum Naturalium Universitatis Nankaiensis 1989(3): 77-88. (!) 1990. Three new species of genus Adelpho- coris Reuter from China (Insecta, Heteroptera: Miridae). Reichenbachia 27: 97-100. 1992. Hemiptera: Miridae, pp. 203-206. Jn Huang, F. S., ed., Insects of Wuling Mountains Area, Southwestern China. Science Press, Beijing. (!) Zheng, L.-Y. and L.-J. Liang. 1991. Mirid bugs prey- 473 ing on persimmon leafhoppers from China. Acta Scientiarum Naturalium Universitatis Nankaien- sis 1991(3): 84-87. (!) Zheng L.-Y. and G.-Q. Liu. 1992. Hemiptera: Mir- idae, pp. 290-305. In Peng, J. W., ed., Iconogra- phy of Forest Insects from Hunan. Hunan People’s Press, Changsha, China. (!) 1993. Hemiptera: Miridae, pp. 175-179. In Huang, C. M., ed., Animals of Longqu Mountain. Chinese Forestry Press, Beijing (!) Zheng, L.-Y. and X.-J. Wang. 1982. New species of Lygus bugs from China (Hemiptera: Miridae). En- tomotaxonomia 5(1): 47-59. (!) . 1983. New species and new records of Lygus (subg. Apolygus) from China (Hemiptera: Miri- dae). Acta Zootaxonomica Sinica 8(4): 422-433. (!) Zheng, L.-Y. and C. Yu. 1990. A new genus of the Lygus complex from China, with descriptions of six new species (Hemiptera: Miridae). Entomo- logica Scandinavica 21: 159-171. 1992. Notes on Chinese species of Lygus (s. str.) Hahn with descriptions of three new species (Hemiptera: Miridae). Acta Zootaxonomica Sin- ica 17(3): 352-359. (!) Zou, H.-G. 1983. A new genus and three new species of Pilophorini Reuter from China (Hemiptera: Miridae). Acta Zootaxonomica Sinica 8(3): 283- 287. (!) 1985. Three new specie of Halticus Hahn from China (Hemiptera: Miridae). Acta Zootax- onomica Sinica 10(3): 304-308. (!) 1986. Three new species of Dryophilocoris Reuter from China (Hemiptera: Miridae). Ento- motaxonomia 8(3): 209-214. (!) 1987a. Descriptions of two new species of Zanchius from China (Hemiptera: Miridae). Acta Zootaxonomica Sinica 12(3): 297-300. (!) . 1987b. Two new species of Pseudoloxops Kirk. from China (Hemiptera: Miridae). Acta Zootax- onomica Sinica 12(4): 289-391. (!) . 1987c. Anew species of Pilophorus from Chi- na (Hemiptera: Miridae). Entomotaxonomia 9(2): 107-108. (!) 1989. New species and new records of Mir- idae from China. Acta Zootaxonomica Sinica 1 4(3): 327-332. (!) PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 474-475 REPORTS OF OFFICERS MEMBERSHIP REPORT List oF NEw MEMBERS FoR 1994 George J. Balogh Matthew D. Moran Alessandra Rung de Paula Baptista John K. Moulton Nell Benton Dawn Southard Michael J. Firko David L. Wagner R. Wills Flowers : ie here Letters were sent to each of the applicants after his/her name was read at a regular meeting. Several letters were sent in re- sponse to requests for membership appli- cations. All members are urged to invite their colleagues and students to become members of the Society. Phillip A. Furr David G. Furth Rodney S. Hanley Lynn S. Kimsey Jeff B. Knight Peter W. Kovarik Jon A. Lewis Andrea Lucci Charles D. Michener Ralph P. Eckerlin, Membership Chairman SUMMARY FINANCIAL STATEMENT FOR 1994 Special General Publications Total Fund Fund Assets Assets: November 1, 1993 $10,239.78 $114,999.53 $125,239.31 Total Receipts for 1994 62;382.32 4,574.04 66,956.36 Total Disbursements for 1994 59,926.59— 840.00- 60,766.59— Assets: October 31, 1994 12,695.51 ITS. 733257 131,429.08 Net Changes in Funds $ 2,455.73 $ 3,734.04 $ 6,189.77 Audited by the Auditing Committee, November 23, 1994, consisting of Natalia J. Van- denberg, Warren E. Steiner, Jr., and Michael E. Schauff, Chairman. Presented to the membership at the meeting of December 1, 1994. Respectfully submitted, Norman E. Woodley, 7reasurer VOLUME 97, NUMBER 2 EDITOR’S REPORT Seventy-eight articles, eight notes, six book reviews, and two obituaries were pub- lished in 1994 for a total of 779 pages. This is an increase of nine articles and 119 pages over 1993. Page charges for book reviews, obituaries, and approximately three articles (up to 15 pages each) per issue were defrayed by the Society. I thank Gary L. Miller for his efforts in obtaining book reviews and A. G. Wheeler, Jr. for the excellent copy editing of two memoirs now in production. F. Christian Thompson continued his laborious efforts 475 to format and code these two nearly com- pleted memoirs scheduled to go to the print- er in early 1995. I also give special thanks to Marie Blair for her continued assistance in routing manuscripts and aiding with cor- respondence. As usual, the Society is indebted to the many specialists who served as reviewers for the more than 86 manuscripts received in 1994. Their services are critical to main- taining the high quality of papers published in the Proceedings. Thomas J. Henry, Editor PROC. ENTOMOL. SOC. WASH. 97(2), 1995, pp. 476-480 SocIETY MEETINGS 994th Regular Meeting— May 5, 1994 The 994th Regular Meeting of the Ento- mological Society of Washington was called to order by President Paul J. Spangler at the Log Lodge, Beltsville, Maryland, at 8:10 pm on May 5, 1994. Thirteen members and six visitors were present. Minutes of the April meeting were read by the Recording Sec- retary M. Alma Solis and approved as read. President Spangler called for reports from officers. Ralph Eckerlin, Membership Chairman, reported four new members: R. Wills Flowers, Agricultural Research Pro- grams, Florida A. & M. University, Talla- hassee, Florida; Jon A. Lewis, Systematic Entomology Laboratory, Washington, D.C.; Charles D. Michener, Snow Entomological Museum, University of Kansas, Lawrence, Kansas; Matthew D. Moran, Department of Biology, University of Delaware, Newark, Delaware. President Spangler called for announce- ments, notes, or exhibits. John H. Fales re- ported finding for the first time in Calvert County, Maryland, the following butterflies: Erynnis brizo (May 15, 25 at Lusby), Hes- peria metae (June 16 at Plum Point), and Satyrium liparops strigosum (June 22 at St. Leonard). Also, in 1993 the Monarch but- terfly recovered from the poor populations in 1992. Monarch butterflies migrating northward in 1994 were seen on April 18, 21, 26. He also distributed copies of a list of “Butterflies known from the Western Shore of Southern Maryland northward to the Largo area in Prince Georges County and the South River in Anne Arundel Coun- tye. Nathan Schiff, Program Chairman, intro- duced the speaker for the evening, Dr. Suz- anne Batra, Bee Research Laboratory, ARS, USDA, whose talk was entitled “A Buzz About Fuzz.’ Many bees are utilized as pol- linators, for example, bumblebees, horn- faced bees, Anthophora sp., Osmia sp., and Colletes sp. Honeybees remain the most im- portant pollinators of crops. They are fast, show a high degree of fidelity, fly long range, and carry much pollen, mainly through the use of their hairy bodies. There are many different kinds of hairs: branched or plu- mose, stiff, sturdy, straight setae, twisted hairs (spirals), barbed, flat-tipped, rough and straight, and flattened, leaflike or spatulate. Bees use these hairs for a wide variety of important functions. They collect pollen, as a lump of nectar-moistened pollen on the corbicula, held in place by setae, a mass of loose pollen on the hind legs, held in place by plumose hair, or loose pollen held be- neath the abdomen by spiralled, flat-tipped, or rough hairs. In courtship and mating the males of Anthophoridae and Megachilidae may have modified legs with elaborate brushes of specialized hairs, which they brush over the female’s face, antennae, and eyes; they hold onto her with stiff setae in- side their other legs. They collect and dis- tribute pheromones with velvety patches of flattened hair, usually on the feet or legs. Other specialized setae on various parts of the body collect nectar or floral oils, spe- cialized flattened hairs lay down waterproof Dufour’s gland secretions for nest construc- tion, and one genus uses “‘foot-pats” for communication. Our visitors were introduced and the meeting was adjourned at 10:45 pm. After the meeting refreshments were provided by John Neal and an anonymous individual. M. Alma Solis, Recording Secretary 995th Regular Meeting—June 2, 1994 The 995th Regular Meeting was held at a banquet with the Maryland Entomological Society at the Associates Court at the Na- tional Museum of Natural History, 10th & VOLUME 97, NUMBER 2 Constitution, Washington, D.C. Jeffrey R. Aldrich was Master of Ceremonies. The speaker for the evening was Professor Lin- coln P. Brower whose presentation was en- titled “The Magnificent Migration of the Monarch Butterfly.”” Over 150 people at- tended and the banquet was adjourned at 9:45 pm. M. Alma Solis, Recording Secretary 996th Regular Meeting— October 6, 1994 The 996th Regular Meeting of the Ento- mological Society of Washington was called to order by President Paul J. Spangler in the Waldo Schmitt Room, National Museum of Natural History at 8:10 pm on October 6, 1994. Thirteen members and three visi- tors were present. Minutes of the May meet- ing were read by Don Anderson and ap- proved as read. President Spangler called for reports from officers. Membership Chairman, Ralph P. Eckerlin, read the names of the following applicants for membership: Alessandra Rung de Paula Baptista, Departamento de Zoologia, Laboratorio de Entomologia, Universidad Federal do Rio de Janeiro, Brazil; George J. Balogh, Portage, Michi- gan; Nell Benton, Alexandria, Virginia; J. H. Frank, Entomology and Nematology De- partment, University of Florida, Gaines- ville, Florida; Phillip A. Furr, Albemarle, North Carolina; Michael J. Firko, Colum- bia, Maryland; David G. Furth, Depart- ment of Entomology, Smithsonian Institu- tion, Washington, D.C.; Lynn S. Kimsey, Department of Entomology, University of California, Davis, California; Jeff B. Knight, Nevada Division of Agriculture, Reno, Ne- vada. Two new members, David Furth and Michael Firko, were present. President Spangler called for notes or ex- hibits. Nathan Schiff brought a hepialid cat- erpillar, Korscheltellus gracilis (Grote), he found underneath moss mats from Spruce Knob, West Virginia, and a sawfly, Ato- mocera decepta Rohwer, he found on Hi- 477 biscus moscheutus and asked if anyone knew of any more specimens for study. John Neal brought up the 1000th Regular Meeting, Dave Furth suggested an archival meeting, and Russell Stewart suggested a buffet. Wal- ter Sheppard and John Heraty volunteered to be on an ad hoc committee for the 1000th Regular Meeting. President Spangler asked for volunteers for refreshments: J. Neal (November), R. Eckerlin (December), M. Firko (May). Nathan Schiff, Program Chairman, intro- duced the speaker for the evening, Dr. Wal- ter S. Sheppard, whose talk was entitled “Genetic Diversity in the Honey Bee.” Dr. Sheppard described the biogeography and biology of Apis species around the world with beautiful photos of honey bees. He il- lustrated unusual nesting habits worldwide, such as giant honey bees out in the open in trees and water towers and dwarf honey bee honey combs for sale in a Bangkok market. Although the original distribution of Apis mellifera L. was from Scandinavia to Africa with 25 subspecies, the species has been in- troduced worldwide and in some areas dis- placing Apis cerana. This species, also known as the eastern cavity nesting bee, was the most important bee for honey production in India and Asia until A. mellifera was in- troduced. A. mellifera lamarckii in modern Egypt are kept in mud tubes just as in an- cient Egypt. The tubes were kept on barges and the honey sold on the Nile. Bee parts found in a 3000 year old piece of wax found alongside mummified Egyptians in the Egyptian Museum Torino were collected for DNA sequencing to compare with the mod- ern /amarckii group. Dr. Sheppard also dis- cussed the relationships between the species and subspecies using DNA sequencing re- sults. Our visitors were introduced and the meeting was adjourned by President Span- gler at 9:30 pm. After the meeting refresh- ments were provided by contributors who wish to remain anonymous. M. Alma Solis, Recording Secretary 478 997th Regular ‘\ieeting—November 3, 1994 The 997: meeting of the Entomological Society of Washington was called to order by Dr. Paul J. Spangler, President, in the Waldo Schmitt Room of the Natural His- tory Building at 8:10 pm on November 3, 1994. Twelve members and six guests were present. Minutes of the October meeting were read by Ms. Hollis Williams and ap- proved as read. President Spangler called for reports from officers. Dr. Ralph P. Eckerlin, Membership Chairman, reported no new members. The Nominating Committee consisting of Dr. E. Eric Grissell (Chairman), Dr. Donald M. Anderson, and Dr. Donald R. Davis pro- vided a slate of officers for the following year: Dr. Ralph P. Eckerlin, President-Elect; Ms. Darlene D. Judd, Recording Secretary; Ms. Hollis B. Williams, Corresponding Sec- retary; Mr. James Pakaluk, Custodian; Dr. Norman E. Woodley, Treasurer; Dr. Na- than M. Schiff, Program Chairman; Dr. M. Alma Solis, Membership Chairman; Mr. Thomas J. Henry, Editor; Dr. David Smith, Associate Editor. The slate was read by Dr. Anderson. Additional nominations will be accepted and the entire slate voted on at the Annual Meeting on December 1. Dr. John Heraty, who is a member of the Ad Hoc Committee for the 1000th Regular Meeting with Dr. Steve Sheppard, made two sug- gestions. The committee suggested either a restaurant or a wine and cheese reception in the museum or elsewhere. Funds for a speaker are available and the society needs suggestions. President Spangler called for notes and specimens. Dr. Raymond J. Gagné showed some slides of mass larval migration of black fungus gnat larvae (Diptera: Sciaridae). This phenomenon may be more common than indicated by the few references in the sci- entific literature. The migrations are ephemeral and seeing one depends mainly on luck. During his 30 years’ experience in Washington, Dr. Gagné has had several calls about long snakelike streams of worms PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON crawling through a backyard, but always long after the fact, and the observers had taken no specimens or photographs. On Septem- ber 29, 1994, in Silver Spring, Maryland, Ms. Louise Rickard, an amateur naturalist, observed two rings of migrating sciarids on the front walk of her residence, watched their progress, took photographs, and saved spec- imens in alcohol. The larger ring was about 60 cm in circumference, about 1.2 cm wide, and 0.6 cm high. When she disturbed a ring, the larvae reformed it, but near nightfall the larvae in one ring that was left undisturbed finally crawled off into the adjacent lawn and disappeared. Steffan (1966, Univ. Calif. Pubs. Entomol. 44: 1-77) reviewed sciarid mass migration reports. Two reports are particularly good, one by Beebe (1949, High Jungle: 261-269) and one by Becker (1914, Psyche 21: 94-95). Both reported that movement of the mass was jerky and ef- fected mainly by the larvae at the top crawl- ing forward over those on the bottom. Beebe noticed that his larvae were of several sizes, evidently belonging to several instars, in- dicating that their dispersal is related to the larvae having depleted a food source. The present case 1s especially noteworthy be- cause Ms. Rickard noticed and saved a larg- er, differently shaped larva crawling along with the sciarids. It was Muscina stabulans (Fallén) (Diptera: Muscidae), a carnivorous maggot that preys heavily on many kinds of gregarious larvae in such media as dung and mushrooms. This is the first record of another species associated with mass mi- gration of sciarids. Dr. Schiff, Program Chairman, intro- duced the speaker for the evening, Dr. Bruce McPheron of Pennsylvania State Univer- sity, whose talk was entitled “Ode to St. Bush or Why Are There So Many Kinds of Tephritid Flies.”” Dr. McPheron studies the evolution of fruit fly diversity and is inter- ested in how it reflects insect diversity in general. While most fruit flies are associated with fruits, as their name indicates, species in this family exhibit a variety of habits: several hundred species are leaf miners, for VOLUME 97, NUMBER 2 example, on parsnip; a whole subfamily is associated with flower heads and seeds of Compositae; Blepharoneura is associated with Cucurbitaceae where partitioning of resources has been shown. Specifically Dr. McPheron is studying the relationship be- tween host plant use and the number of fruit fly species. A single species studied exten- sively, such as the Mediterranean fruit fly, has a diversity of hosts such as coffee, hot peppers, and tropical almond, but shows no evidence for substructuring of host plant choice. But where Anastrepha is introduced, its distribution 1s tied in space and host plant use to the presence of native fruit fly species. In northern Brazil it infests only one plant species, but in southern Brazil where other Anastrepha species occur it is not found on the same host plant as in northern Brazil. Although this is ecologically interesting, there is no evidence that host plants are a driving factor in the population differenti- ation between the fruit fly species. In 1865, Benjamin Walsh speculated that in Rhag- oletis pomonella (Walsh), there was a link- age between use of hosts and population divergence. One hundred years later, in 1965, Guy Bush published his revision of the North American genus Rhagoletis. He compared morphological evolution with host plant use and identified a correlation between morphological species groups and host plant use, suggesting a host plant shift within the natural distribution of a species, the occurrence of reproductive isolation, and sympatric evolution. Bush listed character- istics of organisms having undergone sym- patric evolution: close association with the host plant; mating on the host plant; larva spending its entire life cycle within one fruit and tied into the production of fruit. Dr. McPheron studied R. pomonella on two major hosts with overlapping distributions, hawthorns and apples. He collected fruits, reared the larvae to adults, conducted ge- netic analyses with allozyme electrophore- sis. He found that R. pomonella populations feeding on the different host plants were ge- netically different. 479 Our visitors were introduced and the meeting was adjourned at 9:35 pm. After the meeting refreshments were provided by Dr. John W. Neal, Jr., President-Elect. M. Alma Solis, Recording Secretary 998th Annual Meeting—December 1, 1994 The 998th Annual Meeting of the Ento- mological Society of Washington was called to order by President Paul J. Spangler in the Naturalists’ Center of the Natural History Building at 8:00 pm on December 1, 1994. Fifteen members and four guests were pres- ent. Minutes of the November meeting were read by Recording Secretary Alma Solis and approved as read. President Spangler called for officers’ re- ports. Membership Chairman, Ralph D. Eckerlin, reported no new members. He re- ported a total of 19 new members for 1994 and encouraged more new members. The report by the Editor, Thomas Henry, was read by M. Alma Solis. Seventy-eight arti- cles were published for a total of 779 pages. T. Henry also thanked Gary L. Miller, Book Review Editor, A. G. Wheeler, Jr., Special Publications Editor, and F. C. Thompson for their service to the society. He thanked the reviewers of the journal articles and Ma- rie Blair for assistance. The report by the Custodian, Jim Pakaluk, was read by Paul Spangler. He thanked Sarah Donahue and Tami Carlow who helped with mailings and record keeping. The report by the Corre- sponding Secretary, Hollis Williams, was read by Paul Spangler. She reported writing fourteen letters to new members, speakers, and contributors on behalf of the society. M. Alma Solis read the financial report by Treasurer Norman Woodley that showed the society to be solvent. The report was examined and approved by the Audit Com- mittee, Michael E. Schauff, Chairman, War- ren E. Steiner, Jr., and Natalia J. Vanden- berg, Members. Outgoing President Span- gler thanked Don Anderson for his help, John Neal for organizing the banquet, Tom Henry as Editor of the journal, the officers, 480 and the members of the Auditing, Nomi- nating, and Ad Hoc 1000th Meeting Com- mittees. The slaic of officers for 1995 by the Nom- inating (ommittee was presented by Dave Smith President—John W. Neal, Jr. President-Elect— Ralph Eckerlin Recording Secretary — Darlene Judd Corresponding Secretary— Hollis B. Wil- liams Treasurer— Norman E. Woodley Program Chairman—Nathan M. Schiff Membership Chairman—M. Alma Solis Custodian— James Pakaluk Editor—Thomas J. Henry The motion to accept the slate was made by Ted Spilman, seconded by John Heraty, and was voted upon and unanimously ac- cepted by the members present. President Spangler called for notes and specimens. Nathan Schiff reported that the meetings will be held at the Naturalists’ Center until April, possibly May. Ted Spil- man brought a book entitled The Beetles of Northeastern North America, Volume I: In- troduction; Suborders Archostemata and Adephaga, by N. M. Downie and Ross H. Arnett, Jr. (1994, The Sandhill Crane Press, Gainesville, Florida, $160.00 for 3 vol- umes). Nathan Schiff brought specimens of male and female acrocerid flies, or small- headed flies. The female lays thousands of eggs, and the larvae crawl up the legs of spiders and live as ectoparasites on the book lungs. Ralph Eckerlin brought in an article in the Metro section of the Washington Post describing the removal of a cockroach from the ear of a student from George Washing- ton University. William Bickley brought to the attention of the society that Manya Stoetzel, a member of this society, has been elected President-Elect of the Entomologi- cal Society of America. Curtis Sabrosky re- ported the death in Brazil of José Carvalho, a specialist in Miridae (Heteroptera). Program Chairman, Nathan Schiff, intro- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON duced the speaker for the evening, Dr. L. P. S. (Bas) Kuenen, USDA, ARS Bee Research Laboratory, whose talk was entitled “Flights of Fancy? Factors Mediating Male Moths’ Flight Toward a Pheromone Source.” Dr. Kuenen described his work on factors af- fecting the movement of male moths toward a pheromone source. A video produced at Cardé’s laboratory showed the structure and function of a wind tunnel, a sustained flight tunnel that allows the controlled study of flight by insects. The male moths regulate their velocity by visually looking at the moving floor pattern, and can be slowed down if necessary by regulating the speed of the floor pattern. The male moths fly up- wind toward a pheromone source. If the pheromone plume is ended abruptly in the wind tunnel, the male continues flying to- ward the pheromone source, but exhibits casting behavior. As the male moth flies upwind it doesn’t move up or down and just moves in the same plane with a char- acteristic side to side movement. This work was done with the gypsy moth in the wind tunnel, but studies in a forest environment were also done. In nature casting behavior is also apparent when the male leaves the pheromone plume, but it is more complex due to the lateral shifts in the wind. He has also investigated pupal size as a factor af- fecting flight and found that smaller males fly faster or appear to fly faster. He has also tested the hypothesis that velocity equals distance over time. In locusts, studies had shown that they fly faster at higher altitudes. In moths, he found that they fly slower at higher altitudes which is believed to be be- cause of other visual cues. Our visitors were introduced, new mem- bers present were introduced, and President Paul Spangler transferred the gavel to Pres- ident-Elect John W. Neal, Jr. John Neal ad- journed the meeting at 9:25 pm and refresh- ments were provided by Ralph Eckerlin. M. Alma Solis, Recording Secretary PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS ‘Cynipid Galls of the Eastern United States, by Lewis H. Weld. 124 pp. 1959 _o $ 5.00 Cymipia: Galls-ot the southwest, by dewis Hi Weld? 35 pp: 1960. 2 i a re 3.00 BS OLOMAerS OMIGY I pid sali Sls oo Sha wh kU Fe eo aN eee Poe SO LO a 6.00 Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. A Uae fy PASS Ty bY yt (NS at ISM DSS es uA EE CRT SPs SAA a Ae et 2 Sane Ds ee OY BN RIA ee 1 mst Ay SED 1.00 A Short History of the Entomological Society of Washington, by Ashley B. Gurney. 15 pp. 1976 _ 1.00 Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. SO SY As gh Le DE gel RL NR AG ae ANE OF IN! NILES Ne ES De A eT AT Ae a 1.50 Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. BaP VS RAN HIGHLY Sek Uo 1 Sy ten Mota aac an rk hn ae OE IN ae NL SS NR ae tS 2.00 A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael |edp'ysfelg UUM i Pree) 6) 6] yo ICS) O pantie Rr Ae ak 2D gente s NANeTnen ne Ara Ol sci ELE L.A AD NSB arc Se PARR STN WA 10.00 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 $15.00 No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving. (out of SIS! 8) Bb dS Ab ea Davies OE pital ALE Ming aE ilo ee eR RI ecg Can Rhea Pe Pak Be Peet eA print) No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. yi HRTEM GL a hae eRe ea eg LE aie ae ee er 15.00 No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952. _.- 15.00 No.5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957. _. 15.00 No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi akahasits230ippyy VOGOL Le se 50 Ny val id he eke ENG Pe SENG Peed ek Oe ap ee ak eee Pie 15.00 No.7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. ESTA Sloss a NEON la RN 8) LEER RE Pe ee Od ADRES UR aR ASAE OR NIN ae 11.00 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 _w-2-- Peed 12.00 No.9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. PEOVER NO IS Nae Lat OAL NE AAPA IES a ted RO aac a Na a ERO OY Ls ka WUE AE, Mee eBay HAL 10.00 No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W. Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982 0. 11.00 No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 (0) BLSed Us fol pes oe PLR Os 2 a ta sew On sen ER OMS ICR Ata) Pea NGM AN RETNA ELMAN HAs Paste 18.00 No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. NTS OY 0 PNAS! eh Bh EAM Sa SRN ea Oa can EON DEUS Sy Nees. 8 WR ak be APES a yD pn neo oe 5.00 No. 13. An Identification Manual for the North American Genera of the Family Braconidae (Hyme- noptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987... 18.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $25.00 per volume to non-members and $13.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10 per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Custodian, Entomological Society of Washington, c/o Department of Entomology, MRC 168, Smithsonian Institution, Washington, D.C. 20560. CONTENTS (Continued from front cover) SCHUH, RANDALL T., PER LINDSKOG, and I. M. KERZHNER—Europiella Reuter (Het- eroptera: Miridae): Recognition as a Holarctic group, notes on synonymy, and description of a new species, Europiella carvalhoi, from North America SCUDDER, G. G. E.—The first record for Bothynotus pilosus (Boheman) (Hemiptera: Miridae) in the Nearctic Region SCHWARTZ, MICHAEL D.—Metasequoiamiris carvalhoi, a new genus and species of conifer- inhabiting Mirini from China (Heteroptera: Miridae: Mirinae) SLATER, JAMES A.—New genera and species of Rhyparochrominae from West Africa (He- miptera; Lygaeidae) described in honor of J. C. M. Carvalho STONEDAHL, GARY M. and DAMIR KOVAC—Carvalhofulvius gigantochloae, a new genus and species of bamboo-inhabiting Fulviini from West Malaysia (Heteroptera: Miridae: Cylapinae) WHEELER, A. G., JR.—Plant bugs (Heteroptera: Miridae) of Phlox subulata and other narrow- leaved phloxes in eastern United States YASUNAGA, TOMOHIDE—A new genus of mirine plant bug, Carvalhopantilius, with two new species from Taiwan (Heteroptera, Miridae) ZHENG, LE-YI—A list of the Miridae (Heteroptera) recorded from China since J. C. M. Car- valho’s *‘World Catalogue”’ MISCELLANEOUS REPORTS OF OFFICERS SOCIETY MEETINGS a ee otis, Mae cae = Se ee ee ee . Oh. ae VOL.-97 JULY 1995 NO. 3 | (ISSN 0013-8797) \- YG | = 7X PROCEEDINGS A LT. ‘Cj 1999 of the ENTOMOLOGICAL SOCIETY otf WASHINGTON PUBLISHED QUARTERLY CONTENTS AALBU, ROLF L., THEODORE J. SPILMAN, and KIRBY W. BROWN—The systematic status of Amblycyphrus asperatus, Threnus niger, Pycnomorpha californica, Emmenastus rugosus, and Biomorphus tuberculatus Motschulsky (Coleoptera: Tenebrionidae) ........... 481 ADAMSKI, DAVID—Review of the Blastobasidae of the Republic of The Seychelles (Lepi- Moptenay GelechiOidea) sree ieme niet sa in Uk a Ney te ITU TAURINE ae WoC col aa ae Ha ee Acta OM hae EERE 489 BOHART, RICHARD M.—A review of New World Entomognathus with descriptions of seven Hew species (ia vimenoprera.,Spoecidae, Crabronini) | 3/0085. ce ee uke ae eee ioneen sek canta 500 BYERS, GEORGE W.—New species of Nearctic snow crane flies of the genus Chionea (Dip- LET AM MPU GAG > anne te excise en crs io Sones churns, MSM URL eu Ue wae TE TS Wen NMR SR Uae 508 FROESCHNER, RICHARD C.—Nicholas A. Kormilev: A list of his entomological publications SUNCE UF Cia yg ofS LEED oP RMA SOME AI SA Rots MELA RA aS AWA ne RN Wier 515 GOEDEN, RICHARD D., DAVID H. HEADRICK, and JEFFREY A. TEERINK—Life history and description of immature stages of Valentibulla californica (Coquillett) (Diptera: Te- phritidae) on Chrysothamnus nauseosus (Pallas) Britton in southern California ............. 548 GRIMALDI, DAVID A. and DALTON DE SOUZA AMORIM—A basal new species of Ol- biogaster (Diptera: Anisopodidae) in Dominican amber, and its systematic placement ..... 561 HEYDON, S. L.—The North American species of Systasis Walker (Hymenoptera: Pteromali- HU, G. Y. and J. H. FRANK—Structural comparison of the chorion surface of five Philonthus SPECIES (MC OlSOplel ad MUADUYUMIGae iii Gi asians en aay REIS sha MC Tay ae Cali ea uO ein untae bg 582 KIMSEY, LYNN S.—New amisegine wasps from southeast Asia (Hymenoptera: Chrysididae) 590 KONDRATIEFF, BORIS C. and CHARLES H. NELSON—A review of the genus Remenus Ricker (Plecoptera: Perlodidae), with the description of two new species ..................-. 596 LANDRY, JEAN-FRANCOIS and DAVID L. WAGNER—Taxonomic review of apple-feeding species of Phyllonorycter Hiibner (Lepidoptera, Gracillariidae) in North America .......... (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1995 JOHN W. NEAL, JR., President NORMAN E. WOODLEY, Treasurer RALPH P. ECKERLIN, President-Elect NATHAN SCHIFF, Program Chairman DARLENE D. Jupp, Recording Secretary M. ALMA SOLIS, Membership Chairman Hous B. WILLIAMS, Corresponding Secretary PAUL J. SPANGLER, Past President JAMES PAKALUK, Custodian THOMAS J. HENRY, Editor DAVID R. SMITH, Associate Editor Gary L. MILLER, Book Review Editor Publications Committee DONALD R. DAVIS TerrRY L. ERWIN F. CHRISTIAN THOMPSON A. G. WHEELER, JR., Special Publications Editor Honorary President Curtis W. SABROSKY Honorary Members LOUISE M. RUSSELL ALAN STONE KARL V. KROMBEIN All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, MRC-168, Smithsonian Insti- tution, Washington, D.C. 20560. 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 8 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of See 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, MRC-168, Smith- sonian Institution, Washington, D.C. 20560. Members in good standing receive the Proceedings of the Ento- mological 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 P. 218 OF THE JANUARY, 1991 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. Editor: Thomas J. Henry, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- West, Beltsville, MD 20705. Special Publications: A. G. Wheeler, Jr., Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, PA 17110-9408 Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 19 September 1995 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 481-488 THE SYSTEMATIC STATUS OF AMBLYCYPHRUS ASPERATUS, THRENUS NIGER, PYCNOMORPHA CALIFORNICA, EMMENASTUS RUGOSUS, AND BIOMORPHUS TUBERCULATUS MOTSCHULSKY (COLEOPTERA: TENEBRIONIDAE) Ro.F L. AALBU, THEODORE J. SPILMAN, AND KirBY W. BROWN (RLA) Essig Museum of Entomology, 211 Wellman Hall, University of California, Berkeley, California 94720; (TJS) 4903 Jamestown Road, Bethesda, Maryland 20816; and (KWB) 4570 E. 3rd. N., Joseph City, Arizona 86030. Abstract.—The systematic status of Amblycyphrus asperatus, Threnus niger, Pycno- morpha californica, Emmenastus rugosus, and Biomorphus tuberculatus Motschulsky (Co- leoptera: Tenebrionidae) is resolved. The following changes are proposed: Amblycyphrus is placed as a junior synonym of Cryptoglossa Solier (Cryptoglossini); Amblycyphrus asperatus Motschulsky is placed as a junior synonym of Cryptoglossa spiculifera pectoralis (Blaisdell) [new combination and status]; Threnus Motschulsky is placed as a synonym of Argoporis Horn (Cerenopini), Threnus niger Motschulsky becomes Argoporis niger niger (Motschulsky) [new combi- nation], Argoporis constanzae constanzae Berry 1s placed as a junior synonym of Argoporis niger niger (Motschulsky), Argoporis constanzae inflata Berry becomes Argoporis niger inflata Berry [new combination]; Emmenastus Motschulsky is placed as a synonym of Oxycara Solier [Tentyriini]; Emmenastus rugosus Motschulsky becomes Oxycara rugosa (Motschulsky) [new combination]; Biomorphus Motschulsky is placed as a synonym of Helops Fabricius; Biomorphus tuberculatus Motschulsky is placed as a junior synonym of Helops attenuatus LeConte (Helopini); Pycnomorpha Motschulsky is retained as a valid genus with Pycnomorpha californica Motschulsky as type and two species added, Pyc- nomorpha gibbicollis (Horn) [new combination, transferred from Trichiasida], and Pyc- nomorpha tumidicollis (Blaisdell) [new combination, transferred from Stenomorpha] (As- idini). Keys are provided to Pycnomorpha and its species. Key Words: Coleoptera, Tenebrionidae, Motschulsky, Amblycyphrus, Threnus, Pycno- morpha, Emmenastus, Biomorphus, classification A number of genera have historically been included in catalogs as belonging to the North American fauna but have remained a mystery to later workers. Five of these are genera described by Motschulsky in the late 19th century. All five species, described by Motschulsky in 1845, 1870, and 1872 were known only from the type specimen. The types were deposited in the Moscow State Museum, then the Imperial University Mu- seum, in Moscow, Russia. These five spec- imens remained unknown for more than a hundred years since their original descrip- tion. Recently, through the kind help of Dr. N. B. Nikitsky of the Moscow State Mu- seum, Russia, these five holotypes were bor- rowed by one of us (RLA) and examined by us, finally solving the mystery of these gen- 482 era. Two occur in California, two occur in Baja California and one is not North Amer- ican. The taxonomic position of these gen- era is discussed below. Amblycyphrus and Threnus While working on a Ph.D. dissertation (Aalbu 1985) concerning the systematics of the tribe Cryptoglossini, it became neces- sary to determine the status of genera and species previously included in catalogs as belonging to this tribe. These included the Motschulsky genera Amblycyphrus and Threnus. Amblycyphrus Motschulsky 1870: 401. Amblycyphrus asperatus Motschulsky 1870: 404. Champion 1895: 46 (catalog); Ge- bien 1910: 121, 1937: 701 (catalog). Threnus Motschulsky 1870: 404. Bradley 1930: 184 (classification) Threnus niger Motschulsky 1870: 406. Ge- bien 1911b: 611, 1937: 701 (catalog); Leng 1920: 224 (catalog); Arnett 1960: 652, 672 (classification, distribution); Papp 1961: 106 (catalog); Blackwelder and Arnett 1975: 73.20 (catalog); Arnett 1985: 347 (list). Motschulsky (1870: 401-404) assigned Amblycyphrus, as a ‘““Melasoma of the tribe of the Akisites,” mentioning that this genus was most closely related to Centrioptera Mannerheim. He distinguished Amblycy- phrus from Centrioptera by the following characters: (1) “retractile labrum, which is often entirely hidden under the clypeus”’; (2) ‘““mandibles, which are more distinctly den- ticulate”’; (3) “less thickened, non-monili- form antennae with a non-transverse sub- apical segment”’; (4) “finely punctate elytral striae with the asperities being more obtuse but more pronounced”; (5) “more parallel prosternum, terminating arrow-shaped and not emarginate”’; (6) “non-bilobed meso- sternum”’; and (7) “more slender legs with longer apical tarsal segment,” all unreliable, non-diagnostic characters. Motschulsky also distinguished this genus from Asbolus LeConte mentioning “the genus Asbolus of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON M. LeConte has a generally shorter form, with the asperities of the elytra tuberculate and much more developed, the truncate, very transverse, apical segment, the visible labrum etc.” Unlike Amblycyphrus, Motschulsky as- signed Threnus, as a ““Melasoma of the tribe of the Centriopterides’’ but compared this genus not to Centrioptera but to Cerenopus LeConte [Tenebrioninae: Cerenopini] (La- cordaire, in 1859 had included Cerenopus in the Centriopterides). Motschulsky distin- guished Threnus from Cerenopus by: (1), antennae distinctly enlarged toward ex- tremity; (2), elytra not enlarged posteriorly; (3), epistoma less produced posteriorly; and (4), dissimilar dentition on inflated parts of femorae and tibiae. Upon examination of these types, several bibliographical problems became apparent. The following observations were made: (1) The specimen labeled “*7Threnus niger Mots., Calif.’ (green label with black hand- written ink) is an Argoporis (Cerenopini), specifically Argoporis constanzae constan- zae Berry 1980, a species which occurs in Baja California Sur. The problem in this case is that the genus Argoporis Horn, was also described in 1870 in Horn’s Revision (p. 325). (2) The other specimen is labeled “‘Cen- trioptera asperata Mots., Calif.,” not Am- blycyphrus asperatus (also green label with black handwritten ink). This specimen is not Centrioptera asperata Horn, as one might suspect, but what Blaisdell (1921: 199) described as Centrioptera dulzurae. The problem in this case is that Centrioptera as- perata Horn was also described in 1870 (p. 279) which makes these homonyms. As the exact date of both publications cannot be positively determined, following the Rules of Zoological Nomenclature, both articles are dated December 31, 1870. Be- cause less disruption of current literature and classification will result, Horn 1870 is selected as having priority over Motschul- sky 1870. As part of the systematic treatment of the VOLUME 97, NUMBER 3 tribe Cryptoglossini, a number of changes in combination and/or status of various genera and species resulted. These changes are discussed in detail in Aalbu (in press). Some of these changes, pertinent to a dis- cussion of Amblycyphrus, are listed here. Amblycyphrus asperatus Motschulsky is a synonym [of Cryptoglossa spiculifera pec- toralis (Blaisdell)] and a secondary hom- onym [of Cryptoglossa asperata asperata (Horn) 1870] and is retained in the Cryp- toglossini. The following changes are proposed: Cryptoglossa Solier 1836: 680 Amblycyphrus Motschulsky 1870: 401 NEW SYNONYMY Cryptoglossa spiculifera pectoralis (Blais- dell) 1921: 198 NEW COMBINATION and NEW STATUS Amblycyphrus asperatus Motschulsky 1870: 404 NEW SYNONYMY Centrioptera dulzurae Blaisdell 1921: 199 NEW SYNONYMY Argoporis Horn 1870: 325 Threnus Motschulsky 1870: 404 NEW SYNONYMY Argoporis niger niger (Motschulsky 1870: 406) NEW COMBINATION Argoporis constanzae constanzae Berry 1980: 35 NEW SYNONYMY Argoporis niger inflata Berry 1980: 36 NEW COMBINATION Emmenastus Emmenastus Motschulsky 1845: 75. Emmenastus rugosus Motschulsky 1845: 76. Lacordaire 1859: 59 (classification); Gemminger and Harold 1870: 1934 (cat- alog); Gebien 1910: 18, 1937: 588 (cat- alog); Leng 1920: 221 (catalog); Black- welder 1945: 512 (Emmenastus) (cata- log); Arnett 1960: 665 (distribution); Papp 1961: 100 (catalog); Blackwelder and Ar- nett 1975: 73.10 (catalog). Emmenastus rugosus Motschulsky has been doubifully recorded from Alaska. Em- menastus was described as a new genus by 483 Motschulsky (1845: 75) with two new spe- cies: compactus (1845: 76) from Kamchatka in eastern Siberia and rugosus (1845: 76) from Sitka in Russian America, now Alas- ka. Because it had not been collected since its original description E. compactus was doubtfully included in the Kamchatka fau- nal list. That problem was solved by Bo- gachev (1968: 889) when he examined three type specimens of EF. compactus in the Zoo- logical Museum of University of Moscow and determined that compactus is actually a species of Oxycara Solier previously known as O. cribrata Wollaston 1867; it occurs not on Kamchatka but on the Cape Verde Islands off western Africa. Even though Motschulsky’s published type lo- cality was Kamchatka, the type specimens bore labels indicating Tenerife. (Bogachev did not find the type of E. rugosus in Mos- cow University or the Zoological Institute.) Soon after Motschulsky described Em- menastus rugosus, Mannerheim (1852: 287, 288, 291, 387) discussed the lack of Mela- somes (Tenebrionidae) in the area of Sitka and their abundance in California, said he did not receive specimens of E. rugosus from Motschulsky, doubted that the species oc- curs at Sitka, and indicated the doubt with question marks in his list of species. Then Mannerheim (1853: 110-112) said he ex- amined Motschulsky’s specimens but doubted that the type specimen of EF. ru- gosus came from Sitka; he placed F. rugosus in Blapstinus near B. pulverulentus Man- nerheim, saying that he had specimens from northern California. LeConte (1866: 106) and Horn (1870: 268, 402) considered Em- menastus rugosus to be unrecognizable. Nevertheless, they along with Champion (1884: 8) applied the generic name Em- menastus to previously described species (by LeConte) and to new species that are at pres- ent in the tribe Eurymetopini. These species were subsequently placed as types of or as species in other new genera by Casey (1907: 287) with only E. rugosus remaining 1n the genus but rejected as part of the American fauna. Blackwelder (1945: 512) placed Em- 484 menastus (attributed in error to Champion) as a synonym of Hylocrinus Casey. This synonymy is not explained, but the first spe- cies listed by Champion (1884: 9) is Em- menastus longulus (LeC.) which was later designated the type of Hylocrinus by Casey (1907: 289). We have studied a specimen of FE. rugosus from the Zoological Museum of the Uni- versity of Moscow. The specimen has 3 small handwritten labels: “type”; ‘““Tenerife ?”; ““Emmenastus rugosus Mots Tenerife?” The first label is on uncoated card, the other two on light blue coated card. The handwriting on these labels is different from that on the two examples of Motschulsky’s labels pub- lished by Korchefsky (1937: pl. 16, fig. 23; pl. 21, fig. 33). We consider the specimen to be the type, even though Motschulsky’s published locality, Sitka, does not appear on its labels. (As mentioned above, Bo- gachev had similar difficulties with labels on the original specimens of EF. compactus.) The type of Emmenastus rugosus differs from E. compactus as described by Boga- chev (1968: 889), in the following ways: Pronotum having lateral margin with a thin, unraised flange; surface moderately convex. Elytra in dorsal view gradually widened from base to half-length, laterally curved gradu- ally and then bluntly triangular to apex; dor- sal surface moderately convex and widely embracing body; epipleura relatively broad, gradually narrowed posteriorly; surface with 4 or 5 irregular longitudinal furrows from which emanate short irregular lateral fur- rows. Prosternum without depression and erect hairs of male. Protibia with spurs long, the longer spur extending to apex of second tarsal article. Motschulsky did not designate a type spe- cies for Emmenastus. The fate of the generic name depends on the designation of a type species and the fate of that species. LeConte (1866: 106) stated that FE. rugosus 1s the type species, and Horn (1870: 268) agreed. Casey (1907: 287) says that E. compactus 1s the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON type species because “‘it was so intended by Motschulsky.”” Casey was wrong; perhaps he thought that Motschulsky’s placement of compactus before rugosus was an intention to designate. Gebien (1937: 588) in his cat- alogue stated that compactus 1s the type spe- cies. Surely LeConte’s designation of 1866 was the first and therefore valid designation. It appears that E. rugosus, like E. com- pactus, should be placed in Oxycara (Ten- tyriini). However, we have not been able to synonymize it with other described species of Oxycara. We make this placement es- pecially because of two distinctive charac- ters on the pro- and mesosternal processes: (1) Prosternal process broad, apex acute, in lateral view horizontal and projecting pos- terior to coxae so as to appear shelf-like; (2) mesosternum between coxae flat, broad, and distinctly lowered from remainder of me- sosternum, anterior half becoming strongly grooved, and finally with distinct notch on anterior border for reception of apex of prosternal process. The following changes are proposed: Oxycara Solier 1835: 254 Emmenastus Motschulsky 1845: 75 NEW SYNONYMY Oxycara rugosa (Motschulsky 1845: 76) NEW COMBINATION Biomorphus Biomorphus Motschoulsky 1872: 38. Brad- ley 1930: 323 (classification) Biomorphus tuberculatus Motschulsky 1872: 40. Gebien 1911la: 463, 1941: 811(666) (catalog); Leng 1920: 236 (catalog); Ar- nett 1960: 688 (distribution); Papp 1961: 130 (catalog); Blackwelder and Arnett 1975: 73.71 (catalog); Arnett 1985: 348 (list). Biomorphus tuberculatus is definitely North American. It has been placed in the Tenebrionini in most catalogs. Of the five under consideration here, its taxonomic po- sition is perhaps the easiest to solve. It clear- VOLUME 97, NUMBER 3 ly falls within the range of variation found in Helops attenuatus LeConte (Helopini). The following changes must be made: Helops Fabricius 1775: 257 Biomorphus Motschulsky 1872: 38 NEW SYNONYMY Helops attenuatus LeConte 1851: 137 Biomorphus tuberculatus Motschulsky 1872: 40 NEW SYNONYMY Pycnomorpha Pycnomorpha Motschulsky 1870: 398. Ge- bien 1910: 126 (synonymy) Pycnomorpha californica Motschulsky 1870: 399. Champion 1895: 251 (catalog); Ge- bien 1910: 127 (catalog); Arnett 1960: 674 (distribution); Papp 1961: 110 (catalog); Blackwelder and Arnett 1975: 73.29 (cat- alog); Arnett 1985: 348 (list). The type specimen of Pycnomorpha cal- ifornica is in relatively poor condition. It was apparently found dead. It is encrusted with dirt, dust and old glue and missing its left mesothoracic and metathoracic legs and its right prothoracic and mesothoracic legs. Additionally, the left prothoracic leg has been reglued on to the specimen but not quite at the right position. The specimen is labeled “‘Pycnomorpha californica Mots., California” and “California,” two green la- bels with black handwritten ink. Motschulsky, in his description of Pyc- nomorpha, compared the genus to Steno- morpha Solier. Based on Solier’s illustra- tion, he distinguished the genus by the transverse labium, obtuse mandibles with pronounced middle tooth, more protruding ligula, prominent pronotum, legs not hairy, and elytra carinate on lateral margin. None of these characters are diagnostic. Fortu- nately, a number of specimens obviously conspecific with the type have been recently collected. This has made possible a more thorough comparison with other genera. From the description alone coupled with the cited locality (Nova-Helvetia, an early 485 name for modern Sacramento) one of us (KWB) assumed that this species was the same as Stenomorpha capitosa (Horn), the only similar species known from the Sac- ramento area. However, examination of the Motschulsky specimen and the recently col- lected conspecific specimens clearly show that the locality cited must be in error and the actual location is central Baja California. Under the present classification of genera in the tribe Asidini, Pycnomorpha must be re- tained as a valid genus. Couplets in the Brown (1971: 28) key to genera of Asidini should be modified as follows: 10(9). | Postgenal process projecting well beyond middle of mentum, often greatly thick- ened or swollen (Fig. 19); mentum in tight, broad contact with postgenal process; pronotum dilated laterally; elytral disc costate (1 sp., central Mexico) Zaleucus Champion 10’. Postgenal process rarely projecting be- yond middle of mentum; if process pro- jects beyond middle of mentum, then el- ytra disc not costate (see couplet 24) .. 11 23(22). Humeri of elytra prominent and reflexed, body length under 16 mm. (see couplet Dil) Ratan ceo etre try Mee ek Oe Asidopsis Casey (part) 23% Humeri of elytra inconspicuous, not re- EXE GM cc een one eR res 24 24(23). Mentum large, often in parallel contact with postgenal process; postgenal process thickened, rarely projecting beyond mid- point of mentum, base opposite gular pedestal abruptly bent, forming a quad- rate open space; pronotum often inflated and gibbose; body glabrous; maxillary palpi weakly sexually dimorphic (3 spp., Baja California) Pycnomorpha Motschulsky 24’. Mentum smaller, not in contact with postgenal process which is usually thin; base of postgenal process opposite gular pedestal gradually bent, not forming a distinct quadrate open space ......... 34 Apical segment of maxillary palpus great- ly enlarged and sexually dimorphic, recti- triangular in the female (Fig. 3), larger and scalene with proximal angle pro- longed in the male (Fig. 2); ligula large and tumid (Fig. 25); body rarely hirsute (85 spp., & sub spps.; SW Canada, W. & Central U.S., Mexico) *Stenomorpha Solier 34(24’). 486 34’. Apical segment of maxillary palpus smaller, only moderately enlarged and often not notably sexually dimorphic, at most recti-triangular in the male (Fig. 24); disc of pronotum and elytra with setae present, body often hirsute; ligula small, not tumid (Fig. 6) (20 spp.; SW. ULS., Mexico) eeenereeee Trichiasida Casey (part) Pycnomorpha shares characteristics with the genera Stenomorpha and Trichiasida. It may be distinguished by the following: It differs from Stenomorpha and Tri- chiasida by the thicker postgenal process, larger and thicker mentum, its lateral edge parallel to the postgenal process; proximal edge of postgenal process abruptly bent op- posite gular pedestal forming a notable quadrate pocket, and by a more horizontal antennal shelf with a deeper and longer groove near the eye. It further differs from Stenomorpha by a more slender antenna, smaller tomentose areas of the tenth antennal segment, and weaker sexual dimorphism of maxillary palpi. It further differs from Trichiasida by the smaller tenth antennal segment, and gla- brous body. Two additional described species from Baja California are clearly congeneric with Pycnomorpha californica. They are Tri- chiasida gibbicollis (Horn) (originally de- scribed in Asida) and Stenomorpha tumi- dicollis Blaisdell. One might note that Horn (1880: 152) proposed Asida gabbi as a re- placement name for Asida gibbicollis which was preoccupied in Asida; Casey (1912: 178) in placing A. gabbi into his new genus 77i- chiasida, restored the name as T. gibbicollis. The following changes are proposed: Pycnomorpha gibbicollis (Horn 1870: 288) NEW COMBINATION Pycnomorpha tumidicollis (Blaisdell 1943: 226) NEW COMBINATION The following key will separate the three species: 1. Lateral edge of pronotum thin, barely re- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON flexed; posterior pronotal gibbae smoothly rounded; posterior pronotal angles obtuse, lat- eral margin of elytra rounded or carinate at basal 4 only; body surface sculpture dull or shiny 1’. Lateral edge of pronotum thick, strongly re- flexed; posterior pronotal gibbae sharply car- inate; posterior pronotal angles acute, lateral margin of elytra entirely carinate; body sur- face sculpture’shiny ........- gibbicollis (Horn) 2. Lateral margin of elytra rounded; body sur- face sculpture shiny tumidicollis (Blaisdell) 2a. Lateral margin of elytra rugosely carinate at basal 4; body surface sculpture dull ne ect ao ae californica Motschulsky A satisfactory number of specimens each species has been examined to verify that the characters are sufficiently constant to justify maintaining the above three distinct spe- cies. Pycnomorpha tumidicollis is from northern to central Baja California, califor- nica from central, and gibbicollis southern. Several undescribed species from Baja Cal- ifornia are on hand. Some species of Sten- omorpha from southern California (River- side and San Diego Counties) may also be- long in Pycnomorpha. Further work on these genera 1S 1n progress. ACKNOWLEDGMENT Special appreciation is extended to Dr. N. B. Nikitsky of the Moscow State Museum, USSR, who kindly sent the types of Ambly- cyphrus, Threnus, Pycnomorpha, Emmen- astus and Biomorphus Motschulsky. LITERATURE CITED Aalbu, R. L. 1985. Systematics and Biology of the Cryptoglossini (Coleoptera: Tenebrionidae). Un- published Ph.D. dissertation, The Ohio State Uni- versity. 329 pp. . (Inpress.) The pimeliine tribe Cryptoglossini: Classification, biology and inferred phylogeny (Coleoptera: Tenebrionidae). Knull Series, Ohio Biological Survey Bulletin. Arnett, R. H., Jr. 1960. The Beetles of the United States. Catholic University Press, Washington, D.C. 1112 pp. . 1985. American Insects—A Handbook of the Insects of America North of Mexico. Van Nos- trand Reinhold, New York. XIII, 850 pp. VOLUME 97, NUMBER 3 Berry, R. L. 1980. A revision of the North American genus Argoporis (Coleoptera: Tenebrionidae: Cer- enopini). Ohio Biological Survey, Knull Series 1: 1-106. Blackwelder, R. E. 1945. Checklist of the Coleopter- ous Insects of Mexico, Central America, the West Indies, and South America. United States Nation- al Museum Bulletin 185, part 3: 343-550. Blackwelder, R. E.and R. H. Arnett, Jr. 1975. Check- list of the Beetles of Canada, United States, Mex- ico, Central America and the West Indies (Red Version). Vol. 1, Pt. 5, Family 73 (Tenebrionidae): 73.1-73.86. Blaisdell, F. E. 1921. New species of Melyridae, Chrysomelidae and Tenebrionidae (Coleoptera) from the Pacific Coast with notes on other species. Stanford University Publications, Biological Sci- ences 1(3): 136-231. 1943. Contributions toward a knowledge of the insect fauna of Lower California. No. 7. Co- leoptera; Tenebrionidae. Proceedings of the Cal- ifornia Academy of Sciences, Series 4, 24: 171- 287. Bogachev, A. V. 1968. The taxonomic position and geographical origin of the darkling beetle Em- menastus compactus Motsch. (Coleoptera, Teneb- rionidae). Entomologicheskoe Obozrenie 47: 889- 891 [in Russian]. Entomological Review 47: 542- 543 [English translation]. Bradley, J.C. 1930. A Manual of the Genera of Bee- tles of America North of Mexico. Daw, Illston and Co., Ithaca, N.Y. 360 pp. Brown, K. W. 1971. Redefinition of the Genera Pe- lecyphorus and Philolithus with a key to the genera of the tribe Asidini (Coleoptera: Tenebrionidae). Coleopterists Bulletin 25(1): 17-30. Casey, T. L. 1907. A revision of the American com- ponents of the tenebrionid subfamily Tentyriinae. Proceedings of the Washington Academy of Sci- ences 9: 275-522. 1912. A revision of the American genera of the tenebrionid tribe Asidini. Memoirs on the Co- leoptera III: 70-214. Champion, G. C. 1884. Biologia Centrali-Ameri- cana, Insecta, Coleoptera (Tenebrionidae). 4(1): 1- 88. . 1895. A list of Tenebrionidae supplementary to the “Munich” catalogue. Mémoiers de la So- ciété Entomologique de Belgique 3: 1-264. Fabricius, J. C. 1775. Systema Entomologiae. Lip- siae. 30 + 832 pp. Gebien, H. 1910. Coleoptorum Catalogus, Pars 15, Tenebrionidae I: 1-166. 191la. Coleoptorum Catalogus, Pars 28, Te- nebrionidae III: 355-583. 487 . 1911b. Coleoptorum Catalogus, Pars 37, Te- nebrionidae IV: 587-742. 1937. Katalog der Tenebrioniden. Teil I. Publicasioni del Museo Entomologico Pietro Ros- si, Duino. 2: 505-883. 1941. Katalog der Tenebrioniden. Teil II. Mitteilungen der Miinchener Entomologischen Gesellschaft 31(2): 803-834 (658-689). Gemminger, M. and B. Harold. 1870. Catalogus Co- leopterum, Vol. VII: Tenebrionidae. Monachii, pp. 1801-2189. Horn, G. H. 1870. Revision of the Tenebrionidae of America, north of Mexico. Transactions of the American Philosophical Society (new series) 14: 253-404, illus. 1880. Contributions to the Coleopterology of the United States, No. 3. Transactions of the American Entomological Society 8: 139-154, pl. 3. . 1894. The Coleoptera of Baja California. Pro- ceedings of the California Academy of Sciences, Series 2, 4: 302-449. Korchefsky, R. 1937. Erklérungen der Tafeln I- XXXVIII. Entomologische Beihefte Berlin-Dah- lem 4: 511-533. (Published in the volume with Horn & Kahle’s large article on collections.) Lacordaire, T. 1859. Histoire Naturelle des Insectes. Genera de Coléoptéres ... Tome 5. Roret, Paris, pp. 1-750. LeConte, J. L. 1851. Description of new species of Coleoptera from California. Annals of the Lyceum of Natural History, New York 5: 125-216. 1866. New species of North American Co- leoptera. Smithsonian Miscellaneous Collections 66(167): 87-177. Leng, C. W. 1920. Catalog of the Coleoptera of Amer- ica. Cosmos Press, Cambridge, Massachusetts. 470 pp. Mannerheim, C. G. 1852. Zweiter Nachtrag zur Ka- fer-Fauna der nord-amerikanischen lander des russischen Reiches. Bulletin de la Société Impér- iale des Naturalistes de Moscou, 25(1): 283-387. 1853. Dritter Nachtrag zur Kafer-Fauna der nord-amerikanischen lander des russischen Reich- es. Bulletin de la Société Impériale des Naturalistes de Moscou 26: 95-273. Motschulsky, V. 1845. Remarques sur la collection de Coléoptéres Russes. Bulletin de la Société Im- périale des Naturalistes de Moscou 18(1): 3-82. . 1870. Enumération des nouvelles espéces de coléoptéres rapportés de ses voyages. Bulletin de la Société Impériale des Naturalistes de Moscou 43(1): 379-407. (10th article). illus. 1872. Enumeération des nouvelles espéces de coléoptéres rapportés de ses voyages. Bulletin de la Société Impériale des Naturalistes de Moscou 45(2): 23-55 (11th article). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 488 1961. Checklist of Tenebrionidae of Solier, A. J. 1836. Essai sur les Collaptérides (suite). Annales de la Société Entomologique de France 5: Papp:..€2 8s: America, north of the Panama canal (Notes on North American Coleoptera, No. 14). Opuscula 635-684. Entomologica 26(1-2): 97-140. Wollaston, T. V. 1867. Coleoptera Hesperidum, Be- Solier, A. J. 1835. Essai sur les Collaptérides (suite) ing an Enumeration of the Coleopterous Insects of the Cape Verde Archipelago. John Van Voorst, Annales de la Société Entomologique de France 4: London. 39 + 285 pp., 1 map. 249-419. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 489-499 REVIEW OF THE BLASTOBASIDAE OF THE REPUBLIC OF THE SEYCHELLES (LEPIDOPTERA: GELECHIOIDEA) DaAvip ADAMSKI 6033 Majors Lane, Apt. #2, Columbia, Maryland 21045. Abstract. — Blastobasis legrandi is described from misidentified specimens collected from The Republic of The Seychelles by Legrand in 1959. Blastobasis acarta Meyrick and B. intrepida Meyrick are redescribed, and a lectotype is designated for the former species. A key, illustrations of adults and of male and female genitalia are provided. Key Words: chelles About sixty islands and islets make up the present Republic of The Seychelles. They are divided into three island types: the gra- nitic islands or Seychelles proper, which form a compact group of mountainous is- lands emerging from a submarine plateau between 4-5 degrees south of the equator and 56 degrees east of Greenwich; the sand cays which are located south-west of the Seychelles proper, but also include isolated islands along the northern and southern edge of the Seychelles plateau; the elevated reefs of the Aldabra group. The granitic islands are situated about 930 km from Madagascar, 1600 km from East Africa and Mauritius, and over 1700 km from India. Aldabra lies 800 km from Mahé of the granitic islands, and is much closer to Madagascar and East Africa. Several expeditions to the Seychelles that included collecting Lepidoptera have oc- curred since the French coleopterist, Charles Alluaud, visited the islands in 1892. The earliest of these expeditions yielded several undescribed Lepidoptera (Lionnet 1984). The first major collections of Lepidoptera from the Seychelles were amassed by The Percy Sladen Trust Expeditions of 1905 and 1908. These British collections represented Lepidoptera, Gelechioidea, Blastobasidae, Blastobasinae, Blastobasis, Sey- 111 species, of which 90 species were con- sidered endemic to the Seychelles (Meyrick 1911). All the microlepidoptera collected from The Percy Sladen Trust Expeditions were studied by E. Meyrick (Lionnet 1984). Another major collection of Lepidoptera of the Seychelles was made by Henry Le- grand of The Muséum National D’Histoire Naturelle, Paris. He visited the Seychelles in 1956 and 1958-60 and collected, more than 3500 specimens, and recognized 363 species, of which 117 were new to science (Legrand 1965). Two subsequent collections of Lepidop- tera of The Seychelles were made by Amer- ican Lepidopterists. In 1968, Jay Shaffer participated in an international collecting expedition to Aldabra Atoll. This expedi- tion was one of several sponsored by The Royal Society of London from 1966 to 1980. In 1986, I participated in a Smithsonian sponsored expedition to the granitic Sey- chelles and to Aldabra. Both expeditions yielded many Lepidoptera, most of which had been recorded previously from the Sey- chelles and from the region. The specimens collected on the latter two expeditions are deposited in the United States National Museum. The purpose of this paper is to 490 review the Blastobasidae of The Republic of the Seychelles Islands. The Methuen Handbook of Colour (Kor- nerup and Wanscher 1978) was used as a color standard for the description of the adult vestiture. Genitalia were dissected as de- scribed by Clarke (1941), except mercuro- chrome and chlorazol black were used as stains. Pinned specimens and genital prep- arations were examined with dissecting and compound microscopes. Wing measure- ments were made using a calibrated ocular micrometer. RESULTS Key to the Blastobasidae of the Republic of the Seychelles Malesiey seep nce irate cai soe RGN eC EE: 2 Rem ales eee eve Seep eases 3 Two apical articles of labial palpus normal (Fig. 4); proximal flange of valva with long hairlike setae, elongate valval spine absent, apical pro- cess of lower part of valva normal, aedeagus angled apically (Fig. 7))....... Blastobasis acarta — Two apical articles of labial palpus widened dorsoventrally (Fig. 5); proximal flange of val- va without long hairlike setae, elongate valval spine present, apical process of lower part of valva small, aedeagus straight (Fig. 8) PO er kT ey ata eee Blastobasis intrepida 3. Membrane surrounding ostial area with dense microtrichia (Fig. 9) ......... Blastobasis acarta — Membrane surrounding ostial area without dense microtrichia (Figs. 10-11) 4. Membrane posterior to seventh tergum with two small crescent-shaped sclerites (Fig. 11) ... Pe ee ao oe nee Blastobasis legrandi — Membrane posterior to seventh tergum with- out such sclerotizations; with a round and wrinkled, small invagination within lateral membrane between sixth and seventh seg- ments: (Figs il0)) see seers: Blastobasis intrepida mle Blastobasis legrandi Adamski, NEw SPECIES (Figs. 1, 11) Diagnosis. — Female with membrane pos- terior to seventh tergum bearing two small crescent-shaped sclerites. Head: Vertex and frontoclypeus uniform grayish orange; outer surface of labial palpus PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON grayish orange intermixed with brown scales tipped with white, inner surface grayish or- ange intermixed with white scales; antennal scape and pedicel grayish orange, flagello- meres brown; proboscis grayish brown. Thorax: Tegulaand mesoscutum grayish orange intermixed with brown scales tipped with white; outer surface of legs grayish or- ange intermixed with brown scales tipped with white; tibia and tarsomeres grayish or- ange apically; inner surface of legs grayish orange intermixed with white; forewing (Fig. 1), length 6.4-7.5 mm (n = 3), grayish or- ange intermixed with brownish-orange; scales tipped with white; two brownish-or- ange spots near apical area of discal cell, one spot near middle; undersurface uniform grayish brown; both surfaces of hindwing pale grayish orange; venation similar to B. acarta and B intrepida (Fig. 6). Abdomen: Grayish orange above, white beneath. Male genitalia: Unknown. Female genitalia (Fig. 11): Ovipositor telescopic, in four membranous divisions; ostium within membranous area posterior to seventh sternum, seventh tergum with stout setae; membrane posterior to seventh tergum with two small crescent-shaped sclerotizations; antrum membranous, short, and linear, forming a common inception for ductus seminalis and ductus bursae; ductus bursae long, with two rows of platelike scler- otizations within anterior end; corpus bur- sae membranous, signum hornlike. Holotype. —?, “Seychelles, Mahé, B[eau] Vallon, 20-VI-1959, H. Legrand,” “? gen- italia slide by DA 3273 [green label].’’ Ho- lotype in Muséum National D’Histoire Na- turelle, Paris (MNHP). Paratypes.—@, “Seychelles, Mahé, B[eau] Vallon, 19-IV-1959, H. Legrand,” “*e gen- italia slide by DA 3170 [green label],” ““Sey- chelles, Mahé, B. Vallon, 24-III-1959, H. Legrand,” “‘? genitalia slide by DA 3272 [green label].”’ Paratypes in MNHP. Remarks.—The three specimens of B. /e- grandi were intermixed in Legrand’s series of B. acarta and B. intrepida and can be VOLUME 97, NUMBER 3 491 Fig. 1. Holotype of Blastobasis legrandi Adamski. Fig. 2. Adult of Blastobasis acarta Meyrick. Fig. 3. Adult of Blastobasis intrepida Meyrick. 492 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Oy vA 8 CAN < WY Ae \ RR " UY W Rss \ j Z Fig. 4. Lateral view of head of male Blastobasis acarta Meyrick. Fig. 5. Lateral view of head of male Blastobasis intrepida Meyrick. VOLUME 97, NUMBER 3 493 Fig. 6. Wing venation of Blastobasis intrepida Meyrick. distinguished from female specimens of the latter two species only by examination of genitalia. Etymology.—This species is named in honor of H. Legrand who was devoted to collecting the Lepidoptera of The Sey- chelles. Blastobasis acarta Meyrick, 1911 (Figs. 2, 4, 7, 9) Blastobasis acarta Meyrick, 1911. Trans. Linn. Soc. Lond. 14(2): 286-87. Legrand, H. 1965. Mém. Mus. natn. Hist. nat. Ser. No 4. Diagnosis. — Proximal flange of valva with long setae, apical process of lower part of valva normal in size, aedeagus angled api- cally, membrane surrounding ostial area of female with dense microtrichia. Head (Fig. 4): Vertex and frontoclypeus with brownish-gray scales tipped with white; outer surface of labial palpus mostly brown intermixed with brownish-gray scales, basal and apical areas pale brownish gray or white; inner surface of labial palpus mostly white intermixed with grayish-brown and brown scales; antennal scape and pedicel with gray- ish-brown scales tipped with white; flagel- lomeres brown; male flagellum ciliate; first flagellomere of male with a subconical basal process, forming notch; proboscis grayish brown intermixed with pale-gray scales. Thorax: Tegula and mesoscutum with brownish-gray scales tipped with white; outer surface of legs mostly brown inter- mixed with pale brownish-gray and white scales, tibia and tarsomeres white apically; inner surface of legs mostly white inter- mixed with brownish-gray scales and pale- gray scales; forewing (Fig. 2), length 4.6-8.0 mm (n = 21), brownish gray intermixed with pale-gray scales, some specimens with scales tipped with white; two brown spots near apical area of discal cell, one near middle; undersurface uniform grayish brown; hind- wing with both surfaces pale brownish gray; venation similar to B. /egrandi and B. in- trepida (Fig. 6). 494 Fig. 7. Abdomen: Brownish gray above, white beneath. Male genitalia(Fig. 7): Uncus with some- what widened base, narrowed to apex, an- gled posteriorly; gnathos with two pointed teeth; tegumen setose dorsolaterally, dorsal strut of tegumen absent; diaphragma with microtrichia throughout; proximal flange of valva with dense microtrichia interspersed with several long, hairlike setae; aedeagus angled near base of anellus, anellus setose. Female genitalia (Fig. 9): As in B. la- grandi, except membrane surrounding os- tium with dense microtrichia. Types.—Lectotype here designated: 4, “Lectotype” [round purple-bordered label], “Type” [round red-bordered label], “‘Sey- chelles, Morne Blanc, Mahé, 800 flee]t, [X- 1908, H. Scott,” “Brit[ish] Mus[eum], 1913- 170,” “Blastobasis acarta Meyr{ick], Type 6,” “BM 4 genitilia slide no. 26553.’ Right labial palpus is missing. Lectotype in British Museum (Natural History) (BMNH). Para- lectotypes: 6, ““Paralectotype”’ [round blue- bordered label], ““Silhouette, Seychelles PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Male genitalia of Blastobasis acarta Meyrick (lectotype). Scale line = 0.5 mm. I[sland] [19]08,” ‘‘,” “‘Blastobasis acarta Meyr[ick] 1/1, E. Meyrick det., in Meyrick Coll[ection],” “‘acarta Meyr{ick],” [not dis- sected]. 2, ““Seychelles: Cascade Estate, Mahé, 800 flee]t, [X-1908, H. Scott,” “Brit[ish] Mus[eum], 1913-170.” The spec- imen is missing the abdomen, and the wings are not spread. Paratypes in BMNH. Other specimens examined: ¢, “‘Sey- chelles, Mahé, B[eau] Vallon, 28-II-1959, H. Legrand,” “‘é genitalia slide by DA 3168 [green label],”’ ““¢ wing slide by DA 3176 [green label],”’ , same data as above except: **18-VII-1950, M. Gerber,” ‘3, genitalia slide by DA 3171,” 9, “Seychelles, Mahé, Bleau] Vallon 2-I-1959, H. Legrand,” “‘? genitalia slide by DA 3169,” 722, same data as above except, ““6-V-1960,” “2 genitalia slide by DA 3172,” “20-IV-1956,” “Blas- tobasis intrepida Meyr.” [handwritten la- bel], “‘? genitalia slide by DA 3263,” “10- IV-1956,” “@ genitalia slide by DA 3264,” **12-I-1959,” “9, genitalia slide by DA 3265,” “11-III-1959,” “‘e genitalia slide by DA 3266,” “11-III-1959,” “‘? genitalia slide VOLUME 97, NUMBER 3 Fig. 8. by DA 3269,” “20-IV-1959,” “2 genitalia slide by DA 3270,” °, “Seychelles, Mahé, B[eau] Vallon, 12-VII-1959, M. Gerber,” “e genitalia slide by DA 3282,” 499, same data as above except, “8-I-1960,” “@ genitalia slide by DA 3277,” ‘25-I-1959,” “2 geni- talia slide by DA 3268,” “1-V-1959,” “9 genitalia slide by DA 3271,” “28-IV-1960,” “2 genitalia slide by DA 3276,” 4, ““Aldabra Atoll, 9°24’S, 46°20’E, Takamaka Grove, 12 Feb[ruary] 1968, Jay C. Shaffer,” ¢ ““USNM genitalia slide 81564, DA 3145 [green la- bel], 2 6, same data as above except, “13 Feb[ruary] 1968,” 6, same data as above except, “14 Feb[ruary] 1968,” 2 29, same data as above except, 11 Feb[ruary] 1968,” “‘¢e USNM genitalia slide 81565, DA 3146,” 5 2, same data as above except, “12 Feb[ruary] 1968,” °, same data as above except, “13 Feb[ruary] 1968,” 2 same data as above ex- cept, “14 Feb[ruary] 1968,” 3 2, same data as above except, “15 Feb[ruary] 1968,” “Pond No. 42, near Takamaka Grove,” 4, “Seychelles: Aldabra Atoll, Ile Picard, Set- Male genitalia of Blastobasis intrepida Meyrick (holotype). Scale line = 0.5 mm. tlement, 12-22 Mar[ch] 1986, David Adamski,” “? USNM genitalia slide 81559, DA 3140 [green label],” ¢, same data as above except, “? genitalia slide label 81561, DA 3142,” 3 2, same data as above except, “2 USNM genitalia slide 81560, DA 3141,” “2, USNM genitalia slide 81562, DA 3143,” “2, USNM genitalia slide 81563, DA 3144,” 6, “Seychelles, Mahé, Victoria, Botanical Gardens, 3-8 April] 1986, David Adam- ski,” “¢ USNM genitalia slide 81558, DA 3139 [green label].” Remarks.— The lectotype had previously been labelled, but no designation had been made. Because older specimens are faded, the color description is based upon the most recent specimens collected. Meyrick (1911) states that two males were collected at 1000 feet on Mahé (4 lectotype collected on Morne Blanc, and ¢ paralectotype collected at Cas- cade Estate). Label data show that both specimens were collected at 800 feet. Mey- rick (1911) also states that another paralec- totype male was collected on Silhouette at 496 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cn Fig. 9. Female genitalia of Blastobasis acarta Meyrick. Scale line = 1.0 mm. Fig. 10. Female genitalia of Blastobasis intrepida Meyrick. Scale line = 1.0 mm. VOLUME 97, NUMBER 3 Mare aux Cochons plateau at 1000 feet, but these data are not on the label of the pinned specimen. Blastobasis intrepida Meyrick, 1911 (Figs. 3, 5-6, 8, 10) Blastobasis intrepida Meyrick, 1911. Trans. Linn. Soc. Lond. 14(2) : 287. Legrand, H. 1965. Mém. Mus. natn. Hist. nat. Ser. A, 37: 54. Diagnosis. — Male with two apical articles of labial palpus widened dorsoventrally, an elongate valval spine present, apical process of lower part of valva small, aedeagus straight, apical part of aedeagal sclerite an- gled, lateral membrane with a round and wrinkled, small invagination between sixth and seventh segments. Head (Fig. 5): Similar to B. acarta, ex- cept male with two apical articles of labial palpus widened dorsoventrally (Fig. 5). Thorax: Tegula and mesonotum with brown scales tipped with white, intermixed with pale-brown scales; legs similar to B. acarta; forewing (Fig. 3), length 4.1-6.2 mm (n = 11), grayish brown intermixed with grayish-brown scales tipped with white or grayish-brown scales intermixed with brown scales; basal fascia distinct in specimens with basal area dominated by white scales; two brown spots near apical area of discal cell, one brown spot near middle; undersurface uniform grayish brown; hindwing with both surfaces pale brownish gray; venation sim- ilar to previous species (Fig. 6). Abdomen: Grayish brown above, white beneath. Male genitalia (Fig. 8): Uncus slightly angled posteriorly, apically rounded; gna- thos with two pointed teeth; tegumen with several dorsolateral setae; diaphragma with microtrichia dorsally; upper part of valva fingerlike, narrowed basally; apical process of lower part of valva small; elongate spine projecting about entire length of valva, fused to a subtriangular area (distorted on left val- va in Fig. 8); subtriangular plate with dense 497 Fig. 11. Female genitalia of Blastobasis legrandi Adamski. Scale line = 1.0 mm. 498 microtrichia; aedeagus straight, apical por- tion of aedeagal sclerite angled, anellus se- tose. Female genitalia (Fig. 10): As in previ- ous species except: ostial area membranous throughout; seventh sternum transversly shortened: lateral membrane between sixth and seventh segments with a round, wrin- kled, small invagination. Types. — Holotype, 4, “Holotype” [round red-bordered label], ‘““Type”’ [label is upside down], ‘‘Seychelles: Cascade Estate; Mahé, 800 flee]t; XII-1908, H. Scott,” “Brit[ish] Museum], 1913-170,” “Blastobasis intre- pida Meyr{ick], Type 3,” “BM genitalia slide 26552.” Holotype in BMNH. Other specimens examined.—é, “Sey- chelles, Mahé, B[eau] Vallon, 13-VI-1959, H. Legrand,” “*é genitalia slide by DA 3167 [green label], 4, same data as above except, 11-V-1959, M. Gerber,” 9°, “Seychelles, Mahé, B[eau] Vallon, 11-V-1959, M. Ger- ber,” ““?, genitalia slide by DA 3267,” 799 same data as above except, “20-V-1959,” “2 genitalia slide by DA 3274,” “7-VI- 1960,” ‘°° genitalia slide by DA 3275,” “15- VII-1959,” “2 genitalia slide by DA 3278,” **11-VI-1959,” “e genitalia slide by DA 3279,” “21-IV-1959,” “°° genitalia slide by DA 3280,” “7-VII-1959,” “‘9, genitalia slide by DA 3281,” “21-III-1960,” “e genitalia slide by DA 3283,” 2, same data as above except, “6-VI-1960, H. Legrand [specimen missing abdomen].” Remarks.— Female intrepida can be dis- tinguished from female /egrandi and acarta only by examination of genitalia. Discussion.—Adamski and Brown (1989) provided a phylogenetic classification for the North American Blastobasidae that includ- ed evidence for the monophyly of Blasto- basis Zeller. B. acarta shares many syna- pomorphies of the genus, except for the dor- sal strut of the tegumen and the posterior lobe of the corpus bursae. The cubital area of the hindwing is typical of several Pale- arctic species, e.g. Blastobasis phycidella Zeller, however, it is unlike most New World PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON blastobasids, in which the cubitus is stalked in series (veins M,, M,, CuA, and CuA, are stalked about the same distance from pos- terior wing margin). Similarly, B. intrepida shares several synapomorphies for Blasto- basis. Unlike acarta, B. intrepida does pos- sess the dorsal strut. However, B. intrepida has two characters, an elongate valval spine and the diminutive apical spine of the lower valva, that are found in Neoblastobasis Kuz- netsov and Sinev (1985) and related species recently described from the eastern Pale- arctic Region. The placement of B. legrandi is uncertain because males are not known. A reasonable assessment for the placement of species of Blastobasis awaits a phyloge- netic analysis of the world species. ACKNOWLEDGMENTS I thank K. Sattler of The British Museum (Natural History) for the loan of the types of Blastobasis acarta Meyrick and B. intre- pida Meyrick; G. Luquet of The Muséum National D’Histore Naturelle, Paris, for the loan of specimens of Blastobasidae collect- ed by H. Legrand and M. Gerber; and Ron- ald W. Hodges for providing names for specimens that were misidentified in Le- grand’s collection of Blastobasidae. I also acknowledge funding from the Smithsonian Institution and leadership from Brian Ken- sley, who headed our expedition in 1986. Finally, I express thanks to the Seychelles Islands Foundation for facilitating my field work on Mahé and Aldabra. LITERATURE CITED Adamski, D. and R. L. Brown. 1989. Morphology and systematics of North American Blastobasidae (Lepidoptera: Gelechioidea). Mississippi Agricul- tural Forest Experiment Station Technical Bulletin 165 (Mississippi Entomological Museum No. 1): 1-70. Clarke, J. F. G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bulletin Brooklyn En- tomological Society. 36: 149-161. Kornerup, A. and J. H. Wanscher. 1978. Methuen Handbook of Colour, 2nd ed. Methuen and Co., Ltd., London. 243 pp. VOLUME 97, NUMBER 3 499 Kuznetsov, V. I. and S. Yu. Sinev. 1985. Neoblas- Lionnet,G. 1984. Lepidoptera of Seychelles. Jn Stod- tobasis, a new genus of Blastobasidae (Lepidop- dart, D. R., ed., Biogeography and Ecology of The tera) from the USSR. (in Russian). Zoologicheskii Seychelles Islands. Dr. W. Junk Publishers, The Zhurnal 64: 529-537. Hague. XII+691 pp. Legrand, H. 1965. Lépidoptéres des iles Seychelles Meyrick, E. 1911. Tortricina and Tineina. The Percy et d’Aldabra. Mémoires du Muséum National Sladen Trust Expedition to The Indian Ocean in D’Histoire Naturelle. Series A, Zoologie 37: 1- 1905. Transactions Linnean Society London (2)14: 210. 263-307. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 500-507 A REVIEW OF NEW WORLD ENTOMOGNATHUS WITH DESCRIPTIONS OF SEVEN NEW SPECIES (HYMENOPTERA, SPHECIDAE, CRABRONINI) RICHARD M. BOHART Department of Entomology, University of California, Davis, California 95616-8584. Abstract.—A key is given to the 14 New World species of the genus Entomognathus Dahlbom (1894). Of the seven new species six are assigned to the subgenus Toncahua Pate: alaris (western U.S., Durango, Mexico), apache (Arizona), arizonae (Arizona, Mex- ico), coloratus (Arizona, New Mexico), guerreroi (Guerrero, Mexico), and occidentalis (California, Nevada). One new species is assigned to Entomognathus s.s.: dinocerus (Mo- relos, Mexico). Key Words: cahua The genus Entomognathus Dahlbom had 42' species assigned to it in 1976 (Bohart and Menke). All of these have hairy eyes and occur primarily in the Holarctic and Ethiopian Regions. A revised key to the four subgenera was given by Bohart and Menke (1976: 381). All seven of the previously known American species are in the subge- nus Toncahua Pate (1944) in which the males have only 12 antennal articles and some of the terga of both sexes have their posterior margins concave. In the present paper seven additional species are de- scribed, six in Toncahua, and one in the subgenus Entomognathus Dahlbom. Comments by reviewers, especially those by Karl Krombein, were most helpful. Thanks are also extended to curators of in- stitutions which have lent material (type de- position is indicated by the city involved in capitals): American Museum of Natural History (NEW YORK). ' There are now 45 species known. Marshakov (1976) described two from Asia and nasutus (Gribodo) (1884) was transferred from Lindenius by Leclerq (1989). Entomognathus, Hymenoptera, Sphecidae, Crabronini, New World, Ton- California Academy of Sciences (SAN FRANCISCO). Oregon State University Entomology Museum (CORVALLIS). University of Arizona (TUCSON). University of California Bohart Museum (DAVIS). University of Kansas Snow Museum (LAWRENCE). U.S. National Museum of Natural His- tory (WASHINGTON). Terms used in the key and descriptions which may be unfamiliar are: foveolate: di- vided into small cell-like units; flagello- meres: antennal articles beyond pedicel; MOD: median ocellus diameter; omaulus: carina at widest point of mesothorax; S-I to S-VII: abdominal sterna beyond propo- deum; T-I to T-VII: abdominal terga be- yond propodeum; prepectus: anterior sec- tion of mesopleuron below pronotal lobe. Key TO NORTH AMERICAN ENTOMOGNATHUS 1. Body length less than 4.0 mm (males) or 5.0 mm (females); prescutellar groove foveolate; wings clear or weakly, evenly stained ...... 2 VOLUME 97, NUMBER 3 Body length more than 4.0 mm (males) and at least 5.0 mm (females), prescutellar groove various, wings considerably stained (in most species) . Mesopleuron and scutum rather densely punctate, dull; flagellomeres white beneath in male, more dully in female; wings clear, pro- podeum granulate posteriorly and laterally; pronotal collar thin; Arizona WA etd i arizonae R. Bohart, new species Mesopleuron, at least above, polished; scu- tum with scattered, fine punctures, but pol- ished overall; other characters various . Mesopleural groove behind and adjacent to omaulus finely foveolate (as in Fig. 8), partly obscured by silvery pubescence; pronotal col- lar rounded over; mesopleuron with numer- ous small punctures, especially below, not highly polished; terga with whitish or yellow spots Mesopleural groove behind and adjacent to omaulus coarsely foveolate; pronotal collar narrow, not rounded; mesopleuron highly polished; terga unspotted . Terga partly or entirely rust red except for lateral whitish spots, female scutellum and metanotum all whitish, female clypeal bevel broadly lenticular, New Mexico coloratus R. Bohart, new species Terga black except for small lateral whitish yellow spots in female, absent or weak in male; female scutellum with two whitish yellow spots, small and more lateral in male, meta- notum with posterior linear spots; female clypeal bevel subtriangular, reddish; Arizona, w. Texas, Baja California, Sonora, Durango (Mexico) apache R. Bohart, new species . Propodeum posteriorly with flat, polished ar- eas flanking median furrow, also basolaterally polished, scape yellow in front, Central Amer- [CA ee a nD geometricus Leclerq Propodeum granulate posteriorly on either side of median furrow, basolaterally granulate, scape various . Wings slightly and evenly dusky, scape whit- ish yellow, Florida, North Carolina Bean ice eth oor arenivagus Krombein Wings hyaline, scape black, Mexico mexicanus Cameron . Wings hyaline, prescutellar groove foveolate, notauli somewhat impressed anteriorly to give scutum a tripartite appearance, Mexico .... nt etn gaat ree te BN AE eh evolutionis (Leclerq) Wings slightly or heavily stained, other char- ACLELSEV all OUS Mie ae ot rie ice re 8 . Mesopleuron at its widest point with or with- 10. ile 123 501 out sharp ridge, but without foveolate vertical groove just behind widest point (Fig. 7) ... 9 Mesopleuron at its widest point with omaulus or sharp ridge, adjacent to vertical foveolate groove (Fig. 8) . Mesopleuron at widest point rounded over, no omaulus or sharp edge (Fig. 7); punctation fine and separated, but extensive, surface not polished; prescutellar groove simple, not fov- eolate; scutellum whitish yellow, not 2-spotted, CaliforniasNevada: 2-year occidentalis R. Bohart, new species Mesopleuron at widest point with sharp edge; punctation indistinct, surface polished; pres- cutellar groove with at least a median longi- tudinal ridge; scutellum black or with two yel- low spots Prescutellar groove with four or more foveae separated by longitudinal ridges, markings pale yellow, female mandible weakly incised be- neath, U.S. east of Sierra Nevada, n. Mexico Oe he oa a SE Rin ere texanus (Cresson) Prescutellar groove in two sections separated by a median ridge; markings deep yellow; fe- male mandible sharply excised beneath; Guerrero Mexicome eer aaron guerreroi R. Bohart, new species Large species, about 15 mm long (female): occipital carina ending laterally in sharp tooth; top of clypeal bevel with pair of prominent, forwardly directed processes; wings dark brown; Morelos, Mexico ................. coo SRR Sst Bo dinocerus R. Bohart, new species Medium-sized species, less than 7.0 mm long; occipital carina not toothed laterally; top of clypeal bevel unarmed, wings various but not alitdark(Drownwer ris sare ace 12 Wings brown toward base, clear beyond cel- lular area; prepectal yellow spot present; scu- tellum often all yellow, sometimes two-spot- ted; w. U.S.: 100th meridian west to Arizona; Durango, Mexico alaris R. Bohart, new species Wings weakly stained, no prepectal spot, scu- tellum black or with two yellow spots . Terga black, U.S. east of 100th meridian .. . te EES RAS Corie Pah lenapeorum Viereck Terga with one or more whitish-yellow, lateral spots, U.S. east of 100th meridian Sets tot dae NOP bh Sister ea ETON: memorialis Banks SUBGENUS 7TONCAHUA PATE Toncahua Pate 1944: 341. Generotype Cra- bro texanus Cresson 1887, original des- ignation. 502 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON i Wes . she) E ma tera ily SLeANN ee LN * Vide = aN \\ \ se i \ eet - sed - ig = “yy Sig af SS / LMA NW / | I ANY me J) R\iy \ Ae A ‘ 2, a\ RAW ih \ ae: 7N\ \ < Ly, VEIL 7 Sy \ ~ Yenad ) SAIN Men : /, : \ aah 4; S yh AY ‘eas ; HN. 5 Z NS V=N = / Pll aura / L/ \ / / 1. occidentalis seeocus0O c 5. dinocerus 7. occidentalis 8. alaris | 1 IT T/!! 9. occidentalis 10. dinocerus Figs. 1-10. Species of Entomognathus. | (x37), 2 (* 18), front view of head. 3 (25), 4: (825) 551012); cellular area of forewing. 6 (x 18), clypeus from above. 7 (x37), 8 (< 25), mesopleuron. 9 (x50), mandible, side view. 10 (x25), mandible and end of occipital carina, side view. VOLUME 97, NUMBER 3 Florkinis Leclercq 1956: 2. Generotype En- copognathus evolutionis Leclercq 1956, original designation. The 13 species agree in the following characters: ocelli in a low triangle, male fla- gellum with 10 articles (male of coloratus unknown), clypeus with apical bevel ending in four to six teeth but no projections from upper margin of bevel, forewing recurrent reaching at or slightly before middle of mar- ginal cell (Fig. 3), T-III-IV posterior mar- gins broadly incurved, males customarily with flagellar articles whitish beneath, body length less than 10 mm. Entomognathus (Toncahua) alaris R. Bohart, NEw SPECIES Figs. 4, 8 Female holotype. Length 6.0 mm. Black, marked with pale yellow as follows: scape in front, mandible basally, pronotum all across including lobes, tegula in front, scu- tellum, metanotum, prepectal spot, forefe- mur and midfemur distally, all tibiae and basitarsi outwardly, large lateral spots (ivo- ry yellow) on T-I to T-V; flagellum yellow- ish beneath, S-VI reddish brown, basal 0.75 of forewing (Fig. 4) and all of hindwing brown. Pubescence of head (including eyes) pale. Most body surfaces polished, punc- tation sparse, a little thicker on T-V. Man- dible sharply incised basoventrally (as in Fig. 9); polished clypeal bevel elliptical (as in Fig. 1), 0.4 as long as clypeus whose medial apex has six small teeth; antennal sockets separated by a socket diameter, a little closer to eye; supraorbital fovea in- distinct; prescutellar groove foveolate; omaulus raised sharply, adjacent to narrow foveolate groove (Fig. 8); acetabular carina present; pygidial plate angled at 45°, mod- erately silvery setose. Male. Length 5.0-6.0 mm. Sculpture and markings about as in female. Prepectal spot may be small, band on collar ridge may be broken medially, metanotum may be most- ly black, T-VII and S-VII reddish brown; 503 T-VI closely punctate; clypeal apex with four apical teeth, bevel small. Holotype female (DAVIS), Willcox, Co- chise Co., Arizona, VIII-24-74 (R. M. Bo- hart). Paratypes, Arizona: 6 (DAVIS) same data as holotype; 2 6 (CORVALLIS), 8 mi n. Douglas, Cochise Co., VII-31-46 (H. A. Scullen); 2 (DAVIS), 28 min. Douglas, Co- chise Co., VIII-2-78 (R. W. Brooks); 2 (NEW YORK), 25 mi se. Saunders, Apache Co., VIII-14-72 (J. G. Rozen, R. Mc- Ginley); (NEW YORK); 11 mis. Apache, Cochise Co., VIII-10-73 (M. Favreau); 2 (SAN FRANCISCO), 4 mi sw. Forestdale, Navajo Co., VIII-24-52 (H. B. Leech). Oth- er paratypes: New Mexico: 2? (DAVIS), VIII- 23-64 (M. A. Cazier, et al.); 6 (DAVIS), 18 min. Rodeo, [X-7-76 (R. M. Bohart). Col- orado: 6 (WASHINGTON), (C. F. Baker); 2? (DAVIS), Great Sand Dunes Monument, Alamosa Co., VII-3-76 (H. E. Evans); Tex- as: 6 (DAVIS), Kingsville, Kleberg Co. III- 20-72 (J. E. Gillaspy); 4 ° (SAN FRAN- CISCO, DAVIS), 20 mi. se. Kingsville, Kle- berg Co., V-4-85 (W. J. Pulawski; 2 (SAN FRANCISCO, DAVIS), Junction, Kimble Co., IV-24-86 (W. J. Pulawsk1); 6° ““Tex.” (WASHINGTON); Kansas: 2? (WASHING- TON) Reno Co., VII-12-60; Wyoming: 2 ¢ (WASHINGTON, DAVIS), Glendo, VII- 13-60 (R. J. Lavigne); Mexico: Jalisco: 2 (DAVIS), (Crawford); Durango: 3 6, 2 (DA- VIS), 20 mi w. Durango, VI-17-64 (D. Bol- inger); (CORVALLIS), 76 mis. Durango, VI-16-56 (H. A. Scullen). The moderate size, distinct omaulus, with adjacent foveolate groove, bicolored wings, yellow prepectal spot and extensively yel- low pronotal collar and often scutellum, characterize the species. The bicolored wings and prepectal spot distinguish it from me- morialis. Entomognathus (Toncahua) apache R. Bohart, NEw SPECIES Female holotype. Length 4.5 mm. Black, marked with whitish yellow as follows: scape 504 in front, mandible mostly, pronotal collar except medially, pronotal lobe, tegula part- ly, lateral scutellar spot extending to hind- wing base, postlateral metanotal edge, fem- ora distally, fore- and midtibiae mostly, hindtibia basally, basitarsi, small lateral spots on T-I to IV; wings clear; clypeal bevel and mandible tip dark red, pygidial plate rust red. Pubescence pale. Punctation fine, scattered over upper frons, mesonotum, and mesopleuron, which are, however, polished overall; terga lightly punctate, polished; S-II rather closely punctate but shiny; propo- deum posteriorly granulate but enclosure areolate, laterally granulate but a little shiny. Mandible sharply incised basoventrally; polished clypeal bevel subtriangular, 0.6 as long as clypeus which has six weakly defined apical teeth; antennal sockets 1.0 diameter apart, 0.5 diameter from eye; supraorbital fovea present, narrowly oval; pronotal col- lar rounded over; omaulus present but ad- jacent foveolate groove indistinct; prescu- tellar groove finely foveolate; pygidial plate angled at 45°, densely silvery setose. Male. Length 2.5 mm. Sculpture, punc- tation, and pale markings about as in female except: terga all black, pronotum black ex- cept lobes, scutellum black (lateral pale lines present). Holotype female (NEW YORK), 13 mi sw. Apache, Cochise Co., Arizona, VIII-21- 71 (J. Rozen, M. Favreau). Paratypes: ¢ (SAN FRANCISCO), Baja California Sur, 9 km se. Santa Rita, VIII-25-77 (E. Fisher, R. Westcott); 2 (DAVIS), Dateland, Ari- zona, VII-5-56 (G. Butler); (NEW YORK), near Rodeo, New Mexico, VIII-25-71 (J. Rozen and M. Favreau); 66, @ (DAVIS, TUCSON), Roll, Arizona, VIII-21-54 and X-6-56 (D. Tuttle, G. Butler); ¢ (DAVIS), Ceballos, Durango, Mexico, X-19-68 (G.E. Bohart); 6 (DAVIS), 20 mi. s. Estacion Lla- no, Sonora, Mexico, VIII-17-64 (M.E. Ir- win). This species is similar to arizonae but the mesonotum and mesopleuron of apache are PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON more polished, and the clypeal bevel is sub- triangular rather than broadly oval. Entomognathus (Toncahua) arizonae R. Bohart, NEw SPECIES Female holotype. Length 4.0 mm. Black, marked with whitish yellow as follows: scape in front, mandible basally, pronotal collar laterally, pronotal lobe, tegula partly, lateral scutellar spot, femora distally, tibiae out- wardly, tarsi mostly, small lateral spots on T-I-II; wings clear, abdominal segment VI reddish brown. Pubescence pale. Puncta- tion fine and extensive; vertex, scutum, pro- podeum laterally, dull; terga and S-II with many punctures, somewhat shiny. Mandi- ble sharply incised basoventrally; polished clypeal bevel semicircular, 0.5 as long as clypeus whose medial apex has six small teeth, most lateral one sharp; antennal sock- ets 1.0 diameter apart, 0.5 diameter from eye; supraorbital fovea indistinct; prescu- tellar groove finely foveolate; no distinct omaulus or adjacent foveolate groove at front of pubescent prepectus; acetabular ca- rina present; pygidial plate angled at 45°, moderately silvery setose. Male. Length 3.0 mm. Sculpture, punc- tation, and pale markings about as in fe- male. Mandible often mostly whitish, fla- gellum mostly whitish yellow beneath, teg- ula and post-tegula partly yellow, terga without yellow spots, T-VI closely punctate. Holotype female (DAVIS), 28 mi n. Douglas, Cochise Co., Arizona, VIII-II-78 (R. W. Brooks). Paratypes (all from Cochise Co., Arizona): 3 6 (DAVIS, SAN FRAN- CISCO, WASHINGTON), same data as holotype; 6 (NEW YORK) 13 mi sw. Apache, VIII-14-74 (J., B., and K. Rozen); 2 6 (DAVIS, NEW YORK), 13 mi sw. Apache, VIII-20-71 (J. & D. Ehrenfeld). Other paratypes: 6 2? (DAVIS, WASHING- TON), Phoenix, Maricopa Co., Arizona; ¢ (TUCSON), Avra valley, Arizona, VII-6-55 (G. Butler, F. Werner); 6 (SAN FRANCIS- CO), 7 mi. s. Quartzite, Yuma Co., X-1-82 VOLUME 97, NUMBER 3 (W. J. Pulawski); 6 (SAN FRANCISCO), 3 mi. w. Superior, Pinal Co., VII-12-85 (W. J. Pulawski). Some of the paratypes were collected on Helianthus annuus, others on Euphorbia albomarginata. This small species (3.0—4.0 mm long) has the upper frons, vertex, mesonotum, meso- pleuron, and terga extensively punctate; the propodeal base not polished laterally; and the wings not stained. A somewhat similar species is arenivagus Krombein but it has the mesopleuron polished and its wings are dusky. Entomognathus (Toncahua) coloratus R. Bohart, NEw SPECIES Female holotype. Length 3.0 mm. Black, pale yellow, and rust red; whitish yellow are: scape, mandible on basal half, pronotal col- lar to lobe (all across), tegula partly, post- tegula mostly, scutellum and metanotum (nearly all white), femora distally, tibiae outwardly, lateral spots on T-I to IV, T-V mostly; rust red are: femora mostly, sterna, terga except for pale spots; wings lightly, evenly stained. Pubescence pale. Punctation fine and scattered on head, scutum, meso- pleuron, and abdomen, but these areas pol- ished overall; propodeum shiny laterally. Mandible sharply incised basoventrally; polished clypeal bevel crescentic, 0.3 as long as clypeus whose medial apex has six small teeth, most lateral one largest; anten- nal sockets about 0.7 diameter apart, 0.7 diameter from eye; supraorbital fovea dis- tinct; prescutellar groove foveolate; omau- lus weakly developed, adjacent to a foveo- late groove; acetabular carina present; py- gidial plate angled at 45°, moderately silvery setose. Holotype female (DAVIS), 1 mi. e. Doug- las, Cochise Co., Arizona, VIII-12-62 (M. A. Cazier). Paratypes: 2 2 same data as ho- lotype; 2 (NEW YORK), 4 mi. e. Animas, Hidalgo Co., New Mexico, VIII-24-74 (J. Rozen, M. Favreau). The presence of an omaulus and adjacent 505 groove on this small species together with the extensive rust-red abdominal coloration differentiate coloratus from other known members of the genus in North America. Also remarkable are the entirely whitish scutellum and metanotum, along with an all-dark clypeus. Entomognathus (Toncahua) guerreroi R. Bohart, NEw SPECIES Female holotype. Length 7.0 mm. Black, marked with deep yellow as follows: scape in front, mandible basally, inner streak on foretibia and midtibia, inner distal spot on hindtibia, large lateral spots which are at- tenuate medially on T-I to III, T-IV-V mostly; reddish brown are: scape partly, pronotal lobe, tegula, post-tegula; wings moderately brown stained, lighter basad. Pubescence silvery on face, off-silvery on terga, microsetae dark and plentiful on wing cells. Punctation faint, most body areas pol- ished, including propodeum anterolaterally. Mandible incised basoventrally; clypeal bevel not polished, irregular above, 0.5 x as long as clypeus whose medial apex has six small teeth; antennal sockets 1.0 diameter apart, 0.5 diameter from eye; prescutellar groove with a single median divider, not foveolate; omaulus represented by a sharp edge at front of prepectus, no adjacent fov- eolate groove; acetabular carina present; py- gidial plate angled at 45°, densely pale-gold- en setose. Male. Length 5.0-6.0 mm. Sculpture about as in holotype, yellow markings often more extensive: clypeal midlobe, four pro- notal spots, two spots on scutellum, tibiae and tarsi mostly, band across T-I, T-VI en- tirely. Some paratype males nearly as dark as type female; clypeal bevel less pro- nounced; and wings a little lighter. Holotype female (DAVIS), 9 mis. Tierra Colorado, Guerrero, Mexico, VII-21-63 (F. D. Parker and L. A. Stange). Paratypes (all from Guerrero, Mexico): 6 6 (DAVIS, SAN FRANCISCO, WASHINGTON), same 506 data as holotype; @ topotype (LAW- RENCE), VIII-5-51; 3 6 (DAVIS), Acapul- 60, -VI1I-19-63' (F. D! Parker and? 7A: Stange). M. guerreroi is similar to texanus in size, but averaging a little smaller. Also, the form of the prepectus, polished vertex, scutum, and mesopleuron are much the same. A striking difference is the deep-yellow of the markings in guerreroi rather than the ivory- yellow of texanus. In female mandibles the basoventral emargination is much more pronounced in guerreroi. Furthermore, the clypeal bevel of female guerreroi is less pol- ished and topped by a series of small bumps. In texanus the bevel is more polished and topped by a simple angle. The type series of guerreroi all have the prescutellar groove divided into two chambers, whereas 1n tex- anus it has five or six foveolae. Entomognathus (Toncahua) occidentalis R. Bohart, NEw SPECIES Figs.i1..3,)7,,9 Female holotype. Length 8.0 mm. Black, marked with whitish yellow as follows: scape, basal half of mandible, median clyp- eal lobe, pronotal collar all across, pronotal lobe, tegula mostly, post-tegula laterally, scutellum, femora distally, tibia and basal three tarsomeres outwardly, T-I with sub- apical band, broken medially, large lateral spots on T-II-III, band on T-IV, median spot on T-V; wings lightly stained, becom- ing clear basad. Pubescence of head (in- cluding eyes) pale. Punctation fine but ex- tensive on head, thorax, S-II, S-V, these ar- eas shiny but hardly polished; T-I to IV, S-III-IV_ polished with little punctation. Mandible sharply incised basoventrally (Fig. 9); polished clypeal bevel elliptical, 0.33 x as long as clypeus whose medial apex has six small teeth; antennal sockets about 1.0 diameter from eye and each other; supra- orbital fovea present but weak; prescutellar groove not foveolate; omaulus absent, wid- est point of thorax sharply rounded but not carinate and no adjacent foveolate groove PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Fig. 7); no acetabular carina; pygidial plate angled at 50°, moderately coppery setose. Male. Length 5.0-7.0 mm. Sculpture and markings about as in female. Hindtibia with large brown outer spot, apex of median clypeal lobe with four teeth, T-I with com- plete yellow band. Holotype female (DAVIS), Ash Moun- tain, Tulare Co., California, VI-26-83 (J. A. Halstead). Paratypes (all from California): 2 6(DAVIS, WASHINGTON), topotypical but taken VI-12-83 and VI-19-83 (D. J. Burdick); 2 6 (DAVIS), Borrego Valley, San Diego Co., IV-19-57 (J. C. Hall); 2 6 (DA- VIS), near Darwin Falls, Inyo Co., V-8-58 (E. G. Linsley) and V-12-69 (J. Powell); 3 (DAVIS), Warthan Canyon Road, Fresno Co., I-19-81 (N. J. Smith); ¢ (DAVIS), 8 mi ne. Auberry, Fresno Co., VI-11-81 (N. J. Smith); ¢ (DAVIS) near Santa Margarita, San Luis Obispo Co., V-22-58 (E. G. Lin- sley) and VI-8-62 (P. D. Hurd); ¢ (DAVIS), Carrizo Creek, Riverside Co., V-31-63 (E. I. Schlinger); 6 (SAN FRANCISCO), Pot- wisha, Sequoia National Park, Tulare Co., VII-1-41 (E. C. Van Dyke). Other para- types, 4 6 (SAN FRANCISCO, DAVIS), Candelaria, Mineral Co., Nevada, VI-23-52 (J. P. Figg-Hoblyn). The sharply rounded front margin of the prepectus, without an omaulus or adjacent groove (Fig. 7), and the extensive but mod- erate punctation on the head and thorax, characterize the species. The absence of an anterior prepectal groove occurs also in tex- anus and guerreroi, but these species have the prepectus sharp anteriorly, and the mesopleuron polished. Also remarkable is the simple, rather than foveolate, prescu- tellar groove in occidentalis. The male para- types from Nevada have slightly denser punctation on the mesopleuron and scutum than California specimens. Subgenus Entomognathus Dahlbom (1844: 295) Entomognathus dinocerus, described be- low, is the first record in the New World of VOLUME 97, NUMBER 3 a species in the typical subgenus. The pos- terior margins of T-III and T-IV are straight. A possibly important difference from E. brevis Vander Linden, the generotype, is that the recurrent vein of the forewing reaches the marginal cell slightly but definitely be- yond its middle (Fig. 5). Entomognathus (Entomognathus) dinocerus R. Bohart, NEw SPECIES Figs. 2.5.6, 10 Female holotype. Length 13 mm. Black, marked with deep yellow as follows: scape in front, two elongate spots on pronotal col- lar, pronotal lobe, scutellum mostly, meta- notum laterally, femora mostly, tibiae and tarsi, lateral spots on T-I to V, those on I separated by 4.0 MOD, irregular band on S-II; reddish brown are: mandible medially, tegula, post-tegula; wings dark brown (Fig. 5). Pubescence off-silvery on lower face, mesopleuron; fulvous on upper face, vertex, scutum, T-IV-V; microsetae of wing cells black. Punctation fine, sparse, body exten- sively polished. Mandible weakly indented basoventrally (Fig. 10), occipital carina toothed apically; clypeal bevel lenticular, surface irregular, shiny, 0.5 as long as clypeus, upper edge with a pair of stout, porrect lobes, lower edge with six irregular teeth, lateral ones large; antennal sockets separated by 1.3 socket diameter, 0.5 di- ameter from eye; scutum with long, well- marked median groove, notauli, and shorter parapsidal lines; prescutellar groove with four foveolae; omaulus sharply raised, ad- jacent to a foveolate groove; acetabular ca- rina absent; recurrent vein of forewing reaching marginal cell beyond its middle 507 (Fig. 5); T-II-IJI-IV posterior margins straight; pygidial plate angled at 50°, densely golden setose. Holotype female (DAVIS), Alpuyeca, Morelos, Mexico, VII-3-51 (P. D. Hurd). In this unusually large species, the tooth at the end of the occipital carina, and the projections on the dorsal rim of the clypeal bevel are reminiscent of some dinosaur fos- sil heads, hence the specific name. Also un- usual are the dark brown wings, grooved scutum, and slightly distal position of the recurrent vein (Fig. 5). The weak indenta- tion of the mandible basoventrally (Fig. 10) is found also in texanus, where it is less pronounced. LITERATURE CITED Bohart, R. M.and A. S. Menke. 1976. Sphecid wasps of the world. A generic revision. vii + 695 pp. University of California Press, Berkeley. Dahlbom, A. G. 1894. Hymenoptera Europaea prae- cipue borealia, etc., Vol. 1, xliv + 528 pp. Lund- borgiana, Lund. Gribodo. 1884. Spedizioni Italiano nell’Africa equa- toriale, risultati zoologici imenotteri. Annali Mu- seo Civico Storia Naturale, Genova (2) 1: 277- S23 Leclerq, J. 1956. Encopognathus (Florkinus, subg. nov.) evolutionis n.sp. Crabronien nouveau du Mexique. Bulletin Institut royal des Sciences na- turelles de Belgique 32(20): 1-12. 1989. In view of a monograph of the genus Lindenius Lepeletier and Brulle. Bulletin de la So- cieté des Sciences de Liége (58) 6: 419-444. Marshakov. 1976. Review of genera of the tribe Cra- bronini in the fauna of the USSR. III. Genus En- tomognathus Dahlbom. Zoologicheskii Zhurnal 55(4): 614-618 (in Russian). Pate, V.S.L. 1944. Conspectus of the genera of pem- philidine wasps. American Midland Naturalist 31: 329-334. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 508-514 NEW SPECIES OF NEARCTIC SNOW CRANE FLIES OF THE GENUS CHIONEA (DIPTERA: TIPULIDAE) GEORGE W. BYERS Snow Entomological Museum, Department of Entomology, University of Kansas, Law- rence, Kansas 66045-2119. Abstract. —Chionea jenniferae, n. sp., from southern California, and C. reclusa, n. sp., from southern Illinois, are described and illustrated. A note on variation in C. scita Walker is appended, and new occurrences of C. scita and C. valga are recorded. Key Words: When the North American species of Chi- onea were dealt with in some detail more than a decade ago (Byers 1983), I would not have predicted the discovery of any addi- tional nearctic species, except perhaps in re- mote boreal forests of Canada. But late in 1993 two remarkable new species were col- lected, one far south of the previously known range of the genus in California and the oth- er near the southernmost range in Illinois. I am indebted to Saul I. Frommer of the University of California, Riverside, and Michael A. Goodrich of Eastern Illinois University, Charleston, for forwarding these interesting insects to me. Discovery of these two species brings to 18 the number of species of Chionea known from North America. Chionea jenniferae, NEW SPECIES Description based on 7 males and 15 fe- males, preserved in alcohol. Diagnosis: A dark brown, southwestern species with 2 or 3 flagellomeres beyond fusion segment; sclerotized parts of ninth abdominal tergum of male widely separated medially by membrane, gonapophyses with pale, upturned apices; eighth abdominal ter- gum of female abruptly narrowed at sides, ninth sternum not divided medially. Chionea, Tipulidae, Limoniinae, distribution, variation, winter insects Head: Dark brown. Antennae (Fig. 1) comprising subcylindrical scape, long-tur- binate or subconical pedicel about as long as scape, fusion segment and three, less of- ten two, additional flagellomeres, if three the basal one shorter than others. Major se- tae on vertex about as long as diameter of eye, black, about six in number, directed forward. Compound eyes of 80-90 om- matidia, nearly same number and size in both male and female. Thorax: Sclerotized surfaces brown to dark brown, almost black in some males; membranous areas paler. Halteres light brownish gray. Sparse (6-8) dark setae on mesonotum, shorter setae on pronotum. Coxae and trochanters brown to dark brown, darker than yellowish brown to brown fem- ora and tibiae. Tarsi approximately same color as tibiae except fifth tarsomere darker. Femoral setae nearly as long as diameter of femur at level of attachment, in four dorsal rows and three often indistinct ventral rows; ventral setae on femora of robust males not in rows. Tibial setae longer than diameter of tibia. Abdomen of male: Terga 1-8 unevenly brown to dark grayish brown; posterior margin of each bearing 16-18 dark setae, 3-4 additional setae at each side and a few VOLUME 97, NUMBER 3 509 Figs. 1-8. Chionea jenniferae, n. sp. 1, left antenna, holotype, left lateral aspect. 2, terminal abdominal segments, male holotype, left lateral aspect (most setae omitted). 3, same, dorsal aspect. 4, gonapophysis, penis or aedeagus, associated apodemes and muscles, male paratype, right lateral aspect. 5, gonapophyses and basal enlargement of penis, dorsal aspect. 6, terminal abdominal segments, female allotype, left lateral aspect. 7, same, dorsal aspect. 8, ninth sternum and genital furca, female paratype, posteroventral aspect. Scale a, Figs. 1, 4-5, 8; scale b, Figs. 2-3, 6-7. 510 scattered, short ones. Corresponding sterna slightly paler. Pleura unevenly light grayish brown. Ninth tergum and sternum (Figs. 2, 3) dark brown, tergum broadly divided dor- somedially, membranous surfaces pale gray- brown, those of ninth and tenth terga con- tinuous. Lateral sclerites of tenth segment (tergal derivatives?) dark brown. Basistyles lighter brown than ninth sternum; disti- styles dark brown, mostly glabrous, widened at base but without distinct basal teeth. Gonapophyses (Figs. 4, 5) dark yellowish brown to brown except apical one-third yel- lowish, each slightly twisted subapically, apex widened and curved upward. Penis (Fig. 4) unusually elongate, conspicuously expanded and only moderately sclerotized at base; basal expansion projecting upward between bases of gonapophyses, then abruptly curved cephalad; slender portion recurved, with two curled apical filaments directed laterad. Abdomen of female: Tergum | uniform- ly dark yellowish brown. Terga 2-7 mostly yellowish brown, darker along anterior edge and very narrowly along posterior edge, at sides and diffusely along mid-line; posterior margin with about 10 black setae, scattered shorter setae elsewhere. Sterna more evenly yellowish brown, with about 20 dark brown to black setae along posterior margin, a few others scattered over surface. Pleural areas unevenly light brown. Tergum 8 abruptly narrowed at each side (Fig. 6); terga 9-10 short, wide (Fig. 7); cerci mostly light yel- lowish brown, darker along dorsal edge, slightly upcurved, with bluntly rounded apex (Fig. 6). Hypovalves light yellowish brown clouded with brown; eighth sternum brown. Genital furca (Fig. 8) lyre-shaped, with slen- der arms bowed outward. Ninth sternum formed of convergent slender prolongations from lower corners of tergum 9, with area of light sclerotization adjoining each; me- dian portion deflected somewhat ventrad. Body length (excluding antennae): Male, 3.51-4.34 mm. (holotype 3.59 mm.); fe- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON male, 3.90-6.48 mm. (allotype 6.34 mm.). Hind femur of holotype 2.39 mm. long, that of allotype also 2.39 mm. Specimens are variously contracted or extended in fluid preservative. Types: Male holotype, female allotype, 6 male and 14 female paratypes, collected from surface of snow in San Gorgonio Wil- derness, near Fish Creek Camp (4.3 km. northeast of Mt. Gorgonio, at elevation ap- proximately 2620 m.), in the San Bernar- dino National Forest, about 46 km. (29 mi.) east of San Bernardino, San Bernardino Co., California, on 13 November 1993, by Greg- ory P. Walker. Holotype, allotype, 2 male and 2 female paratypes in Snow Entomo- logical Museum, University of Kansas, Lawrence; 4 male and 12 female paratypes in collection of University of California at Riverside. In a letter, Dr. Walker describes the type locality as “‘. . . in a coniferous forest dom- inated by Jeffrey pine (Pinus jeffreyi) and white fir (Abies concolor). There is not much of an understory; it is pretty open with scat- tered shrubs and lots of bare soil. The ter- rain... was fairly steep and the ground was completely covered in snow.” At the request of the collector, Dr. G. P. Walker, this species is named for his six- year-old daughter, Jennifer, who first no- ticed the Chioneas, called them to her fath- er’s attention and helped collect them. Chionea jenniferae most closely resem- bles C. carolus Byers of the northern and central Sierra Nevada (Byers 1983: 169-172, figs. 163-170) and is somewhat less similar to C. lyrata Byers (1983) of the northern Sierras. Characteristics shared by jenniferae and carolus include small size, dark color, short antennal flagellum and, in males, the twisted, upturned apices of the gonapo- physes and the shape of the dististyles. Fe- males of these species are similar in the shapes of abdominal terga 8-10 and of the genital furca. Differences include, most sig- nificantly, the complete dorsomedial inter- VOLUME 97, NUMBER 3 ruption or separation of the ninth abdom- inal tergum in the male of jenniferae, a char- acter unique among all known species of Chionea. In carolus, the ninth tergum is deeply emarginate medially, more so than in any other species, yet is clearly contin- uous. While unlike that of any other nearctic species, the penis or aedeagus of the male in jenniferae has a form decidedly like that in C. lutescens Lundstrom and some other western palearctic species that have been differentiated from Chionea and assigned to a genus Niphadobata (cf. Burghele-Balaces- co, 1969: plates 2, 5). This may have a caus- al relationship to the membranous division of the ninth tergum, for (in preserved spec- imens) the penis arches dorsad and is deeply impressed against the underside of both the ninth and tenth terga. The structure of the genital fork and ninth sternum in the female shows no apparent adaptation to either the aedeagal curvature and length or the apical filaments* (i.e. no conspicuous differences from corresponding parts in females of oth- er species). The male keys to C. jellisoni at couplet 10 in my earlier key (Byers 1983: 123); the female goes to couplet 16 but not to either species identified there (p. 126). Chionea reclusa, NEw SPECIES Description based on one male specimen preserved in alcohol. Diagnosis: A light yellowish brown spe- cies with 13-segmented antennae; male dis- tistyles enlarged apically but without black- ened points near base, ninth abdominal ter- gum shallowly emarginate medially, apex of each gonapophysis pointed in dorsal aspect but truncated in lateral aspect. Head: Dark yellowish brown. Antennae (Fig. 9) with subcylindrical scape, pedicel about as long as scape, fusion segment and * In an earlier paper on Chionea (Byers 1983: 78), I stated incorrectly that apical filaments occur in C. valga and C. obtusa; these are present in C. valga and in C. hybrida. Silt ten additional flagellomeres. About 16 dark setae scattered over vertex, directed gener- ally forward. Compound eyes of approxi- mately 110 ommatidia each. Antennae and maxillary palps pale grayish brown; cervical sclerites grayish brown. Thorax: Most sclerotized surfaces yel- lowish brown; sternum, coxae and trochan- ters slightly paler yellowish brown; mem- branous areas and halteres pale buff-gray. About 12 prominent setae on mesonotum; shorter setae on pronotum. Wings minute, smaller than in other species of similar body size. Femora, tibiae and most of basitarsi yellowish brown; apical one-fourth of each basitarsus and all of other tarsomeres gray- ish brown. Most setae on hind coxa in outer vertical row; others on ventro-mesal sur- face. Hind femora only slightly thickened (as other femora), each with four dorsal rows of setae and three ventral rows. (Right mid- dle leg missing from holotype.) Abdomen of male: Terga 1-8 mostly yel- lowish brown, 2-7 grayish brown at each side and narrowly along anterior margin; tergum | darker at sides only; about 14 dark setae along posterior edge of each tergum, other setae at sides; no conspicuous setae on tergum 8. Sterna 2-7 light yellowish brown bordered on sides by light brown; 14-18 dark setae long posterior edge, a few others at each side and elsewhere on pos- terior half of each sternum. Pleural areas pale buff. Genital bulb pale yellowish brown with dark brown setae. Ninth tergum (Fig. 10) broadly rounded, only slightly emargin- ate medially, with 12-14 marginal setae at each side. Dististyles (Figs. 11, 12) without basal teeth, expanded and rounded apically, with blunt, darkly sclerotized, subapical point directed inward and somewhat for- ward. Gonapophyses (Figs. 13, 14) pale yel- lowish brown with brown lateral markings, with truncate dorsal apex and rounded, more ventrolateral apex. Penis slightly expanded near mid-length, bearing thin, rounded, yel- lowish flange at each side shortly before apex. 512 ‘ \ \ PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9-14. Chionea reclusa, n. sp., male holotype. 9, left antenna, left ventrolateral aspect. 10, ninth ab- dominal tergum (medial portion), dorsal aspect. 11, left dististyle and apex of basistyle, mesal aspect. 12, same, left lateral aspect. 13, gonapophysis and penis or aedeagus, associated apodemes and muscles, right lateral aspect. 14, same, dorsal aspect. Scale, all figures. Body length (excluding antennae): Male holotype, 4.90 mm.; length of hind femur, 30 Pimm, Type: Male holotype, collected in Mal- aise trap operated from 15 November to 30 December 1993, 11.2 km. (7 miles) west of Carbondale, Jackson Co., Illinois, by M. A. Goodrich and D. L. Wood. The holotype is in the Snow Entomological Museum, Uni- versity of Kansas, Lawrence. This site is in the edge of the Shawnee National Forest. Dr. Goodrich has de- scribed the habitat as ‘“‘a mature second- growth deciduous forest .. . (with) numer- ous standing dead trees and downed logs in the vicinity.”” The location is T9S, R2W, Sec. 20, NW 4. Chionea reclusa is so named because of its isolation from what appears to be its only close relative, C. wilsoni Byers of north- eastern Alabama (Latin rec/usa = separated, removed). Although the true extent of the ranges of both these species 1s as yet un- known, the collections that provide our only hints are separated by nearly 560 km. (350 miles). VOLUME 97, NUMBER 3 Chionea reclusa, as understood from the male only, belongs to the scita group of spe- cies (Byers 1983: 126) on the basis of its coloration, its elongate antennae, and its vertically oriented gonapophyses without upturned apical hook. It shares with C. wil- soni 13-segmented antennae and the unusu- ally shaped dististyles, which instead of ta- pering to an acute apex as in all other species are apically expanded, rounded and mesally concave. C. reclusa differs from wilsoni in the shapes of the gonapophyses, penis, ninth abdominal tergum, and in several details of the dististyles (cf. Byers 1983: 193, figs. 219- 226). The male keys to C. wil/soni at couplet 3 in the existing key (Byers 1983: 122). Chionea scita Walker Dr. Richard L. Hoffman, of the Virginia Museum of Natural History, Martinsville, sent for my examination a collection of six males and seven females of this species, which he caught in a pitfall trap set in a forest of mixed hardwoods, 25 October to 23 November 1991. The site is at an ele- vation of 1190 m. on a peak of the Blue Ridge called “‘The Priest,” in the George Washington National Forest, 7.2 km. southeast of Montebello, in Nelson Co., Virginia. This collection, while not large, shows considerable variation in dimensions of in- dividuals, such as has been noted in other species (e.g. Byers, 1983: 73, 148). The larg- est male, for example, is nearly twice the size of the smallest in overall length (ratio of 1.98 to 1). Earlier, I have used the hind femur as an indicator of the disproportion- ate development of “robust”? males of Chi- onea as compared to small, slender-legged males. In this case, while the ratio of length of hind femur is only 1.78 to 1, greatest femoral width shows a ratio of 2.75 to 1. The females also show great size difference, largest to smallest being in a ratio of 1.74 to 1 for overall length, but there is less vari- ation in dimensions of the ovipositor (ratios 513 of length of ovipositor, hypovalves, etc., be- ing 1.38—-1.40 to 1). Measurement of those parts of the male external genitalia likely to be in direct con- tact with the female ovipositor reveals much less variation than in body length, etc. Length of dististyle, for example, varies in this sample (largest male: smallest male) in a ratio of 1.25 to 1, and length of anterior edge of basistyle (taken as a measure of in- ner curvature, which is difficult to obtain without dissection) shows a ratio of only 1.05 to 1. The point of all this is that in their apparently haphazard search for mates on the snow surface (or in leaf litter, etc.), males need not go from one female to an- other until they reach one of corresponding overall size. This is only the second reported occur- rence of Chionea scita in Virginia, the other being far to the southwest in Smythe Coun- ty. The species, however, ranges fairly wide- ly, from Vermont westward to Michigan and southward to northern Georgia (but prob- ably only in the Appalachian Mountains, south of Pennsylvania). Two female specimens of this species were recently received from Dr. Peter Adler of Clemson University. These were collected in a Malaise trap set by a spring brook at Tanglewood Spring, near Pendleton, ele- vation 226 m, Anderson Co., South Caro- lina, | December 1987-3 January 1988, by John Morse. They constitute the first record of the occurrence of the genus Chionea in South Carolina. Chionea valga Harris A male of C. valga was recently received for examination from Dr. Charles Parker, research biologist in the Great Smoky Mountains National Park. It was found in a drift net (24-hour sample) set in Noland Creek, Swain Co., North Carolina, eleva- tion 1727 m, 17 February 1993, by G. Sal- ansky. This discovery extends the species’ range about 240 km (140 miles) south west- 514 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ward from its previously known range (Vir- meéridionale (Dipt. Tipulidae). Ann. Soc. Entomol. ginia and northward) and is the first re- France, nouv. ser. 5: 983-1000. é Byers, G. W. 1983. The crane fly genus Chionea in “C1 y in North Car- ; ys corded occurrence of C. valga orth Ca North America. Univ. Kansas Sci. Bull. 52: 59- olina. 195. LITERATURE CITED Burghele-Balacesco, A. 1969. Révision des genres Chionea et Niphadobata en Europe centrale et PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 515-547 NICHOLAS A. KORMILEV: A LIST OF HIS ENTOMOLOGICAL PUBLICATIONS AND PROPOSED TAXA RICHARD C. FROESCHNER National Museum of Natural History, NHB 127, Smithsonian Institution, Washington, DrC20560: Abstract. —A list of the 231 entomological publication titles and the taxonomic names proposed, alone or in coauthorship, by Nicholas A. Kormilev from 1936 to 1992 is given. Names proposed in two orders of insects—for Heteroptera: 1 family; 2 subfamilies; 6 tribes; 105 genera and subgenera; and 1182 species and subspecies; for Neuroptera: | generic name; | subspecies name. Key Words: Nicholas Alexander Kormilev, born Jan- uary 29, 1901, in Yalta in the Crimea, pub- lished the first of his 231 entomological ar- ticles in 1936. Nearly all of these 231 en- tomological articles were on members of the insect group the Hemiptera-Heteroptera, including proposals of one family, two sub- families, six tribes, 105 genera and subgen- era, and 1182 species and subspecies. In the insect order Neuroptera he proposed one new genus and one new subspecies. His early life was made difficult by the Balkan Wars, the Russo-Japanese War, eth- nic battles, revolutions, frequent family moves because of his father’s changes of military assignments, World War I, military service, and World War II; but his perse- verance, his expanding knowledge, and his familiarity with several languages enabled him to obtain employment in a variety of jobs—academic and otherwise. In 1926 he graduated from the School of Agriculture at the University of Zagreb, in Yugoslavia. Then came employment with the Ministry of Education and the Ministry of Agriculture. While working at Skoplje he frequently visited the Natural History Mu- seum, and at the urging of its director he Kormilev, lists, publications, taxa, Heteroptera, Neuroptera published his first paper in 1936. In 1947 he emigrated to Argentina where he became an entomologist with the Museo Argentino de Ciencias Naturales “Bernardino Riva- davia,” Buenos Aires. He was also em- ployed at the Institute de Investigacion Mu- seum, Anex de Colegio Maximo de San Jose, en San Miguel. In 1957 he came to the Unit- ed States and in New York worked at an electric corporation in quality control and production of silver-zinc batteries. In New York he began studies in the collections at the American Museum of Natural History. After retirement in 1968 he concentrated his research on two families of Heteroptera, the flat bugs (Aradidae) and the ambush bugs (Phymatidae). Then, with help from Dr. J. L. Gressitt, he went to Hawaii and there wrote his large monograph of the flat bug subfamily Mezirinae of the Oriental Region and the South Pacific (Kormilev 197 1a). In 1970 he spent some time in Australia before returning to the United States. His early, European collection was de- posited in the Serbian Museum. His later Heteroptera collection and and sizable tiger beetle (Cicindelidae) collection were depos- ited in the Smithsonian Institution’s Na- 516 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Nicholas Kormilev, 1984. tional Museum of Natural History in Wash- ington, D.C. Nicholas, or ““Nick”’ as he is affectionately called by entomological colleagues, has al- ways been an avid scholar who willingly served his science by identifying large num- bers of Heteroptera for others as well as through his numerous publications. But his interests are broader than that, and he often made special efforts to see collections of art and historically important landmarks. Now, because “Nick,” who will be 94 in 1995, told me he has published his last pa- per, it seems appropriate to furnish the sci- entific community with a complete listing of his many entomological publications and proposed names. This paper is arranged in two parts: The first contains a chronological list of Kor- milev’s publications, with coauthors listed in parentheses at the end of each title. The abbreviated journal titles are given in full at the end of the literature section. The sec- ond list contains all taxa he proposed, ar- ranged by family-group, genus-group, and species-group, followed by a date-page ref- erence to the above-mentioned list of pub- lications. List OF PUBLICATIONS 1936a I. Beitrag zur Kenntnis der Verbreitung jugo- slavischer Hemiptera-Heteroptera (Serbien und Siidserbien). Bull. Soc. Sci. Skoplje 17(5): 29-54, 5 figs. 1938a II. Beitrag zur Kenntnis der Verbreitung jugo- slavischer Hemiptera-Heteroptera (Serbien und Siidserbien). Bull. Soc. Sci. Skoplje 18(6): 167-172, 1 fig. 1939a III. Beitrag zur Kenntnis der Verbreitung ju- goslavischer Hemiptera-Heteroptera (Serbien und Siidserbien). Ann. Mus. Serbiae Merid., Skopje 1(2): 21-29, 2 figs. 1939b IV. Beitrag zur Kenntnis der Verbreitung ju- goslavischer Hemiptera-Heteroptera. Bull. Soc. Sci. Skoplje 20(7): 193-199, 6 figs. 1943a V. Beitrag zur Kenntnis der Verbreitung bal- kanischer Hemiptera-Heteroptera (Serbien und Ma- zedonien). Roy. Serbien Acad. Sci. Separate issue 136, Natur. Sci. and Math., 35, Ohridski Sbornik, 6: 123-132, 1 map. 1948a Una especie nueva de la familia Elasmodemi- dae L. & S. de la Republica Argentina. Rev. Soc. Ent. Argent. 14: 141-147, 2 figs. 1949a Notas sobre los Colobathristidae de Bolivia. Notas Mus. La Plata, 14, Zool. 124: 167-176, 3 figs. 1949b A new species of the genus Macrocephalus Swederus from Brasil (Hemiptera-Phymatidae). Rev. Ent. Rio de Jan. 20 (1-3): 583-588, 1 fig. 1949c Una especies nueva del genero Placocoris Mayr en la Argentina (Hemiptera-Pentatomidae). Com. Inst. Nac. Inv. Cien. Nat. Bs. As., Zool. 1(9): 3-12, 3 figs. 1949d Riggiella vianai n. g., n. sp. de la Argentina (Hemiptera-Lygaeidae). Com. Inst. Nac. Inv. Cien. Nat. Bs. As., Zool. 1(11): 3-13, 4 figs. 1949e La familia Colobathristidae Stal en la Argen- tina, con la descripcion de tres especies nuevas neo- tropicales (Hemiptera). Acta Zool. Lilloana 7: 359- 383, 6 figs., 1 map. 1949f Dos especies nuevas del genero Adoxoplatys Breddin, 1903, de Bolivia (Hemiptera-Pentatomi- dae). Notas Mus. La Plata 14, Zool., 128: 313-324, 2 figs. 1950a La subfamilia Oxycareninae Stal en la Argen- tina, con la descripcion de una especie nueva (He- miptera-Lygaeidae). An. Soc. Cien. Argent. 149: 22- 32, 7 figs. 1950b Notes on Neotropical Pentatomidae, with de- scription of one new genus and two new species (Hemiptera). Rev. Brasil. Biol. 10: 339-346, 5 figs. 1950c Notes on Neotropical Phymatidae; diagnosis of new species (Hemiptera). Rev. Ent. Rio de Jan. 21(3): 581-591. 1950d Notas Hemipterologicas. Acta Zool. Lilloana 9: 495-503, 4 figs., 1 pl. 195la Notas sobre Colobathristidae Neotropicales VOLUME 97, NUMBER 3 (Hemiptera), con la descripcion de tres géneros y siete especies nuevos. Rev. Brasil. Biol. 11(1): 63- 84, 17 figs. 1 map. 1951b Notas sobre Phymatidae Neotropicales III, con la descripcion de dos especies nuevas (Hemiptera). Mis. Est. Patol. Reg. Argent. 22(79): 53-58, 4 figs. 1951c Sobre los géneros Vulsirea Spinola (1837), Ra- mosiana Kormilev (1950) y Adoxoplatys Breddin (1903), con la descripcion de tres especies nuevas (Hemiptera-Pentatomidae). Rev. Soc. Ent. Argent. 15: 83-95, 13 figs. 1951d Aradidae Argentinos I. (Hemiptera). Com. Inst. Nac. Inv. Cien. Nat. Bs. As., Zool. 2(6): 83-95, 1 map. 195le Notas sobre Phymatidae Neotropicales II, con la descripcion de cuatro especies nuevas (Hemip- tera). An. Soc. Cien. Argent. 152: 167-178, 4 figs. 1951f Notas sobre Colobathristidae Neotropicales II, con la descripcion de dos especies nuevas (Hemip- tera). Rev. Brasil. Biol. 11(3): 333-339, 3 figs. 1951g Phymatidae Argentinas (Hemiptera), con ob- servaciones sobre Phymatidae en general. Rev. Inst. Nac. Inv. Cien. Nat. Bs. As., Zool. 2(2): 45-110, 8 figs., pls. 1-14, 3 maps. 1951h Notas sobre Hemipteros Bolivianos, con la des- cripcion de un género y tres especies nuevas. Publ. Univ. Cochabamba, Bolivia, Folia Univ. 5: 35-45, 3 figs. 1952a Notas Hemipterologicas II. Dusenia 3(1): 51- 565 pl. 3: 1952b La subfamilia Graphosomatinae Jakovlev en la Argentina (Hemiptera-Pentatomidae), con la des- cripcion de un genero y una especie nuevos. An. Soc. Cien. Argent. 153: 212-218, 3 figs. (senior au- thor with A. A. Piran). 1952c Notes on Neotropical Aradidae, with descrip- tion of one new species. Pan-Pac. Ent. 23(2): 119- 122, 2 figs. 1952d Notas sobre Coreidae Neotropicales con la des- cripcion de dos especies nuevas (Hemiptera). EOS 28(1): 91-99, 7 figs. 1952e Notas sobre Phymatidae Neotropicales IV (He- miptera-Reduvioidea). Mis. Est. Patol. Reg. Argent. 23(81-82): 123-130, 4 figs. 1952f Los Insectos de las Islas Juan Fernandez, 1. Ly- gaeidae (Hemiptera). Rev. Chilena Ent. 2: 7-14, 4 figs. 1952g Una especie nueva del genero Ochterus La- treille, 1807, del Paraguay (Hemiptera-Ochteridae). An. Soc. Cien. Argent. 154: 155-159 (senior author with J. A. De Carlo). 1952h Unaespecie nueva del genero G/yphepomis Berg, 1891, de la Argentina (Hemiptera-Pentatomidae). Rev. Soc. Ent. Argent. 15: 302-306, 4 figs. (as senior author with A. A. Piran). 1953a The first apterous aradid from Argentina (He- miptera). Dusenia 4(2): 125-126, 1 fig. 517 1953b Revision de Micrelytrinae Stal de la Argentina. Con descripcion de un género y siete especies nuevos de Argentina, Brasil y Bolivia (Hemiptera, Coris- cidae). Rev. Soc. Ent. Argent. 16(2): 49-66, 6 figs., 1 map. 1953c Notes on Aradidae from the Eastern Hemi- sphere (Hemiptera). Verh. Naturf. Ges. Basel 64(2): 333-346, 8 figs. 1953d Notes on Colobathristidae from Borneo and Java in the Collections of the Museum of Natural History in Basle (Hemiptera). Bull. Soc. Ent. Suisse 26(4): 287-292. 1953e Notes on Neotropical Aradidae III (Hemip- tera). On some apterous Mezirinae from Brasil. Du- senia 4(4—5): 229-242, pl. 18. 1953f Notas sobre Phymatidae Neotropicales V (He- miptera). Mis. Est. Pat. Reg. Argent. 24(83-84): 57- Gill tie? 1953g Notas sobre Phymatidae Nearcticas I (Hemip- tera). Mis. Est. Patol. Reg. Argent. 24(83-84): 63- 67, 7 figs. 1954a Notas sobre Coreidae Neotropicales II (He- miptera), Merocorinae de la Argentina y paises lim- itrofes. Rev. Ecuat. Ent. Parasit. 2(1-2): 153-186, 7 figs. 1954b Notas sobre Aradidae del hemisferio Oriental V.—Aradidae de Espana e Islas Canarias (Hemip- tera-Aradidae). Rev. Ecuat. Ent. Parasit. 2(1-2): 203- 208, 3 figs. 1954c Notes on Nearctic Phymatidae II (Hemiptera): a new subspecies of Macrocephalus prehensilis (F.) from Florida. Jour. Kans. Ent. Soc. 27(4): 158-159. 1954d Una familia nueva para la fauna Argentina (Hemiptera-Megaridae). An. Soc. Cien. Argent. 157: 47-54, 8 figs., 1 map. 1954e Notes on Neotropical Aradidae IV (Hemip- tera). On some apterous Mezirinae from Brasil II. Dusenia 5(3-4): 125-130, 2 figs. 1954f Notes on Aradidae from the Eastern Hemi- sphere (Hemiptera). II, Aradidae from the Oriental and Australian Regions in the Hungarian National Museum, Budapest, Hungary, Philipp. Jour. Sci. 83(2): 123-136, 5 figs. 1955a Aradidae (Hemiptera) Argentinae II. Acta Zool. Lilloana 13: 207-256, 52 figs., pls. I-III, 2 maps (1953). 1955b Notas sobre Tingidae Neotropicales (Hemip- tera). Rev. Brasil. Biol. 15(1): 63-68, 1 fig. 1955c Notes on Aradidae from Eastern Hemisphere III (Hemiptera). On some Aradidae from Japan, Russian Far East and Formosa (Taiwan). Mushi 28(4): 31-41, pl. 6. 1955d Notes on Aradidae from the Eastern Hemi- sphere IV (Hemiptera-Heteroptera). Aradidae from the Oriental and Australian Regions. —2. Ann. Mag. Nat. Hist. ser. 12, 8: 194-203, 12 figs. 1955e Notas sobre Pentatomoidea Neotropicales II 518 (Hemiptera). Acta Sci. Inst. Inv. de San Miguel, Prov. de Bs. As. 1: 1-15, 7 figs. 1955f Notas sobre Lygaeidae Neotropicales I (He- miptera); la subfamilia Cyminae (Stal) en la Argen- tina. Acta Sci. Inst. Inv. San Miguel, Prov. de. Bs. As. 2: 1-8, 7 figs. 1955g Una curiosa familia de Hemipteros nueva para la fauna Argentina, Thaumastotheriidae (Kirkaldy), 1907. Rev. Soc. Ent. Argent. 18: 5-10, 9 figs. 1955h Notes on Aradidae from the Eastern Hemi- sphere VIII (Hemiptera). Aradidae from the Ori- ental Region VI. Verh. Naturf. Ges. Basel 66(2): 141-146, 3 figs. 1955i La subfamilia Cyrtocorinae Distant en la Ar- gentina (Hemiptera: Pentatomoidea). Rev. Ecuat. Ent. Parasit. 2(3—4): 321-334, 2 pl. 1955} A new myrmecofil family of Hemiptera from the delta of Rio Parana, Argentina. Rev. Ecuat. Ent. Parasit. 2(3—4): 465-477, 1 pl. 1955k Notes on Aradidae from the Eastern Hemi- sphere VI (Hemiptera). Aradidae from the Oriental and Australian Regions-IV. Rev. Ecuat. Ent. Par- asit. 2(3—-4): 485-508, 2 pl. 19551 Notes on Aradidae from the Eastern Hemi- sphere (Hemiptera). VII. Aradidae from Fukien, South China. Quart. Jour. Taiwan Mus. 8(3): 177- 191, pls. 1-2. 1956a Notas sobre Pentatomoidea Neotropicales IV (Hemiptera). Acta Scient. Inst. Inv. de San Miquel, Prov. Bs. As. 3: 1-12, 12 figs. 1956b Notas sobre Pentatomoidea Neotropicales V (Hemiptera). Acta Scient. Inst. Inv. de San Miguel, Prov. Bs. As. 4: 3-7. 1956c Notes on Aradidae from the Eastern Hemi- sphere, IX (Hemiptera). Aradidae from the Ethio- pian Region. Ann. Mag. Nat. Hist. ser. 12, 9: 250- 256, 5 figs. 1956d Notes on Aradidae from the Eastern Hemi- sphere, X (Hemiptera). On some apterous Meziri- nae from India and Indonesia. Philipp. Jour. Sci. 85(2): 283-294, 9 figs. 1956e Notes on Aradidae from the Eastern Hemi- sphere XII (Hemiptera). Mitt. Miinchner Ent. Ges. 46: 42-46, 8 figs. 1957a Notes on Aradidae from the Eastern Hemi- sphere, XI (Hemiptera). On some apterous Mezir- inae from New Guinea. Philipp. Jour. Sci. 85(3): 389-403, 20 figs. (1956) 1957b Notes on American ““Phymatidae” (Hemip- tera). Rev. Brasil. Biol. 17(1): 123-138, 14 figs. 1957c Notas sobre Aradidae Neotropicales VI (He- miptera). An. Soc. Cien. Argent. 162: 148-159, 1 pl. (1956). 1957d Notas sobre Pentatomoidea Neotropicales VI (Hemiptera). An. Soc. Cien. Argent. 163: 47-57, 10 figs. 1957e Notes on Aradidae from the Eastern Hemi- sphere (Hemiptera) XIII. On some Aradidae in the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Drake Collection. Quart. Jour. Taiwan Mus. 10(1): 37-46, 1 pl. 1957f Notes on Aradidae from the Eastern Hemi- sphere XIV (Hemiptera). Aradidae from the Onr- ental and Australian Regions IX. Ann. Mag. Nat. Hist. ser. 12, 10: 265-273, 11 figs. 1957g Notes on Neotropical Aradidae (Hemiptera), VII. Two new apterous Aradidae. Amer. Mus. Nov- it. 1860: 1-7, 4 figs. 1957h Los Insectos de las Islas Juan Fernandez. 30. Lygaeidae (Supplemento) (Hemiptera). Rev. Chi- lena Ent., 5: 385-389, 4 figs. 19571 On some Phymatidae in the American Museum of Natural History (Hemiptera-Heteroptera). Jour. N. Y. Ent. Soc. 65: 33-40, 2 figs. 1957} Notes on oriental Phymatidae (Hemiptera). The oriental Phymatidae in the Drake Collection. Quart. Jour. Taiwan Mus. 10: 63-69, 2 figs. 1958a Concerning the apterous Aradidae of the Amer- icas (Hemiptera). Ann. Ent. Soc. Amer. 51(3): 241- 247, 2 figs. [junior author with C. J. Drake]. 1958b Notes on Aradidae from the Eastern Hemi- sphere XV (Hemiptera). Jour. N. Y. Ent. Soc. 66: 87-97, pl. 5. 1958c Notas Hemipterologicas III. Acta Zool. Lil- loana 16: 55-59. 1959a Notes on Aradidae in the U.S. National Mu- seum (Hemiptera). I. Subfamily Calisiinae. Proc. U.S. Nat. Mus. 109(3413): 209-222, 18 figs. (1958). 1959b Notes on Neotropical Aradidae II (Hemiptera). Proc. Ent. Soc. Wash. 61(2): 61-71, 6 figs. 1959c Notas sobre Aradidae Neotropicales V (He- miptera). Sobre el género Pictinus Stal, 1873. Rev. Soc. Uruguaya Ent. 3: 21-33, pl. 1. 1959d Notas sobre Mantispidae neotropicales I (Neu- roptera). Rev. Soc. Ent. Argent. 21: 1-18, 24 figs. [junior author with G. W. Williner]. 1959e Notes on Neotropical Aradidae. IX (Hemip- tera). Studia Ent. 2(1-4): 309-320, 6 figs. 1960a Notes on Aradidae in the U.S. National Mu- seum (Hemiptera) II. Jour. N. Y. Ent. Soc. 68: 36- 47, 1 pl. 1960b Notes on Aradidae from the Eastern Hemi- sphere X VI (Hemiptera). Proc. Ent. Soc. Wash. 62(2): 106-107. 1960c Notas sobre Aradidae Neotropicales VIII (He- miptera). An. Soc. Cient. Argent. 169(5-6): 83-94, 5 figs. 1960d Notas sobre Aradidae Neotropicales X (He- miptera). Rev. Soc. Uruguaya Ent. 4: 3-17, pl. I. 1960e Notes on Neotropical Aradidae XI (Hemip- tera). Jour. N. Y. Ent. Soc. 68: 208-220, 9 figs. 1960f Notes on Aradidae from the Eastern Hemi- sphere X VII (Hemiptera). Quart. Jour. Taiwan Mus. 13(3-4): 163-174, 2 figs. 196la Notes on Aradidae from the Eastern Hemi- sphere X VIII (Hemiptera). Jour. Ent. Soc. S. Africa 24: 248-252, 2 figs. VOLUME 97, NUMBER 3 1961b Concerning Phylus breviceps Reuter, 1900, and P. balcanicus Kormilev, 1939 (Hemiptera: Miri- dae). Ent. Month. Mag. ser. 4, 256: 77-78, 3 figs. 1962a Notes on American Phymatidae IT (Hemiptera, Reduvioidea). Jour. N. Y. Ent. Soc. 70: 47-58, 15 figs. 1962b Notes on Aradidae in the Naturhistoriska Riks- museum, Stockholm (Hemiptera-Heteroptera). Ar- kiv for Zool., Ser. 2, 15(14): 255-273, 15 figs. 1962c Notes on African and Asiatic Macrocephalinae (Hemiptera: Phymatidae). Amer. Mus. Novit. 2107: 1-15, 20 figs. 1962d Revision of Phymatinae (Hemiptera: Phyma- tidae). Philipp. Jour. Sci. 89(3-4): 287-486, pls. 1- 19 (1960). 1962e Notes on Phymatidae in the British Museum (Nat. Hist.) (Hemiptera-Heteroptera). Ann. Mag. Nat. Hist., ser. 13, 5: 349-367, 20 figs. 1963a Notes on Aradidae in the Naturhistoriska Riks- museum, Stockholm, Hemiptera-Heteroptera. Ar- kiv for Zool. Ser. 2, 15(31): 443-455, 4 figs. 1963b On some Calisiinae in the British Museum (Nat. Hist.) (Hemiptera-Heteroptera, Aradidae). Ann. Mag. Nat. Hist. ser. 13, 5: 601-607, 9 figs. 1963c Notes on American Phymatidae III (Hemip- tera-Heteroptera). Proc. Ent. Soc. Wash. 65(3): 215- 226, 21 figs. 1964a Notes on Aradidae in the U.S. National Mu- seum III. Subfamily Mezirinae (Hemiptera). Proc. U.S. Nat. Mus. 115(3483): 245-258, 7 figs. 1964b Neotropical Aradidae XII (Heteroptera: Ar- adidae). Jour. N. Y. Ent. Soc. 72: 34-39, 6 figs. 1964c New Genera and Species of Queensland Ar- adidae (Hemiptera: Heteroptera). Jour. Ent. Soc. Queensland 3: 42-47, 7 figs. 1964d Neotropical Aradidae XIII (Hemiptera: Ar- adidae). Jour. N. Y. Ent. Soc. 72: 112-119, 17 figs. 1964e The Third Species of Neoanthylla Kormilev, 1951, from Peru (Hemiptera: Phymatidae). Proc. Ent. Soc. Wash. 66(3): 137-140, 4 figs. 1964f Notes on Aradidae in the Naturhistoriska Riks- museum, Stockholm (Hemiptera-Heteroptera). Ark. for Zool. Ser. 2, 16(23): 463-479, 21 figs. 1964g Notes on Neotropical Aradidae XIV (Hemip- tera: Heteroptera). Studia Ent. 7(1-4): 153-160, 8 figs. 1964h Conserning [!] the genera Reeceicus Drake and Signocoris Hoberlandt, Hemiptera-Heteroptera, Aradidae. Quart. Jour. Taiwan Mus. 17(3-4): 187- 191, 2 figs. 1965a Notes on Aradidae in the Naturhistoriska Riks- museum, Stockholm (Hemiptera Heteroptera). Ar- kiv for Zool. Ser. 2, 18(1): 1-8, 5 figs. 1965b Notes on Australian Aradidae (Hemiptera: Heteroptera), with descriptions of new genera and species. Proc. R. Soc. Queensland 77(3): 11-35, 11 figs. 1965c Two new species of the genus Phymata Latreille S19 from Brasil (Hemiptera-Heteroptera, Phymatidae). Studia Ent. 8(1-4): 205-208, 4 figs. 1965d Notes on Neotropical Aradidae XV (Hemip- tera-Heteroptera). Opusc. Zool. 84: 1-7, 10 figs. 1966a On some Aradidae from Africa and Polynesia (Hem.-Heteroptera). EOS 41(2-3): 387-394, 6 figs. 1966b Two new American Aradidae (Hemiptera-Het- eroptera). Psyche 73(1): 26-29, 8 figs. 1966c A new species of the genus Pictinellus Usinger & Matsuda, 1959 (Hemiptera: Aradidae). Proc. Ent. Soc. Wash. 68(4): 306-308, | fig. 1966d Notes on Aradidae in the U.S. National Mu- seum, IV (Hemiptera: Heteroptera). Proc. U.S. Nat. Mus. 119(3548): 1-25, 23 figs. 1966e Aradidae in the South Australian Museum, Adelaide (Hemiptera-Heteroptera). Rec. S. Austral. Mus. 15(2): 275-307, 31 figs. 1966f Notes on Neotropical Aradidae XVI (Hemip- tera-Heteroptera). Studia Ent. 9(1-4): 515-521, 1 fig. 1966g On some Phymatidae from the Old and New Worlds (Hem. Heteroptera). EOS 42(1—2): 275-286, 2 figs. 1967a A new apterous genus and species of Aradinae from Mexico (Hemiptera-Heteroptera, Aradidae). L. A. Co. Mus. Contr. Sci. 122: 2-4, 3 figs. 1967b Notes on Australian Aradidae (Hemiptera- Heteroptera) with descriptions of new species of Calisius Stal and Glochocoris Usinger & Matsuda. Proc. R. Soc. Queensland 79(6): 71-78, 6 figs. 1967c Notes on Aradidae from the Eastern Hemi- sphere XX (Hem.-Heteroptera). EOS 42(1-4): 467- 491, 20 figs. 1967d Anew species of Neuroctenus Fieber from Tan- ganyika (Hemiptera, Aradidae). Opusc. Zool. 92: 1- 3, 3 figs. 1967e Aradidae in the Bishop Museum, Honolulu (Hemiptera-Heteroptera). Pac. Ins. 9(3): 447-479, 25 figs. 1967f Some Aradidae (Hemiptera-Heteroptera) from the Philippine, Bismarck and Solomon Islands. Ent. Medd. 35: 291-300, 8 figs. 1967g Aradidae in the South Australian Museum, Adelaide II (Hemiptera-Heteroptera). Rec. S. Aus- tral. Mus. 15(3): 513-550, 27 figs. 1967h On some Aradidae from Brasil, Argentina and Laos (Hemiptera, Heteroptera). Opusc. Zool. 100: 1-10, 13 figs. 1968a Notes on Aradidae in the U.S. National Mu- seum V, (Hemiptera: Heteroptera). Proc. U.S. Nat. Mus. 125(3651): 1-16, 8 figs. 1968b North and Central American species of Aneu- rus Curtis (Hemiptera: Aradidae). Proc. U.S. Nat. Mus. 125(3657): 1-12, 19 figs. 1968c Notes on Neotropical Aradidae XVII. Aradi- dae in the Field Museum of Natural History, Chi- cago, Illinois (Hem.-Heteroptera). Ann. Soc. Ent. France, n. s. 4(2): 279-289, 12 figs. 520 1968d Aradidae in the Bishop Museum, Honolulu, II (Hemiptera-Heteroptera). Pac. Ins. 10(2): 249-260, 19 figs. 1968e Notes on Neotropical Aradidae XVIII (He- miptera-Heteroptera). Papéis Avul. Zool., Sao Pau- lo 22(6): 47-56, 7 figs. 1968f A new species of the genus Macrocephalus Swederus from Mexico (Hemiptera: Phymatidae). Proc. Ent. Soc. Wash. 70(3): 242-245, 1 fig. 1968g Aradidae in the Bishop Museum, Honolulu II (Hemiptera: Heteroptera). Pac. Ins. 10(3—4): 575- 597, 31 figs. 1 map. 1968h Notes on Aradidae from the Eastern Hemi- sphere XIX (Hemiptera). Khumbu Himal 3(1): 27- 28. 1968i 51. Two new species of Aradidae (Hemiptera- Heteroptera) from Rennell Island. Nat. Hist. Ren- nell Isl. 5: 99-102, pl. 2. 1968} Notes on Aradidae from the Eastern Hemi- sphere XXI. Aradidae in the Field Museum of Nat- ural History, Chicago, Illinois (Hemiptera: Heter- optera). Quart. Jour. Taiwan Mus. 21(3-4): 225- 233 eeigs: 1968k Notes on Neotropical Aradidae XIX (Hemip- tera-Heteroptera). Rev. Facultad Agron. Univ. Centr. Venezuela 1: 43-56, 8 figs. 1969a Aradidae in the Bishop Museum, Honolulu IV, (Hemiptera-Heteroptera). Pac. Ins. 1 1(1): 49-70, 35 figs., 1 map. 1969b Thaicorinae, n. subf. from Thailand (Hemip- tera: Heteroptera: Piesmatidae), Pac. Ins. 1 1(3-4): 645-648, 4 figs. 1970a Two new species of the genus Aradus F., 1803, from Palaearctic Region (Hemiptera: Aradidae). Ann. Naturh. Mus. Wien 74: 201-204, 4 figs. 1971a Mezirinae of the Oriental Region and South Pacific (Hemiptera-Heteroptera: Aradidae). Pac. Ins. Monogr. 26: 1-165, 233 figs., 3 maps. 1971b Aradidae in the Bishop Museum, Honolulu V. (Supplement) (Hemiptera-Heteroptera). Pac. Ins. 12(4): 701-722, 15 figs. (1970). [issue number 4 was marked “*25 December 1970.” but the table of “Contents of Volume 12” gives the date of issue as “30 January 1971”). 1971c New Records of some Oriental Phymatidae, with a new species of Chelocoris Bianchi (Hemiptera: Heteroptera). Pac. Ins. 12(4): 883-886, 1 fig. (1970). [see note under 1971b]. 1971d Ochteridae from the Oriental and Australian Regions (Hemiptera-Heteroptera). Pac. Ins. 13(3- 4): 429-444, 23 figs., 2 maps. 197le Aradidae de la Hacienda El Limon, D. F. (He- miptera-Heteroptera). Mem. Soc. Cient. Nat. La Salle 31(89): 165-174, 3 figs. 1971f Key to American species of the genus Mezira (Hemiptera: Aradidae). Proc. Ent. Soc. Wash. 73(3): 282-292. 1972a Two new species of North American Flat Bugs PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Hemiptera: Aradidae). Bull. S. Calif. Acad. Sci. 71(2): 91-93, 5 figs. 1972b A new species of ambush bugs from Arizona (Hemiptera: Phymatidae). Bull. S. Calif. Acad. Sci. 71(2): 93-95, 1 fig. 1972c Aradidae in the Bishop Museum, Honolulu, VI (Hemiptera: Heteroptera). Pac. Ins. 14(3): 553-570, 9 figs. 1972d Mezirinae of the Oriental Region and South Pacific, Supplement (Hemiptera-Heteroptera: Ar- adidae). Pac. Ins. 14(3): 571-583, 17 figs. 1972e A new species of the genus Ochterus Latreille, 1807, from New Guinea (Hem.-Het.: Ochteridae). Pac. Ins. 14: 585-587, 3 figs. 1973a Ochteridae from western and southern Africa (Hemiptera: Heteroptera). Occ. Pap. Calif. Acad. Sci. 106: 1-9, 11 figs., 1 map. 1973b Aradidae from the Oriental Region and South Pacific (Hemiptera-Heteroptera). Pac. Ins. 15(1): 67— 83, 19 figs. 1973c Aradidae from the Oriental and Australian Regions in the Museum National d’Histoire Natu- relle, Paris (Hem.-Heteroptera.). Ann. Soc. Ent. France, n. s. 9(1): 193-209, 15 figs. 1973d Notes on Neotropical Aradidae XXI (Hem. Heteroptera). Ann. Soc. Ent. France, n. s. 9(2): 433- 439, 5 figs., pl. 10. 1973e Anew species of the genus Aneurus Curtis, 1825, from Nepal (Hemiptera: Aradidae). Ent. Rec. 85: 141-146, pl. 10. [senior author with E. Heiss]. 1973f Neotropical Aradidae in the Museum National d’Histoire Naturelle, Paris. Ann. Soc. Ent. France, n. s. 9(3): 735-746, 7 figs. 1973g On some Aradidae (Hemiptera: Heteroptera) from Nepal and Thailand. Ber. nat.-med. Ver. Inns- bruck 60: 61-77, 25 figs, 6 photos. [senior author with E. Heiss]. 1974a Aradidae from the tropical Africa and Mada- gascar in the Museum National d’Histoire Natu- relle, Paris. (Hem. Heteroptera). Ann. Soc. Ent. France, n. s. 10(1): 59-77, 5 figs. 1974b A new genus of Calisiinae from Kameroon (Hem., Aradidae). EOS 48: 295-299, 2 figs. 1974c Aradidae in the Rijksmuseum van Natuurlijke Historie, Leiden (Hemiptera-Heteroptera). Zool. Meded. 48(21): 233-247, 11 figs. 1975a Neotropical Aradidae in the collections of the California Academy of Sciences, San Francisco (He- miptera-Heteroptera). Occ. Papers Cal. Acad. Sci. 122: 1-28, 14 figs. 1975b Notes on the Neotropical Aradidae, XX (Hem. Heteroptera). EOS 49: 219-224, 5 figs. (1973). 1975c Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen Museums in Basil. Hemiptera: Fam. Aradidae. Ent. Basil. 1: 93-112, 25 figs., 8 photos. [senior author with E. Heiss]. 1976a Onsome Aradidae from the Old and New World VOLUME 97, NUMBER 3 (Hemiptera-Heteroptera). Zool. Scripta 5: 67-78, 23 figs. 1976b Three new species of Aradidae from Japan and Taiwan (Hemiptera-Heteroptera). Dtsch. Ent. Zeitschr. n. f., 23 (1-3): 221-228, 16 figs. [senior author with E. Heiss]. 1976c On new and little known species of Aradidae, I (Hemiptera, Heteroptera). Zeitschr. Arbgem. Os- terr. Ent. 28(1-3): 39-44, 10 figs., 3 photos. [senior author with E. Heiss]. 1976d Notes on Aradidae from China and India (He- miptera-Heteroptera). Pac. Ins. 17(1): 87-90, 4 figs. 1976e A new genus and seven new species of Neo- tropical Aradidae (Hemiptera-Heteroptera). Rev. Brasil. Biol. 36: 735-743, 12 figs. 1977a Anew species of the genus Over/aetiella Schou- teden from Sumatra (Hemiptera: Aradidae). Proc. Ent. Soc. Wash. 79(3): 354-357, 3 figs. 1977b A new genus and a new species of the Carven- tinae from Surinam (Hemiptera, Aradidae). Zool. Meded. 52(1): 1-6, 4 figs., 1 pl. [senior author with P. H. van Doesburg]. 1977c New species of Aradidae from the Philippines and Borneo (Hemiptera: Heteroptera). Pac. Ins. 1 7(2— 3): 261-163, 5 figs. 1977d Aradidae from the Oriental and Australian Regions in the collections of the California Academy of Sciences, San Francisco (Hemiptera: Heterop- tera). Trans. Amer. Ent. Soc. 103: 603-622, 33 figs. 1977e Five new Aradidae from the Oriental Region (Insecta: Heteroptera). Ber. nat.-med. Ver. Inns- bruck 64: 97-106, 13 figs. [senior author with E. Heiss]. 1978a Two new species of American Aradidae (He- miptera). Proc. Ent. Soc. Wash. 80(2): 228-233, 7 figs. 1978b Aradus frigidus Kiritshenko, 1913, and Aradus italicus Kormilev, 1970 (Hemiptera: Aradidae). Ann. Naturh. Mus. Wien 81: 505-506, 6 figs. 1978c Aradidae in the Bishop Museum, Honolulu VII (Hemiptera: Heteroptera). Pac. Ins. 18(1—2): 53-60, 13 figs. 1978d Two new species of Aradidae from Brasil (He- miptera, Heteroptera). Zeitschr. Arbgem. Osterr. Ent. 30(1-2): 29-32, 4 figs. [senior author with E. Heiss]. 1978e On some Aradidae from the Oriental and Aus- tralian Regions (Hemiptera). Philipp. Jour. Sci. 105(4): 235-254, 12 figs. (1976). 1978f A new species of the genus Neasterocoris Usin- ger and Matsuda from Ecuador (Hemiptera: Arad- idae). Rev. Brasil. Ent. 22(3—4): 199-201, 2 figs. 1979a A new apterous species of Aradidae from Ke- nya (Hemiptera). Pan-Pac. Ent. 55(1): 57-60, 2 figs. 1979b A new species of the genus Aradus F., 1803, from Alaska (Insecta: Heteroptera). Ber. nat.-med. Ver. Innsbruck 66: 47-52, 4 figs., 8 photos. [senior author with E. Heiss]. 1979c New Aradidae from Ceylon and Malaya in the 521 British Museum (N.H.) (Heteroptera). Orient. Ins. 13(1-2): 155-162, 20 figs. [senior author with E. Heiss]. 1979d Four new species and lectotype designation of some neotropical Aradidae (Insecta, Heteroptera). Ent. Arb. Mus. Frey 28: 93-118, 35 figs. [senior author with E. Heiss]. 1979e Dos nuevas especies y algunas especies poco conocidas de Aradidae de Brasil (Hemiptera). An. Inst. Biol. Univ. Nac. Auton Mexico, 50, Ser. Zool. (1): 341-345, 8 figs. 1980a Homonymy in the Aradidae (Hemiptera). Pac. Ins. 22(3—4): 328. 1980b Notes on American Aradinae (Hemiptera: Ar- adidae). Proc. Ent. Soc. Wash. 82(1): 99-107, 9 figs. 1980c Sobre especies Argentinas del genero Aneurus Curtis, 1825 (Hemiptera: Aradidae). Acta Zool. Lil- loana 36(1): 53-56, 10 figs. 1980d Three new species of Aradidae from Mexico (Hemiptera). Proc. Ent. Soc. Wash. 82(4): 695-699, 7 figs. 1980e A new species of the genus Daulocoris Usinger and Matsuda, 1959, from Sumatra (Hemiptera: Ar- adidae). Sociobiology 5(1): 21-24, 3 figs. 198la Aradidae (Hemiptera) from Ceylon. Ceylon Jour. Sci. (Biol. Sci.) 14(1-2): 122-131, 9 figs. 1981b A new micropterous species of Carventus Stal from Chile (Hemiptera: Aradidae). Proc. Ent. Soc. Wash. 83(2): 296-299, 3 figs. 1981c Onsome Neotropical species of the genus Mac- rocephalus (Hemiptera: Phymatidae). Sociobiology 6(2): 214-220, 2 figs. 1982a Six new species of Neotropical Aradidae (He- miptera). Proc. Ent. Soc. Wash., 84(3): 480-488, 14 figs. 1982b Records and Descriptions of North American and Oriental Aradidae (Hemiptera). Wasmann Jour. Biol. 40(1-2): 1-17, 24 figs. 1982c On Mezira granulata (Say) group (Hemiptera: Aradidae). Jour. Nat. Hist. 16: 775-779, 6 figs. 1982d Five new species of the Neotropical Aradidae (Hemiptera). Rev. Brasil. Ent. 26(3-4): 269-275, 9 figs. 1982e Onsome Aradidae in the collections of the Uni- versity of Georgia, Athens, Georgia, U.S.A. (He- miptera). Jour. Ga. Ent. Soc. 17(3): 333-337, 5 figs. 1982f Records and descriptions of Central American Aradidae (Hemiptera). Wasmann Jour. Biol. 40(1- 2): 26-44, 21 figs. 1982¢ A new species of Neuroctenus from nests of termites (Hemiptera: Aradidae). Sociobiology 7: 25— 28, 4 figs. 1983a New oriental aradid bugs in the collection of the British Museum (Natural History) (Insecta: He- miptera). Jour. Nat. Hist. 17: 437-469, 34 figs. 1983b A new species of the genus Phymata Latreille from Mexico. Jour. Ga. Ent. Soc. 18(1): 138-141, 5 figs. 522 1983c Onthe homonymy of Hybocoris Kormilev, 1982 (Hemiptera: Aradidae). Proc. Ent. Soc. Wash. 85(4): 690. 1983d Two new species of Neotropical Aradidae and notes on the genus G/yptocoris Harris and Drake (Hemiptera). Proc. Ent. Soc. Wash. 85(4): 818-821, 5 figs. 1984a A new species of the genus Chinessa Usinger and Matsuda, 1959, from Papua New Guinea (He- miptera: Aradidae), Pan-Pac. Ent. 60(2): 76-78, 2 figs. 1984b Keys to the genera and descriptions of new taxa of macrocephaline ambush bugs (Heteroptera: Phy- matidae). Jour. Nat. Hist. 18: 623-637, 11 figs. 1985a On some aradid bugs described by Francis Walker (Insecta: Hemiptera). Jour. Nat. Hist. 19: 1045-1047, 2 figs. 1985b Replacement name for Carventus parvus Kor- milev (Hem. Aradidae). Ent. Month. Mag. 121: 98. 1985c Anew species of the genus Notapictinus Usinger and Matsuda from Venezuela (Hemiptera: Aradi- dae). Rev. Brasil. Ent. 29(1): 153-154, 2 figs. 1986a The first find of a Mesozoic aradid bug fossil in Northeast Siberia (Hemiptera: Aradidae). Jour. Nat. Hist. 20: 279-282, 3 figs., 1 table. [senior au- thor with Y. A. Popov]. 1986b Notes on Phymatidae (Heteroptera). Zool. Meded. 60(2): 113-127, 11 figs. [senior author with P. H. van Doesburg]. 1986c Notes on Asiatic and African Macrocephalinae (Heteroptera: Phymatidae) with description of a new genus. Zool. Meded. 60(18): 273-276, 3 figs. [senior author with P. H. van Doesburg]. 1986d Notes on American Macrocephalinae with de- scriptions of two new species (Hemiptera: Phyma- tidae). Pan-Pac. Ent. 62(4): 303-310, 4 figs. 1986e Two new genera and seven new species of Ar- adidae (Heteroptera). Jour. N. Y. Ent. Soc. 94(2): 249-261, 10 figs. 1986f Two new species of Macrocephalinae from Mexico (Hemiptera, Phymatidae). Rev. Brasil. Ent. 30(2): 299-302, 6 figs. 1987a Notes on North and Central American Lo- phoscutus spp. (Hemiptera: Phymatidae, Macroce- phalinae). Proc. Ent. Soc. Wash. 89(4): 701-705, 2 figs. 1987b On African Macrocephalinae (Hemiptera, Phy- matidae). Rev. France Ent. n. s. 9(4): 159-161, 1 fig. 1987c Flat bugs of the world: A synonymic list (Het- eroptera: Aradidae). Entomography 5: 1-246, 3 figs. [senior author with R. C. Froeschner]. 1988a Notes on Agreuocoris Handlirsch (Hemiptera: Phymatidae) Ann. Soc. Ent. France, n. s. 24(4): 115- 116. 1988b On some Phymatidae of the Natural History Museum Basel (Hemiptera). Ent. Basil. 12: 63-76, 8 figs. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1988c Four new species of the genus Lophoscutus Kor- milev, 1951, from Latin America (Hemiptera: Phy- matidae, Macrocephalinae). EOS 64: 141-146, 5 figs. 1989a A new species of the genus Lophoscutus Kor- milev from Mexico (Hemiptera: Phymatidae: Ma- crocephalinae). Pan-Pac. Ent. 65(4): 451-453, 1 fig. 1989b Corrections for two ambush bugs described from Pakistan (Hemiptera: Phymatidae). Bishop Mus. Occ. Papers 29: 149-150. 1989c The preliminary study of the Phymatidae in Inner Mongolia, China (Hemiptera: Phymatidae). Zool. Res. 10(4): 341-347, 15 figs. [second author with Nonnaizab and B. Qi]. 1989d A new species of the genus Lophoscutus Kor- milev from the Dominican Republic (Hemiptera, Phymatidae). Mitt. Miinch. Ent. Ges. 79(4): 61-63, 4 figs. 1990a Phymatidae or ambush bugs of the world: A synonymic list with keys to species, except Lopho- scutus and Phymata (Hemiptera). Entomography. 6: 1-75, 8 figs. [junior author with R. C. Froeschner]. 1990b Case 2700. Carcinochelis Fieber, 1861 (Insecta, Heteroptera): proposed designation of Carchochelis alutaceus Handlirsch, 1897, as the type species. Bull. Zool. Nomencl. 47(1) March 1990: 30-31. [second author with Richard C. Froeschner]. 1990c A new species of the genus Chelocoris from Vietnam (Hemiptera: Phymatidae: Carcinocori- nae). Bishop Mus. Occas. Papers 30: 298-300. 1990d Two new species of the genus Lophoscutus Kor- milev, 1951 (Hemiptera, Phymatidae). Rev. Brasil. Ent. 34: 221-225. 1990e A new homonym in the Carventinae (Hemip- tera: Aradidae). Pan-Pac. Ent. 66: 325. 1990f Notes on American Phymatidae, with descrip- tion of a new species of Phymata Latreille. Anales Inst. Biol. Univ. Nac. Auton. Mexico, Ser. Zool. 61: 99-106. 1991a Notes on Caribbean Phymatidae (Hemiptera: Heteroptera). Zool. Meded. 65: 277-285. [senior au- thor with P. H. van Doesburg]. 1992a A New Homonym in the Mezirinae (Hemip- tera: Aradidae). Jour. N. Y. Ent. Soc. 100(1): 184. 1992b Notes on Macrocephalinae (Heteroptera: Phy- matidae). Zool. Meded. 66: 417-421. [senior author with P. H. van Doesburg]. List OF JOURNALS CITED BY ABBREVIATIONS Acta Sci. Inst. Inv. San Miguel.—Acta Scientifica de los Institutos de Investi- gacion de San Miguel, Province de Bue- nos Aires. Acta Zool. Lilloana.—Acta Zoologica Lil- loana. VOLUME 97, NUMBER 3 Amer. Mus. Nov.—American Museum Novitates. An. Inst. Biol.—Anales Instituto del Biolo- gia, Universidad Nacional Autonoma de Mexico, serie Zoologica. An. Soc. Cien. Argent.—Anales de la So- ciedad Cientifica Argentina. Ann. Ent. Soc. Amer.—Annals of the En- tomological Society of America. Ann. Mag. Nat. Hist.—Annals and Maga- zine of Natural History. Ann. Naturh. Hofsmus. Wien.—Annalen des Naturhistorischen Museums in Wien. Ann. Soc. Ent. France.—Annales de la So- ciété entomologique de France, Nouvelle série. Ann. Mus. Serbiae Merid., Skoplje.—An- nales Musei Serbiae Meridionalis, Skop- le (Yugoslavia). Ark Zool.—Arkiv f6r Zoologi. Ber. nat.-med. Ver. Innsbruck.— Berichte des naturwissenschaftlich-medizinischen Vereins in Innsbruck. Bishop Mus. Occ. Papers.— Bishop Muse- um Occasional Papers. Bull. S. Cal. Acad. Sci.—Bulletin of the Southern California Academy of Sci- ences. Bull. Soc. Ent. Suisse. — Bulletin Société En- tomologique Suisse. Bull. Soc. Sci. Skoplje.— Bulletin de la So- ciété Scientifique de Skoplje (Yugosla- via). Bull. Zool. Nomencl.— Bulletin of Zoolog- ical Nomenclature. Ceylon Jour. Sci.—Ceylon Journal of Sci- ence. Com. Inst. Nac. Inv. Cien. Nac.—Comun- icaciones del Instituto Nacional de In- vestigacion de las Ciencias Naturales, Ciencias Zoologicas. Dscht. Ent. Zeitschr.—Deutsche entomo- logische Zeitschrift. Dusenia. — Dusenia. Ent. Arb. Mus. Frey.—Entomologische Ar- beiten Museum G. Frey. Ent. Basil.—Entomologica Basiliensia. Ent. Medd.—Entomologiske Meddelelser. 523 Ent. Month. Mag.—Entomologist’s Month- ly Magazine. Ent. Rec.—Entomologist’s Record. Entomography.— Entomography EOS.— Official title for Revista Espanola de Entomologia. Jour. Ent. Soc. Queensland.—Journal of the Entomological Society of Queensland. Jour. Ent. Soc. S. Africa.—Journal of the Entomological Society of Southern Afri- ca. Jour. Ga. Ent. Soc.—Journal of the Georgia Entomological Society. Jour. Kans. Ent. Soc.—Journal of the Kan- sas Entomological Society. Jour. N. Y. Ent. Soc.—Journal of the New York Entomological Society. Jour. Nat. Hist.—Journal of Natural His- tory. Khumbu Himal.—published by Universi- tatsverlag Wagner, Innsbruck-Miinchen. L. A. Co. Mus. Contr. Sci.—Los Angeles County Museum Contributions to Sci- ence. Mem. Soc. Cien. Nat. La Salle. —Memorias de la Sociedad de Ciencias Naturales La Salle. Mis. Est. Patol. Reg. Argent.— Misio de Es- tudios de Patologia Regional Argentina. Mitt. Miinch. Ent. Ges.— Mitteilungen der Miinchner Entomologischen Gesell- schaft. Mushi.— Mushi [official title of publication of the Fukuoka Entomological Society). Nat. Hist. Rennell Isl.—The Natural His- tory of Rennell Island, British Solomon Islands. Notas Mus. La Plata, Zool. — Notas del Mu- seo de la Plata, Zoologica. Occ. Pap. Cal. Acad. Sci.—Occasional Pa- pers of the California Academy of Sci- ences. Opusc. Ent.—Opuscula Entomologica. Orient. Ins.—Oriental Insects. Pac. Ins.—Pacific Insects. Pac. Ins. Monogr.—Pacific Insect Mono- graphs. Pan-Pac. Ent.—Pan-Pacific Entomologist. 524 Papéis Avul. Zool.— Papéis Avulso de Zoo- logia, Sao Paulo. Philipp. Jour. Sci.—The Philippine Journal of Science. Posebna Izdanja. — Posebna Izdanja, Srpska Akademija Nauka, Belgrade [vol. 136] (Monographii Srpska Akademeiji Nauka [vol. 35] or Monographiarum Regia Ac- ademia Serbica [vol. 35]. Proc. Ent. Soc. Wash.— Proceedings of the Entomological Society of Washington. Proc. R. Soc. Queensland.— Proceedings of the Royal Society of Queensland. Proc. U.S. Nat. Mus.—Proceedings of the United States National Museum. Psyche— Psyche, a Journal of Entomology. Publ. Univ. Cochabamba.— Publicaciones de la Universidad de Cochabamba, Folia Universitaria. Quart. Jour. Taiwan Mus.— Quarterly Jour- nal of the Taiwan Museum. Rec. S. Austral. Mus.— Records of the South Australian Museum. Rev. Brasil. Biol.—Revista Brasileira de Biologia. Rev. Brasil. Ent.— Revista Brasileira de En- tomologia. Rev. Chilena Ent.— Revista Chilena de En- tomologia. Rev. Ecuat. Ent. Parasit.— Revista Ecuato- riana de Entomologia Y Parasitologia. Rev. Ent.— Revista de Entomologia, Rio de Janeiro. Rev. Facultad Agron. Univ. Centr. Vene- zuela.— Revista de la Facultad de Agron- omia, Universidad Central de Venezuela. Rev. France Ent.—Revue Francais d’En- tomologie. Rev. Inst. Nac. Inv. Cien. Nat.— Revista del Instituto Nacional de Investigacion de las Ciencias Naturales. Rev. Soc. Ent. Argent.—Revista de la So- ciedad Entomologica Argentina. Rev. Soc. Uruguaya Ent.— Revista de la So- ciedad Uruguaya de Entomologia. Roy. Serbian Acad. Sci.—Royal Serbian Academy of Sciences, Special Issues, Nat- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ural History & Mathematical Papers, Ohridski Zbornik. Sociobio. — Sociobiology. Stud. Ent.—Studia Entomologica. Trans. Amer. Ent. Soc.— Transactions of the American Entomological Society. Verh. Naturf. Ges. Basel.— Verhandlungen der Naturforschenden Gesellschaft in Ba- sel. Wasmann Jour. Biol.—The Wasmann Journal of Biology. Zeitschr. Arbgem. Osterr. Ent.— Zeitschrift der Arbeitsgemeinschaft Osterreichisher Entomologen. Zool. Meded.—Zoologische Mededelingen, Riksmuseum van Natuurliyke Historie te Leiden. Zool. Res.— Zoological Research. Zool. Scripta.— Zoologica Scripta. List OF NAMES PROPOSED Order Heteroptera Family-group Names Aradidae Llaimacorini Kormilev and Froeschner, 1987c> 7. and .197 (misspelled ‘““Llaimocorini” on p. 197) Tretocorini Kormilev and Froeschner, 1987c: 6 and 93 Coreidae Merocorini Kormilev, 1954a: 156 Meropachyni [sic] Kormilev, 1954a: 156 and 159 Spathophorini Kormilev, 1954a: 156 and 174 Lygaeidae Robinsonocorini Kormilev, 1952f: 9 Piesmatidae Thaicorinae Kormilev, 1969b: 645 Thaumastotheriidae Discocorinae Kormilev, 1955g: 7 Vianaididae Kormilev, 1955j: 466 Genus-group Names Alydidae Bactrocoris Kormilev, 1953b: 55 VOLUME 97, NUMBER 3 Aradidae Aegisocoris Kormilev, 1967g: 521 Aellocoris Kormilev, 1964c: 53 Aneurus (Aneurillus) Kormilev, 1968d: 259 Apaniocoris Kormilev, 1983a: 443 Aparilocoris Kormilev, 1983c: 690 [new name for preoccupied Hybocoris Kormilev, 1982] Apterocoris Kormilev and Doesburg, 1977b: 1 Aradiolus Kormilev, 1967a: 2 Arbanatus Kormilev, 1951: 180 Argocoris Kormilev, 1967g: 519 [preoc- cupied, see Pseudoargocoris Kormi- lev, 1992: 184] Aspisocoris Kormilev, 1967g: 515 Atactocoris Kormilev, 1964d: 114 Axapisocoris Kormilev and Heiss, 1979c: tert) Biroana Kormilev, 1957a: 395 Caecicoris Kormilev, 1957a: 398 Chapadia Kormilev, 1960a: 42 Chelysosoma Kormilev, 1956d: 283 Chrysodaspis Kormilev, 1971a: 99 Clavicornia Kormilev, 1960f: 167 Delnocoris Kormilev, 1982f: 41 Dihybogaster Kormilev, 1953d: 230 [in generic heading misspelled “‘Dyhi- bogaster’’| Dimifaia Kormilev, 1971a: 96 Diphyllonotus Kormilev, 1959b: 61 [““Dyhibogaster’ —see Dihybogaster] Drakeida Kormilev, 1958b: 89 Erineocoris Kormilev, 1960f: 163 Eunotoplocoris Kormilev, 1957g: 4 Euricoris Kormilev, 1957a: 393 Forficulassa Kormilev, 1960a: 40 Froeschnerissa Kormilev, 1986e: 253 Glyptoaptera Kormilev, 1965b: 18 Glyptomorpha Kormilev, 1977d: 412 [preoccupied, see Morphocoris Kor- milev, 1980a: 328] Gnostocoris Kormilev, 1967e: 449 Halaszfya Kormilev, 1960e: 210 Heissia Kormilev, 1986e: 251 Hybocoris Kormilev, 1982b: 7 [preoccu- 525 pied, see Aparilocoris Kormilev, 1983c: 690 Kaulocoris Kormilev, 1971b: 718 Kelaino Kormilev, 1963a: 451 Kema Kormilev, 1955h: 141 Kiritshenkiana Kormilev, 1976a: 71 Kiritshenkiessa Kormilev, 1971a: 10 Leiocoris Kormilev, 1971a: 112 Leurocoris Kormilev, 1971b: 720 Libiocoris Kormilev, 1957a: 390 Limonocoris Kormilev, 1971le: 166 Llaimacoris Kormilev, 1964d: 117 Magdalenia Kormilev, 1983a: 445 Mapiri Kormilev, 1959b: 69 Micromezira Kormilev, 1967c: 488 Morphocoris Kormilev, 1980a: 328 [new name for the preoccupied G/ypto- morpha Kormilev, 1977d: 412] Odontonotus Kormilev, 1955c: 34 Paracalisiopsis Kormilev, 1963b: 602 and 605 Paracalisius Kormilev, 1974b: 296 Paracarventus Kormilev, 1964c: 42 Parahesus Kormilev, 1960a: 46 Paramezira Kormilev, 1974c: 246 Parapictinus Kormilev, 1956d: 291 Pararhombocoris Kormilev, 1990e: 325 [new name for the preoccupied Rhombocoris Kormilev, 1965b: 18] Parartabanus Kormilev, 1972d: 573 Pelecoris Kormilev, 1971a: 97 Probatoceps Kormilev, 1965b: 21 Pseudoargocoris Kormilev, 1992a: 184 [new name for preoccupied Argocoris Kormilev, 1967: 519] Pseudopictinus Kormilev, 1966f: 517 Rhombocoris Kormilev, 1965b: 17 [pre- occupied, see Pararhombocoris Kor- milev, 1965b: 18] Rhynchomirus Kormilev, 1976e: 742 Rustem Kormilev, 1957e: 39 Santaremia Kormilev, 1960a: 44 Scirrhocoris Kormilev, 1965b: 26 Scironocoris Kormilev, 1957a: 401 Usingerida Kormilev, 1955h: 142 Usumbaraia Kormilev, 1956c: 253 Zemira Kormilev, 1971la: 31 [preoccu- 526 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pied, see Zimera Kormilev, 1980a: 328] Zimera Kormilev, 1980a: 328 [new name for preoccupied Zemira Kormilev, 1971a: 31] Coreidae Paralycambes Kormilev, 1954a: 174 and 180 Colobathristidae Carvalhoia Kormilev, 1951a: 67 Labradoria Kormilev, 1951a: 80 Neocolobathristes Kormilev, 1951a: 70 Paraelopura Kormilev, 1953d: 290 Perudella Kormilev, 1949a: 170 Lygaeidae Karamania Kormilev, 1938a: 168 Riggiella Kormilev, 1949d: 4 Robinsonocoris Kormilev, 1952f: 9 Pentatomidae Daimonocoris Kormilev, 1951h: 37 Harpagogaster Kormilev, 1957d: 48 Neoadoxoplatys Kormilev, 1956b: 4 Neoleprosoma Kormilev, 1952b: 213 Ornithosoma Kormilev, 1957d: 52 Prionotocoris Kormilev, 1955e: 7 Ramosiana Kormilev, 1950c: 340 Willinerinia Kormilev, 1950d: 496 Phymatidae Agdistocoris Kormilev, 1962e: 356 Bakerinia Kormilev, 1962c: 9 Phymata (Euryphymata) Kormilevy, 1962d: 308, 324 and 452 Hoberlandtiana Kormilev and Doesburg, 1986c: 273 Kelainocoris Kormilev, 1963c: 215 and 22 Macrocephalus (Lophoscutus) Kormilev, 1951g: 101 Phymata (Neoanthylla) Kormilev, 1951g: 56 Phymata (Neophymata) Kormilev, 1962d: 308, 324 and 450 Parabotha Kormilev, 1984b: 629 Paraphymata Kormilev, 1962d: 326 and 464 Phymatispa Kormilev, 1951g: 83 Phymatocoris Kormilev and Doesburg, 1991la: 279 Piesmatidae Thaicoris Kormilev, 1969b: 646 Thaumastotheriidae Discocoris Kormilev, 1955g: 8 Vianaididae Vianaida Kormilev, 1955j: 468 Species-group Names Acanthosomatidae Sniploa shajovsoii Kormilev, 1952a: 52 Alydidae Bactrocoris plaumanni Kormilev, 1953b: 56 Bactrophya argentina Kormilev, 1953b: 53 Cydamus bolivianus Kormilev, 1953b: 59 Cydamus delpontei Kormilev, 1953b: 62 Cydamus lizeri Kormilev, 1953b: 64 Cydamus minor Kormilev, 1953b: 61 Cydamus seai Kormilev, 1953b: 65 Aradidae Acantharadus lobulatus Kormilev, 1953c: 337 Acaraptera (Lissaptera) denticeps Kor- milev, 1966e: 306 Acaraptera (Nesiaptera) denticulata Kor- milev, 1968g: 589 Acaraptera (Acaraptera) dimorpha Kor- milev, 1966e: 303 Acaraptera (Nesiaptera) gibbosa Kormi- lev, 1968g: 590 Acaraptera (Acaraptera) minuta Kormi- lev, 1966e: 305 Acaraptera (Nesiaptera) ovata Kormilev, 1968g: 588 Acaraptera (Nesiaptera) rotundata Kor- milev, 1968g: 589 Acaraptera (Acaraptera) solomonensis Kormilev, 1971b: 716 Acaraptera (Nesiaptera) tuberculata Kor- milev, 1968g: 590 Acaricoris austeris Drake and Kormilev, 1958a Acaricoris barroanus Drake and Kormi- lev, 1958a: 244 Acaricoris clausus Drake and Kormilev, 1958: 244 Acaricoris dureti Kormilev, 1953a: 125 VOLUME 97, NUMBER 3 Acaricoris haitiensis Kormilev, 1968a: 2 Acoryphocoris antennatus Kormilev, 1971a: 141 Acoryphocoris angusticornis Kormilev, 1971a: 140 Acoryphocoris brevicornis Kormilev, 197 1a: 141 Acoryphocoris denticulatus Kormilev, 1971a: 140 Acoryphocoris duodecimus Kormilev, 1983a: 466 Acoryphocoris minor Kormilev, 1983a: 465 Acoryphocoris oviventris Kormilev, 197 1a: 139 Aegisocoris granulatus Kormilev, 1967g: 533. Aellocoris breviceps Kormilev, 1964c: 45 Aellocoris undulatus Kormilev, 1964c: 45 Aglaocoris drakei Kormilev, 1968a: 5 Aneurus (Aneura) aibonitensis Kormilev, 1968b: 4 Aneurus ampliatus Kormilev, 1976a: 71 Aneurus angulatus Kormilev, 1965b: 14 Aneurus assamensis Kormilev, 1977d: 606 Aneurus aterrimus Kormilev, 1982a: 482 Aneurus barberi Kormilev, 1960e: 218 Aneurus (Iralunelus) bergi Kormilev, 1980: 55 Aneurus bhutanensis Kormilev and Heiss, 1975c: 95 Aneurus bimaculatus Kormilev and Heiss, 1977e: 97 Aneurus bishopi Kormilev, 1872c: 556 Aneurus bispinosus Kormilev, 1960c: 93 Aneurus bolivianus Kormilev, 1960e: 217 Aneurus (Aneurillua) borneensis Kormi- lev, 1971b: 710 Aneurus bosqui Kormilev, 1967h: 4 Aneurus brailovskyi Kormilev, 1982a: 480 Aneurus brevirostris Kormilev, 1967a: 70 Aneurus burmensis Kormilev, 1976a: 70 Aneurus bucki Kormilev, 1965d: 2 Aneurus carioca Kormilev, 1968e: 48 Aneurus (Aneurus) championi Kormilev, 1968b: 9 Aneurus cheesmani Kormilev, 1967c: 471 27 Aneurus (Aneurillus) consimilis Kormi- lev, 1982b: 4 Aneurus (Iralunelus) costaricensis Kor- milev, 1982f: 30 Aneurus crenulatus Kormilev, 1957e: 45 Aneurus doesburgi Kormilev, 1974c: 234 Aneurus equatoriensis Kormilev, 1973d: 433 Aneurus fritzi Kormilev, 1960e: 218 Aneurus (Aneurus) froeschneri Kormilev, 1968b: 7 Aneurus (Aneurillus) gracilis Kormilev, 1973c: 558 Aneurus granuliger Kormilev, 1978c: 53 Aneurus (Aneurus) gressitti Kormilev, 1968d: 254 Aneurus (Aneurus) hainanensis Kormi- lev, 1968d: 252 Aneurus haitiensis Kormilev, 1968a: 6 Aneurus herediensis Kormilev, 1982f: 27 Aneurus guanacastensis Kormilev, 1982f: 29 Aneurus insularis Kormilev, 1971b: 709 Aneurus krombeini Kormilev, 1981a: 123 Aneurus (Iralunelus) longicornis Kormi- lev, 1982f: 32 Aneurus mexicanus Kormilev, 1980d: 695 Aneurus (Aneurus) micronesicus com- munis Kormilev, 1968d: 257 Aneurus (Iralunelus) monrosi Kormilev, 1980c: 54 Aneurus mysorensis Kormilev, 1977d: 604 Aneurus nasutus Kormilev, 1966d: 7 Aneurus nepalensis Kormilev and Heiss, 1973: 143 Aneurus nipponicus Kormilev and Heiss, 1976b: 224 Aneurus nitidulus Kormilev, 19551: 189 Aneurus oviventris Kormilev and Heiss, 1976b: 224 Aneurus papuasicus Kormilev, 1967c: 474 Aneurus (Aneurus) pisoniae Kormilev, 1968b: 6 Aneurus plaumanni Kormilev, 1965d: 1 Aneurus (Aneurus) proximus Kormilevy, 1968d: 257 Aneurus pusillus Kormilev, 1968b: 3 528 Aneurus pygmaeus Kormilev, 1966d: 6 {neurus robustus Kormilev, 1957e: 44 Aneurus (Aneurus) rugosiceps Kormilev, 1968d: 256 Aneurus (Aneurus) schuhi Kormilev, 1982d: 269 Aneurus (Aneurus) solomonensis Kormi- lev, 1968d: 258 Aneurus striatus Kormilev, 1972c: 557 Aneurus (Aneurus) sublobatus Kormilev, 1968d: 254 Aneurus subsimilis Kormilev, 1968d: 255 Aneurus superbus Kormilev, 1967c: Aneurus sutteri Kormilev, 1953c: 334 Aneurus tainguensis Kormilev, 1971b: TO Aneurus taiwanensis Kormilev, 1972c: SES) Aneurus tonkinensis Kormilev, 1973c: 193 Aneurus vauriei Kormilev, 1864g: 156 Aneurus (Aneurus) viethamensis Kormi- lev, 1968d: 252 Apaniocoris micropterus Kormilev, 1983a: 444 Aparilocoris venezuelanus Kormilev, 1983d: 818 Aphelocoris alatus Kormilev, 1971a: 136 Aphelocoris carinatus Kormilev, 1967c: 485 Aphelocoris confusus Kormilev, 1971a: 136 Aphelocoris minutissimus Kormilev, 1983a: 464 Aphelocoris quadridentatus Kormilev, 1971a: 134 Aphleboderrhis alata Kormilev, 1960e: 216 Aphyseteres nausutus Kormilev, 1986e: 254 Apteraradus rossi Kormilev, 1977d: 610 Apteraradus similis Kormilev, 1976a: 72 Apteraradus vietnamensis Kormilev, 1968g: 592 Apterocoris surinamensis Kormilev and Doesburg, 1977b: 3 Aradiolus chemsaki Kormilev, 1980b: 100 Aradiolus paradoxus Kormilev, 1967:a: 4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Aradus alaskanus Kormilev and Heiss, 1979b: 47 Aradus angusticornis Kormilev, 1974a: 61 Aradus (Aradus) barberi Kormilev, 1966d: 4 Aradus (Aradus) brevicornis Kormilev, 1980b: 105 Aradus (Aradus) burmensis Kormilev, 1976a: 69 Aradus canariensis Kormilev, 1954b: 204 Aradus capensis Kormilev, 1956c: 250 Aradus (Aradus) carolinensis Kormilev, 1964f: 476 Aradus coloradensis Kormilev, 1964f: 477 Aradus creticus Kormilevy and Popov, 1986a: 280 Aradus (Aradus) denticulatus Kormilev, 1982e: 334 Aradus erraticus Kormilev, 1966e: 280 Aradus esakii Kormilev and Heiss, 1976b: 224 Aradus foliaceus Kormilev, 1957e: 38 Aradus (Aradus) fuscicornis Kormiley, 1966e: 278 Aradus (Aradus) granuliger Kormilev, 1980b: 103 Aradus italicus Kormilev, 1970a: 201 Aradus kiritshenkoi Kormilev, 1970a: 203 Aradus lawrencei Kormilev, 1966b: 27 Aradus malaisei Kormilev, 1976a: 67 Aradus maroccanus Kormilev, 1867c: 468 Aradus nevadensis Kormilev, 1972a: 91 Aradus occidentalis Kormilev, 1980b: 106 Aradus (Quilnus) oregonicus Kormilev, 1978a: 229 Aradus ovatus Kormilev, 1966d: 3 Aradus (Aradus) oviventris Kormilev, 1966b: 26 Aradus nipponicus Kormilev, 1955c: 33 Aradus saileri Kormilev, 1966d: 2 Aradus sinensis Kormilev, 19551: 177 Aradus (Aradus) testaceus Kormilevy, 1980b: 104 Aradus tonkinensis Kormilev and Heiss, 1976c: 39 Aradus (Quilnus) usingeri Kormiley, 1978a: 230 VOLUME 97, NUMBER 3 Aradus (Aradus) vietnamensis Kormiley, 1967e: 453 Arbanatus abdominalis Kormilev, 197 La: 147 Arbanatus affinis Kormilev and Heiss, 1975c: 100 Arbanatus angustus Kormilev, 1972d: 579 Arbanatus antennatus Kormilev, 1972d: 578 Arbanatus brachypterus Kormilev, 197 1a: 157 Arbanatus castaneus Kormilev, 1971a: 150 Arbanatus distinctus Kormilev, 1971la: 154 Arbanatus elegantulus Kormilev, 1971a; 151 Arbanatus grandis Kormilev, 1978c: 59 Arbanatus gressitti Kormilev, 197 1a: 149 Arbanatus inermis Kormilev, 19551: 181 Arbanatus longicornis Kormilev, 197 1a: 146 Arbanatus longirostris Kormilev, 197 1a: 152 Arbanatus longiscutum Kormilev, 1971a: 147 Arbanatus longulus Kormilev, 1971a: 146 Arbanatus minutus Kormilev, 1971a: 153 Arbanatus parallelus gracilis Kormilev, 1971la: 152 Arbanatus philippinensis Kormilev, 1971a: 144 Arbanatus piliferus Kormilev, 1971a: 155 Arbanatus pilosulus Kormilev, 197 1a: 156 Arbanatus polynesicus Kormilev, 1971a: 155 Arbanatus robustus Kormilev, 1971a: 156 Arbanatus scabrosus Kormilev, 1971la: 153 Arbanatus simplex Kormilev, 197 1a: 148 Arbanatus simulans Kormilev, 197 1a: 148 Arbanatus subparallelus Kormilev, 197 1a: 149 Arbanatus tahitiensis Kormilev, 197 1a: 156 Arbanatus tonkinensis Kormilev, 1973c: 208 Arbanatus tricolor Kormilev, 1971a: 150 Argocoris grossi Kormilev, 1967g: 521 529 Arictus acuminatus Kormilev, 1971a: 108 Arictus brachycephalus Kormilev, 1968): 230 Arictus ceylonensis Kormilev, 1981a: 128 Arictus gamma Kormilev, 1972d: 581 Arictus gracilis Kormilev, 1983a: 462 Arictus grandis Kormilev, 1976a: 74 Arictus gressitti Kormilev, 1971a: 110 Arictus longicornis Kormilev, 197 1a: 111 Arictus monteithi Kormilev, 1965b: 32 [published August 31, 1965, ap- peared previously (1965a: 5, dated August 26, 1965) asa nomen nudum] Arictus sedlaceki Kormilev, 1971a: 110 Arictus taiwanicus Kormilev, 197 1a: 109 Arictus wagneri Kormilev and Heiss, L977e2100 Artabanellus mcnamarai Kormilev, IG67ex 17 Artabanus australis Kormilev, 1958b: 91 Artabanus bellicosus Kormilev, 1976a: 73 Artabanus bilobiceps papuasicus Kormi- leva L96792°527 Artabanus brachypterus Kormilev, 197 1a: 18 Artabanus brevipennis Kormilev, 1972d: 574 Artabanus burmensis Kormilev, 1983a: 450 Artabanus decemspinosus Kormilev, 1978e: 235 Artabanus degeneratus Kormilev, 197 1a: 16 Artabanus denticeps Kormilev, 1971a: 23 Artabanus fijiensis Kormilev, 1971a: 19 Artabanus fukienensis Kormilev, 19551: 182 Artabanus gazellensis Kormilev, 197 1a: 1, Artabanus gloriosus Kormilev, 1971a: 24 Artabanus halaszfyi Kormilev, 1955d: 199 Artabanus inermis Kormilev, 1955d: 201 Artabanus longiceps Kormilev, 1978e: 238 Artabanus magnificus Kormilev, 197 1a: 25 Artabanus mcfarlandi Kormilev, 1967g: 528 530 Artabanus micropterus Kormilev, 197 1a: 15 Artabanus montanus Kormilev, 197 1a: 20 Artabanus oviventris Kormilev, 1978e: 237 Artabanus rennellensis Kormilev, 19681: 99 Artabanus robustus Kormilev, 1971a: 21 Artabanus sexdentatus Kormilev, 197 1a: 23 Artabanus similis Kormilev, 1978e: 241 Artabanus singaporensis Kormilev, 1978e: 240 Artabanus solomonensis Kormilev, 1978c: 57 Artabanus sumatrensis Kormilev, 1973b: [2 Artabanus superbus Kormilev, 197 1a: 24 Artabanus truncatus Kormilev, 1967g: S27 Artabanus tuberculatus Kormilev, 1867g: a2 Artabanus umboiensis Kormilev, 197 1a: 18 Artabanus victor Kormilev, 1971a: 23 Artabanus vietnamensis Kormilev, 1973b: 7h Artagerus martinezi Kormilev, 1959b: 67 and 68 Artagerus plaumanni Kormilev, 1959b: 67 and 69 Aspisocoris termitophilus Kormilev, 1867g: 517 Atactocoris farri Kormilev, 1864d: 115 Axapisocoris brachypterus Kormilev and Heiss, 1979c: 157 Axapisocoris secundus Kormilev and Heiss, 1979c: 159 Barcinus truncatus Kormilev, 1955d: 196 Bergrothiessa confusa Kormilev, 1966f: S15,2516 Bergrothiessa grossa Kormilev, 1959e: 315 Bergrothiessa paranensis Kormilev, 1964b: 34 Bergrothiessa plaumanni Kormilev, 1960e: 208 Bergrothiessa rufa Kormilev, 1959e: 317 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Biroana armigera Kormilev, 1971b: 718 Biroana eurycephala Kormilev, 1957a: 397 Brachyrhynchus pauper Kormilev and Froeschner, 1987c: 118 [new name for preoccupied Mezira modesta Kormilev, 1972d: 583] Caecicoris oviventris Kormilev, 1957a: 399 Calisiopsis brasiliensis Kormilev, 1959a: 222 Calisiopsis minutus Kormilev, 1959a: 221 Calisiopsis planiceps Kormilev, 1976a: 69 Calisius africanus Kormilev and Heiss, 1976c: 41 Calisius ashlocki Kormilev, 1967e: 460 Calisius australis Kormilev, 1959a: 218 Calisius bilobatus Kormilev, 1959a: 214 Calisius borneensis Kormilev, 1986e: 250 Calisius brachypterus Kormilev, 1967e: 476 Calisius brasiliensis Kormilev, 1967h: 2 Calisius brevicornis Kormilev, 1967: 74 Calisius breviscutatus Kormilev, 1967b: 73 Calisius caledonicus Kormilev, 1971b: 707 Calisius confusus Kormilev, 1955a: 216 Calisius diffusus Kormilev, 1971b: 706 Calisius discrepans Kormilev, 1967e: 473 Calisius distinctus Kormilev, 1967e: 464 Calisius excelsus Kormilev, 1967e: 472 Calisius farri Kormilev, 1964g: 153 Calisius fuscus Kormilev, 1967e: 454 Calisius gracilicornis Kormilev, 1971b: 705 Calisius gracilis Kormilev, 1959a: 211 Calisius granuliger Kormilev, 1967e: 469 Calisius gressitti Kormilev, 1971b: 704 Calisius grossi Kormilev, 1966e: 284 Calisius hackeri Kormilev, 1959a: 219 Calisius hebridensis Kormilev, 1972c: 554 Calisius himalayensis Kormilev and Heiss, 1975c: 94 Calisius histrionicus Kormilev, 1971b: 708 Calisius homalanthi Kormilev, 1967e: 477 Calisius insignis Kormilev, 1959a: 215 VOLUME 97, NUMBER 3 Calisius leai Kormilev, 1966e: 290 Calisius liliputianus Kormilev, 1967e: 466 Calisius longiventris Kormilev, 1959a: 216 Calisius magdalenae Kormilev, 1966e: 288 Calisius minutus Kormilev, 1967e: 463 Calisius montanus Kormilev, 1967e: 462 Calisius nasutus Kormilev, 1967e: 461 Calisius notabilis Kormilev, 1966e: 291 Calisius orientalis Kormilev, 1971b: 703 Calisius ornatus Kormilev, 1967e: 455 Calisius pallidus Kormilev, 1967e: 470 Calisius pangoensis Kormilev, 1976e: 736 Calisius parvus Kormilev, 1967e: 474 Calisius pilosulus Kormilev, 1967e: 474 Calisius pulcher Kormilev, 1976e: 737 Calisius pusillus Kormilev, 1967e: 468 Calisius quadridentatus Kormilev, 1967e: 466 Calisius robusticornis Kormilev, 1967b: 75 Calisius septimus Kormilev, 1966e: 288 Calisius seychellensis Kormilev, 1963b: 602 Calisius sordidus Kormilev, 1967e: 467 Calisius tasmanicus Kormilev, 1963b: 604 Calisius testaceus Kormilev, 1967e: 461 Calisius texasanus Kormilev, 1968k: 45 Calisius variegatus Kormilev, 1968k: 44 Calisius varius Kormilev, 1967: 475 Calisius zimmermani Kormilev, 1967e: 471 Camerarius aberrans Kormilev, 1969a: 69 Camerarius armatus Kormilev, 1953c: 335 Camerarius armigera Kormilev, 1976a: RS Camerarius ceylonensis Kormilev, 198 1a: 125 Camerarius indicus Kormilev, 1972c: 566 Camerarius intermediarius Kormilev, 1969a: 68 Camerarius solomonensis Kormilev, 1972E2 S67 Camerarius wappersi Kormilev, 1954f: 128 531 Carventus australis Kormilev, 1958b: 87 Carventus biroi Kormilev, 1954f: 125 Carventus brachypterus Kormilev, 1966e: 301 Carventus chilensis Kormilev, 1981b: 296 Carventus crassus Kormilev, 1969a: 64 Carventus depressus Kormilev, 1969a: 59 Carventus dilatatus Kormilev, 1974a: 62 Carventus dollingi Kormilev, 1985a: 1046 Carventus elegantulus Kormilev, 1967c: 478 Carventus elongatus Kormilev, 1965b: 16 Carventus gracilis Kormilev, 1969a: 59 Carventus grandis Kormilev, 1972c: 559 Carventus gressitti Kormilev, 1969a: 61 Carventus guineensi Kormilev, 1974a: 63 Carventus hebridensis Kormilev, 1972c: 564 Carventus horvathi Kormilev, 1954f: 123 Carventus longiceps Kormilev, 1969a: 58 Carventus longiventris Kormilev, 1967f: 292 Carventus luteomarginatus Kormilev, 1955d: 194 Carventus malayensis Kormilev, 1966e: 295 Carventus malekulensis Kormilev, 1973b: 68 Carventus micropterus Kormilev and Heiss, 1979c: 155 Carventus milleri Kormilev, 1967c: 478 Carventus milleri borneensis Kormilev, 1967c: 480 Carventus minusculus Kormilev, 1969a: 61 Carventus minutus Kormilev, 1955k: 486 Carventus ovatus Kormilev, 1966e: 296 Carventus oviventris Kormilev, 1967f: 294 Carventus papuasicus Kormilev, 1969a: 58 Carventus parvulus Kormilev, 1985b: 98 [new name for preoccupied Carven- tus parvus Kormilev, 1973] Carventus parvus Kormilev, 1973b: 70 [preoccupied, see Carventus parvulus Kormilev, 1985b: 98] Carventus peterseni Kormilev, 1967f: 295 Carventus philippinensis Kormiley, 1969a: 58 532 Carventus pusillus Kormilev, 1972c: 565 Carventus pygmaeus Kormilev, 1972c: 563 Carventus quatei Kormilev, 1969a: 65 Carventus robustus Kormilev, 1966e: 298 Carventus sinensis Kormilev, 1969a: 62 Carventus stolidus Kormilev, 1969a: 63 Carventus taiwanensis Kormilev, 1969a: 63 Carventus variegatus Kormilev, 1969a: 64 Carventus vicinus Kormilev, 1972c: 561 Carventus vietnamensis Kormilev, 1969a: ei Carventus zimmermani Kormilev, 1969a: 39 Chapadia alata Kormilev, 1960a: 44 Chelonocoris tomentosus Kormilev, 1983a: 447 Chelonocoris usingeri Kormilev, 1956d: 287 Chelysocoris arachnoides Kormilev, 1956d: 288 Chelysosoma halaszfyi Kormilev, 1956d: 285 Chiastoplonia antennata Kormilev, 1967c: 484 Chiastoplonia angulata Kormilev, 1954f: 135 Chiastoplonia bellicosa Kormilev, 1954f: 132 Chiastoplonia chinai Kormilev, 19551: 184 Chiastoplonia confusa Kormilev, 1978e: 247 Chiastoplonia froeschneri Kormilev, 1978e: 250 Chiastoplonia liliputiana Kormilev, 19551: 185 Chiastoplonia lobata Kormilev, 1971a: 138 Chiastoplonia luzonica Kormilev, 1978e: 251 Chiastoplonia macarthuri Kormilev, 1978e: 248 Chiastoplonia minuta Kormilev, 1965b: 30 Chiastoplonia ovata Kormilev, 1954f: 134 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Chiastoplonia pauperula Kormilev, 1971la: 137 Chiastoplonia pusilla Kormilev, 1967c: 483 Chiastoplonia solomonensis Kormilev, 1978e: 249 Chinessa acuminata Kormilev, 1971la: 126 Chinessa armata Kormilev, 1971a: 120 Chinessa arnaudi Kormilev, 1984a: 76 Chinessa bellicosa Kormilev, 1971a: 121 Chinessa brachyptera Kormilev, 1971a: 119 Chinessa ferox Kormilev, 1971a: 127 Chinessa forfex Kormilev, 1971a: 120 Chinessa fulgida Kormilev, 1971a: 120 Chinessa gressitti Kormilev, 1971a: 129 Chinessa horribilis Kormilev, 1971a: 122 Chinessa iniqua Kormilev, 1971la: 129 Chinessa iniqua ceramensis Kormilev, 1971a: 130 Chinessa kokodensis Kormilev, 1983a: 461 Chinessa lobulata Kormilev, 1971a: 123 Chinessa lobuliventris Kormilev, 197 1a: 124 Chinessa major Kormilev, 1983a: 460 Chinessa modesta Kormilev, 1971a: 122 Chinessa necopinata Kormilev, 1971a: 125 Chinessa paralobulata Kormilev, 197 1a: 123 Chinessa quadridentata Kormilev, 197 1a: 125 Chinessa robusta Kormilev, 1971a: 127 Chinessa wauensis Kormilev, 1972d: Chrysodaspis angulatus Kormilev, 197 1a: 101 Chrysodaspis armatus Kormilev, 1976a: el, Chrysodaspis aurivillii Kormilev, 1976a: chen Chrysodaspis grandis Kormilev, 1973b: 81 Chrysodaspis maai Kormilev, 197 1a: 101 Chrysodaspis malayensis Kormilev and Heiss, 1979c: 160 VOLUME 97, NUMBER 3 Chrysodaspis ovatus Kormileyv, 197 1a: 101 Cinyphus alatus Kormilev, 1960d: 4 Cinyphus amazonicus Kormilev, 1960c: 83 Cinyphus andinus Kormilev, 1960d: 5 Cinyphus meziroides Kormilev, 1960d: 8 Cinyphus ovatus Kormilev, 1960d: 6 Cinyphus peruvianus Kormilev, 1960d: 7 Cinyphus saileri Kormilev, 1964a: 246 Cinyphus terminalis Kormilev, 1968k: 53 Cinyphus venezuelanus Kormilev, 1968k: 2 Clavicornia usingeri Kormilev, 1960f: 169 Clavicornia usingeri granulata Kormilev, 1967b: 75 **Coloborhinus” —see Coloborrhinus Coloborrhinus meziroides Kormilev, 1973f: 742 Coloborrhinus peruvianus Kormilev, 1975a: 10 Crimia inermis Kormilev, 1953c: 336 Crimia tertia Kormilev, 1983a: 463 Ctenoneurus aberrans Kormilev, 1967¢: Soi Ctenoneurus australis Kormilev, 1965a: 3 Ctenoneurus bakeri Kormilev, 1978e: 242 Ctenoneurus borneensis Kormilev, 197 la: By Ctenoneurus camerounensis Kormiley, 1974a: 69 Ctenoneurus cochereaui Kormilev, 197 1a: 55 Ctenoneurus coghiensis Kormilev, 197 1a: 60 Ctenoneurus fijiensis Kormilev, 197 1a: 59 Ctenoneurus quineensis Kormilev, 1974a: 69 Ctenoneurus halaszfyvi Kormilev, 1958b: 92 Ctenoneurus hammaensis Kormiley, 197 1a: 58 Ctenoneurus insignis Kormilev, 1971a: 56 Ctenoneurus longiceps Kormilev, 197 1a: 51 533 Ctenoneurus lugubris Kormilev, 197 1a: 55 Ctenoneurus major Kormilev, 1971a: 54 Ctenoneurus malayensis Kormilevy, 1973b: 73 Ctenoneurus mandrakaensis Kormilev, 1974a: 70 Ctenoneurus minutus Kormilev, 1967c: 481 Ctenoneurus myersi Kormilev, 1953c: 344 Ctenoneurus pendergrasti Kormilev, 1971a: 58 Ctenoneurus philippinensis Kormilev, 197 1a: 56 Ctenoneurus piligerus Kormilev, 1977d: 614 Ctenoneurus prominens Kormilev, 197 1a: 62 Ctenoneurus punctiventris Kormilev, 197 1a: 54 Ctenoneurus samoaensis Kormilev, 1971a: 61 Ctenoneurus simulans Kormilev, 197 1a: 61 Ctenoneurus solomonensis Kormilev, 197 ta: 52 Ctenoneurus spiculifer Kormilev, 197 1a: 60 Ctenoneurus sumatrensis Kormilev, 1973b: 74 Ctenoneurus terminalis Kormilev, 1974a: 68 Ctenoneurus wolffi Kormilev, 19681: 101 Ctenoneurus zimmermani Kormilev, NO WMias52 Daulocoris australis Kormilev, 1973c: 200 Daulocoris bicaudatus Kormilev, 197 1a: 102: 103 Daulocoris confusus Kormilev, 1973c: 201 Daulocoris formosanus Kormilev, 197 la: 104 Daulocoris gazellensis Kormilev, 197 1a: 104 Daulocoris nepalensis Kormilev and Heiss; 1973e¢71 Daulocoris robustus Kormilev, 197 1a: 105 Daulocoris sumatrensis Kormilev, 1980e: 23 534 Delnocoris micropterus Kormilev, 1982f: Dihybogaster incrustatus Kormilev, 1953e: 231 Dihybogaster plana Kormilev, 1959e: 309 and 310 Dihybogaster plaumanni Kormilev, 1959e: 309 and 311 Dimifaia lobulata Kormilev, 1971: 97 Dimorphacantha borneensis Kormilev, 1986e: 256 Diphyllonotus brachypterus Kormilev, L957c tS Diphyllonotus explanatus Kormilev, 1959b: 62 Drakeida incrustata Kormilev, 1958b: 90 Drakiessa confusa Kormilev, 1965b: 25 Drakeiessa minor Kormilev, 1963a: 446 Drakiessa parva Kormilev, 1965b: 24 Drakiessa tertia Kormilev, 1964c: 47 Dundocoris natalensis Kormilev, 1961a: 250 Dundocoris. stuckenbergi Kormilev, 196l1a: 249 Dysodius equatorianus Kormilev, 1975a: 9 Emydocoris stali Kormilev, 1963a: 448 Eretmocoris dominicus Kormilev, 1968a: 3 Erineocoris lobulatus Kormilev, 1960f: 164 Euchelonocoris hoberlandti Kormilev, 1974a: 65 Eunotoplocoris ruckesi Kormilev, 1957g: 4 Euricoris gloriosus Kormilev, 1965b: 22 Euricoris occultus Kormilev, 1957a: 395 Euricoris squalidus Kormilev, 1968g: 595 Forficulassa lobulata Kormilev, 1060a: 42 Froeschnerissa heveli Kormilev, 1986e: 254 Glochocoris abdominalis Kormilev, 1967: 78 Glochocoris acutus Kormilev, 1971a: 143 Glochocoris biroi Kormilev, 1960f: 170 Glochocoris borneensis Kormilev, 1986e: 258 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Glochocoris brisbanicus Kormilev, 1967b: 76 Glochocoris cristatus Kormilev, 1960f: 173 Glochocoris elongatus Kormilev, 1960f: 169 Glochocoris longiventris Kormilev, 1967c: 487 Glochocoris montanus Kormilev, 1973b: 77 Glochocoris monteithi Kormilev, 1967b: Tan Glochocoris pusillus Kormilev, 197 1a: 142 Glochocoris pygmaeus Kormilev, 1960f: 172 Glochocoris similis Kormilev, 1967c: 487 Glochocoris truncatus Kormilev, 197 1a: 143 Glochocoris tuberculatus Kormilev, 1960f: 171 Glyptoaptera woodwardi Kormilev, 1965b: 19 Glyptocoris annulatus Kormilev, 1953e: 235 Glyptocoris confusus Kormilev, 1953e: 233 Glyptocoris minutus Kormilev, 1959e: 313 and 314 Glyptocoris plaumanni Kormilev, 1954e: 126 Glyptomorpha malayensis Kormilev, 1977d: 613 Glyptomorpha thailandica Kormilev, 1977d: 613 Gnosotocoris gressitti Kormilev, 1967e: 452 Halaszfya elongata Kormilev, 1960e: 213 Halaszfya ovata Kormilev, 1960e: 213 Heissia rotundata Kormilev, 1986e: 252 Hesus mexicanus Kormilev, 1968c: 281 Hybocoris mexicanus Kormilev, 1982b: 9 Indiaradus cavagnaroi Kormilev, 1977d: 608 Indiaradus minor Kormilev, 1977b: 609 Kaulocoris stylatus Kormilev, 1971b: 720 Kelaino kjellanderi Kormilev, 1963a: 452 Kema acutissima Kormilev, 1971a: 116 VOLUME 97, NUMBER 3 Kema papuasica Kormilev, 1971a: 116 Kiritshenkiana furconotata Kormilev, 1976a: 72 Kiritshenkiessa spinipes Kormilev, 197 1a: 11 Kolpodaptera minuta Kormilev, 1966d: 11 Kolpodaptera rugosa Kormilev, 1966d: 12 Kormilevia aberrans Kormilev, 1963a: 449 Kormilevia gerali Kormilev, 1964b: 36 Leiocoris angulatus Kormilev, 197 1a: 114 Leiocoris borneensis Kormilev, 197 1a: 114 Leiocoris ferrugineus Kormilev, 197 1a: 13 Leiocoris subparallelus Kormilev, 197 1a: 114 Leurocoris caledonicus Kormilev, 1971b: TA Libiocoris lobatus Kormilev, 1968g: 594 Libiocoris pilicornis Kormilev, 1972c: 568 Libiocoris poecilus Kormilev, 1957a: 391 Limonocoris jolyi Kormilev, 1971le Llaimacoris penai Kormilev, 1964d: 118 Magdalenia excelsa Kormilev, 1983a: 446 Mastigocoris brachypterus Kormilev, 198la: 129 Mastigocoris insularis Kormilev, 1983a: 442 Mastigocoris malayensis Kormilev, 1967g: 514 Mastigocoris philippinensis Kormilev, 1972d: 572 Mastigocoris truncatus Kormilev and Heiss; 197 7e: 102 Mastigocoris usingeri Kormilev, 197 1a: 12 Mapiri paradoxa Kormilev, 1959b: 70 Mezira africana Kormilev, 1974a: 75 Mezira alfa Kormilev, 1955k: 496 Mezira amazonica Kormilev, 1962b: 267 Mezira andina Kormilev, 1965d: 6 Mezira argentinensis Kormilev, 195S5a: 224 and 228 Mezira (Zemira) armata Kormilev, 1964f: 463 535 Mezira armigera Kormilev, 1971a: 36 Mezira arnaudi Kormilev, 1975a: 20 Mezira auripilosa Kormilev, 1983d: 821 Mezira auritomentosa Kormilev, 1955k: 488 Mezira barberi Kormilev, 1964a: 256 Mezira beta Kormilev, 1955k: 501 Mezira (Zemira) bhoutanensis Kormilev, 1973c: 199 Mezira birabeni Kormilev, 1955a: 226 Mezira boliviana Kormilev, 1962b: 272 Mezira bonaerensis Kormilev, 1960c: 89 Mezira brachyptera Kormilev, 1964d: 115 Mezira brasiliensis Kormilev, 1976a: 76 Mezira breviceps Kormilev, 1971a: 48 Mezira bridarolli Kormilev, 1960c: 90 Mezira bruchi Kormilev, 1955a: 236 Mezira (Zemira) burmensis Kormiley, 1971a: 48 Mezira carioca Kormilev, 1964a: 255 Mezira championi Kormilev, 1964a: 251 Mezira chemsaki Kormilev, 1982f: 40 Mezira chinai Kormilev, 1955k: 505 Mezira cornigera Kormilev, 1953c: 339 Mezira costalimai Kormilev, 1964a: 253 Mezira (Zemira) crenata Kormilev, 1973b: 79 Mezira crenulata Kormilev, 1968a: 12 Mezira cubana Kormilev, 1960e: 215 Mezira (Zemira) dentipes Kormilev, 1974c: 243 Mezira doesburgi Kormilev and Froesch- ner, 1987c: 150 [new name for pre- occupied Mezira surinamensis Kor- milev, 1974c: 242] Mezira dybasi Kormilev, 1968c: 283 Mezira elegans Kormilev, 1967g: 543 Mezira enigmatica Kormilev, 1967g: 547 Mezira equatoriana Kormilev, 1968a: 9 Mezira eurycephala Kormilev, 1960c: 87 Mezira exarmata Kormilev, 1971a: 37 Mezira formosa Kormilev, 1955a: 226 and 243 Mezira fritzi Kormilev, 1979e: 344 Mezira (Zemira) funebra Kormilev, 197 1a: 46 Mezira funebra sumbaensis Kormilevy, 1973b: 80 536 Mezira ghanaensis Kormilev, 1966a: 393 Mezira gigantea Kormilev, 1955k: 490 Viezira gracilis Kormilev, 1968e: 52 Mezira guanacastensis Kormilev, 198 2a: 487 Mezira guianensis Kormilev, 1964a: 256 Mezira halaszfyi Kormilev, 1960d: 9 Mezira (Mezira) himalayensis Kormilev and Heiss, 1973: 69 Mezira (Zemira) hispida Kormilev, 197 1a: 34 Mezira hoberlandti Kormilev, 1960f: 166 Mezira hondurensis Kormilev, 1982a: 485 Mezira (Zemira) horrida Kormilev, 1977d: 620 Mezira hyperlobata Kormilev, 1962b: 266 Mezira inca Kormilev, 1960d: 12 Mezira (Zemira) incisa Kormilev, 197 la: 40 Mezira (Zemira) incognita Kormilev, 1976a: 76 Mezira incrustata Kormilev, 1973d: 741 Mezira (Zemira) indica Kormilev, 197 1a: 43 Mezira izzardi Kormilev, 1955k: 503 Mezira (Zemira) javanensis Kormilev, 1973¢: 197 Mezira (Zemira) kachinensis Kormilev, 1983a: 458 Mezira (Zemira) kerzhneri Kormilev, LOW las35 Mezira kjellanderi Kormilev, 1962b: 265 Mezira lindemannae Kormilev, 1956e: 42 Mezira lobuliventris Kormilev, 1953c: 340 Mezira longiceps Kormilev, 1977d: 616 Mezira (Zemira) longicornis Kormilev, 1971a: 42 Mezira (Zemira) longirostris Kormilev, 1982b: 15 Mezira luteomaculata Kormilev, 1957f: 271 Mezira luteonotata Kormilev, 1964a: 250 Mezira luzonica Kormilev, 1968): 231 Mezira maculata Kormilev, 1975a: 21 Mezira mekongensis Kormilev, 1983a: 458 Mezira mexicana Kormilev, 1964a: 252 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mezira minor Kormilev, 1960e: 214 Mezira (Zemira) modesta Kormilev, 1972d: 583 [preoccupied, see Bra- chyrhynchus pauper Kormilev and Froeschner, 1987c: 118] Mezira nasalis Kormilev, 1968c: 286 Mezira neonigripennis Kormilev, 1955a: 224 and 238 Mezira neonigripennis misionensis Kor- milev, 1955a: 240 Mezira neonigripennis neonigripennis Kormilev, 1955a: 238 Mezira nepalensis Kormilev and Heiss, 1973g: 66 Mezira nuda Kormilev and Heiss, 1973g: 64 Mezira oblonga Kormilev and Heiss, 1978d: 30 Mezira occidentalis Kormilev, 1968c: 287 Mezira occidentalis ‘“‘variety” appendi- culata Kormilev, 1968c: 288, invalid name, a post-1960 proposal of a “‘va- riety” Mezira paraangustata Kormilev, 1968c: 282 Mezira paraensis Kormilev and Heiss, 1979d: 114 Mezira paragranuligera Kormilev, 195S5a: 224 and 237 Mezira paraguayensis Kormilev, 1968: 10 Mezira paralata Kormilev, 1964a: 257 Mezira (Zemira) parallela Kormilev, 197 1a: 41 Mezira paratropicalis Kormilev, 1980d: 698 Mezira parvicornis Kormilev, 1973d: 435 Mezira pauperula Kormilev, 1962b: 270 Mezira peruviana Kormilev, 1960d: 11 Mezira (Zemira) philippinensis Kormi- lev, 1971la: 43 Mezira pilifera Kormilev, 1967h: 9 Mezira piligera Kormilev, 1971a: 29 Mezira pilosa Kormilev, 1968e: 51 Mezira pilosula Kormilev, 1973c: 196 Mezira placida Kormilev, 1968a: 13 Mezira placida haitiensis Kormilev, 1968a: 15 VOLUME 97, NUMBER 3 Mezira plaumanni Kormilev, 1966f: 520 Mezira proseni Kormilev, 1960c: 92 Mezira proxima Kormilev, 1982a: 486 [preoccupied, see Mezira vicina Kor- milev and Froeschner, 1987c: 160] Mezira pusilla Kormilev, 1968a: 8 Mezira (Zemira) quadridentata Kormi- lev, 1971a: 35 Mezira (Zemira) quadrispinosa Kormi- lev, 1973b: 78 Mezira roberti Kormilev, 1971a: 28 Mezira romani Kormilev, 1962b: 268 Mezira (Zemira) rossi Kormilev, 197 1a: 47 Mezira saltensis Kormilev, 1955a: 230 Mezira sangabrielensis Kormilev, 1962b: 270 Mezira sanmartini Kormilev, 1968a: 9 Mezira sayi Kormilev, 1982c: 777 Mezira (Zemira) serrata Kormilev, 1971a: 41 Mezira sinensis Kormilev, 1972d: 582 Mezira smithi Kormilev, 1982e: 336 Mezira (Zemira) solomonensis Kormi- lev, 197 1a: 47 Mezira spinipes Kormilev, 1965a: 6 Mezira spissigrada Kormilev, 1960c: 88 Mezira (Zemira) stolida Kormiley and Heiss, 1977e: 104 Mezira stysi Kormilev, 1976d: 88 Mezira (Zemira) subinermis Kormilev, 197 1a: 38 Mezira submontana Kormilev, 1983a: 456 Mezira subtriangula Kormilev, 1957f: 269 Mezira sulcata Kormilev, 1958b: 91 Mezira (Zemira) sumatrensis Kormilev, 1971a: 37 Mezira surinamensis Kormilev, 1974c: 242 [preoccupied, see Mezira does- burgi Kormilev and Froeschner, 1987c: 150] Mezira taiwanica Kormilev, 1957f: 267 Mezira tartagalensis Kormilev, 1955a: 224 and 233 Mezira tasmani Kormilev, 1955k: 492 Mezira termitophila Kormilev, 1976d: 88 537 Mezira (Zemira) thailandica Kormilev, 1971a: 39 Mezira timida Kormilev, 1968e: 53 Mezira tomentosa Kormilev, 1955c: 37 Mezira trinidadensis Kormilev, 1957c: 156 Mezira (Zemira) tristis Kormilev, 1973c: 198 Mezira tropicalis Kormilev, 1972a: 92 Mezira usingeri Kormilev, 1955k: 494 Mezira variegata Kormilev, 1968c: 283 Mezira venezuelana Kormilev, 197le Mezira veracruzensis Kormilev, 1968c: 288 Mezira vianai Kormilev, 1955a: 226 and 242 Mezira vicina Kormilev and Froeschner, 1987c: 160 [new name for preoccu- pied Mezira proxima Kormilev, 1982a: 486] Mezira vulcanica Kormilev, 1975a: 19 Mezira wilsoni Kormilev, 1967g: 542 Micromezira australis Kormilev, 1967c: 490 Miorrhynchus angulatus Kormilev, 1966d: 14 Miorrhynchus_ bolivianus Kormilev, 1959b: 64 and 65 Miorrhynchus brasilienssis Kormilev, 1959b: 64 and 66 Miorrhynchus championi Kormilevy, 1959b: 64 and 65 Miorrhynchus jatahyensis Kormilev, 1973f: 745 Miorrhynchus longicornis Kormilev, 1960d: 3 Miorrhynchus paraguayensis Kormilev, 1952a: 54 Miorrhynchus peruvianus Kormilev, 1960d: 4 Miorrhynchus plaumanni Kormilev, 1957c: 154 Miorrhynchus proseni Kormilev, 1959b: 62 and 64 Miorrhynchus schuhi Kormilev, 1976e: 741 Miorrhynchus undulatus Kormilevy, 1966d: 15 538 Miorrhynchus usingeri Kormilev, 1952c: 120 ‘*Nanium’’— see Nannium. Nannium brasiliense Kormilev, 1959e: 319 [genus name misspelled “Nan- ium’’| Neasterocoris costaricensis Kormilev, 1982d: 273 Neasterocoris spangleri Kormilev, 1978f: 199 Neophloeobia tuberculata Kormilev, 1967g: 524 Neuroctenus acutus Kormilev, 1971a: 83 Neuroctenus amazonicus Kormilev, 1960c: 91 Neuroctenus andrei Kormilev, 1975a: 2 [new name for preoccupied Neuroc- tenus villiersi Kormilev, 1973f: 740] Neuroctenus appendiculatus Kormilev, 197 1a: 88 Neuroctenus arizonicus Kormilev, 1982b: 9 Neuroctenus aztequi Kormilev, 1975a: 18 Neuroctenus bhoutanensis Kormilev and Heiss, 1975c: 99 Neuroctenus bicaudatus Kormilev, 197 la: 68 Neuroctenus bilobatus Kormilev, 197 la: 91 Neuroctenus brevirostris Kormilev, 197 la: 79 Neuroctenus burmensis Kormilev, 1976a: 74 Neuroctenus caudatus Kormilev, 1974a: 78 Neuroctenus ceramensis Kormilev, 197 1a: 93 Neuroctenus chilensis Kormilev, 1975a: 12 Neuroctenus colombianus Kormilev, 1968c: 280 Neuroctenus confusus Kormilev, 19551: 188 Neuroctenus consimilis Kormilev, 1974a: 73 Neuroctenus crassicornis Kormilevy, 1971a: 79 Neuroctenus discrepans Kormilev, 1980d: 696 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Neuroctenus dybasi Kormilev, 1971a: 85 Neuroctenus egelidus Kormilev, 1971a: 90 Neuroctenus eurycephalus Kormilev, 1971a: 86 Neuroctenus ferreus Kormilev, 1971a: 95 Neuroctenus ferrugineus Kormilev, 1974a: 74 Neuroctenus foratus Kormilev, 197 1a: 82 Neuroctenus franzi Kormilev and Heiss, 1973: 71 Neuroctenus freudei Kormilev, 1967d: 1 Neuroctenus fulgidus Kormilev, 197 1a: 80 Neuroctenus gracilis Kormilev, 1965b: 29 Neuroctenus grandis Kormilev, 1965b: 28 Neuroctenus granulatus Kormilev, 1973c: 205 Neuroctenus granuliger Kormilev, 1971: 82 Neuroctenus gressitti Kormilev, 197 1a: 72 Neuroctenus handschini Kormilev, 1953c: 342 Neuroctenus hortatrix Kormilev, 1971a: ie Neuroctenus hyalinipennis Kormilev, 97a? 77 Neuroctenus hyalinipennis australicus Kormilev, 1971la: 77 Neuroctenus incautus Kormilev, 1971a: 84 Neuroctenus incisus Kormilev, 1973c: 202 Neuroctenus indonesicus Kormilevy, 197 1a: 92 Neuroctenus insignis Kormilev, 1967h: 6 Neuroctenus kashmirensis Kormilev, 197 1a: 68 Neuroctenus latus Kormilev, 1977c: 261 Neuroctenus lestoni Kormilev, 1966a: 388 Neuroctenus lobulatus Kormilev, 197 1a: 93 Neuroctenus lobuliventris Kormilev, LOT Laie Neuroctenus longiceps Kormilev, 1973c: 206 Neuroctenus longicornis Kormilev, 197 1a: 90 Neuroctenus longissimus Kormilev, 1974c: 240 VOLUME 97, NUMBER 3 Neuroctenus longiusculus Kormilev, L9S7e:-157 Neuroctenus longiventris Kormilev, 1960d: 15 Neuroctenus luchti Kormilev, 197 1a: 87 Neuroctenus luteomaculatus Kormilev and Heiss, 1977e: 103 Neuroctenus luteomarginatus Kormilev, 1958b: 94 Neuroctenus meziroides Kormilev, 1958b: 94 Nauroctenus millironi Kormilev, 197 1a: 88 Neuroctenus minor Kormilev, 197 1a: 92 Neuroctenus minusculus Kormilev, NO 7tar dl Neuroctenus morio Kormilev, 1971a: 85 Neuroctenus muiri Kormilev, 1971a: 94 Neuroctenus necopinatus Kormilev, 1971la: 73 Neuroctenus pauperulus Kormilev, 197 1a: 79 Neuroctenus pavlitzkiae Kormilev, 1956e: 44 Neuroctenus perplexus Kormilev, 1983a: 453 Neuroctenus persimilis Kormilev, 197 1a: 87 Neuroctenus perssoni Kormilev, 1976a: 1S Neuroctenus placidus Kormilev, 1974a: 73 Neuroctenus proseni Kormilev, 1960c: 92 Neuroctenus pygmaeus Kormilev, 1978c: 58 Neuroctenus reynaudi Kormilev, 1973c: 204 Neuroctenus robustus Kormilev, 1973f: 739 Neuroctenus rossi Kormilev, 1975a: 15 Neuroctenus sauteri Kormilev, 1978c: 58 Neuroctenus scabrosus Kormilev, 197 1a: 710 Neuroctenus schlingeri Kormilev, 1975a: 14 Neuroctenus seclusus Kormilev, 197 1a: 81 Neuroctenus sedlaceki Kormilev, 197 1a: 78 539 Neuroctenus sertiventris Kormilev, 197 1a: 73 Neuroctenus sinensis Kormilev, 19551: 186 Neuroctenus singularis Kormilev, 197 1a: 70 Neuroctenus spiniceps Kormilev, 1974a: ie Neuroctenus subandinus Kormilev, 1955a: 249 Neuroctenus subrugosus Kormilev, 1977d: 615 Neuroctenus substitutus Kormilev, 1975a: 17 Neuroctenus sumbaensis Kormilev, 1973b: 76 Neuroctenus sundaicus Kormilev, 1973b: 15 Neuroctenus surinamensis Kormilev, 1974c: 241 Neuroctenus taiwanicus Kormilev, 1955c: 39 Neuroctenus termitophilus Kormilev, 1982g: 25 Neuroctenus testaceus Kormilev and Heiss, 1978d: 29 Neuroctenus thailandicus Kormilev, 1982b: 14 Neuroctenus tritonicus Kormilev, 1973c: 207 Neuroctenus vanduzeei Kormilev, 1975a: 16 Neuroctenus villersi Kormilev, 1973f: 740 [preoccupied, see Neuroctenus andrei Kormilev, 1975a: 2] Notapictinus amazonicus Kormilev, 1968k: 50 Notapictinus angulatus Kormilev, 1964f: 471 Notapictinus araguaensis Kormiley, 1985c: 153 Notapictinus bimaculatus Kormilev, 1979e: 343 Notapictinus christae Kormilev, 1967h: 6 Notapictinus diharpagus Kormilev, 1960d: 13 Notapictinus dissimilis Kormilev and Heiss, 1979d: 109 540 Notapictinus dollingi Kormilev, 1986e: 260 Notapictinus dyscritus Kormilev, 1960d: 14 Votapictinus equatoriensis Kormilev, 1973d: 438 Notapictinus hoguei Kormilev, 1975b: 219 Notapictinus incaicus Kormilev, 1964f: 473 Notapictinus kjellanderi Kormilev, 1964f: 472 Notapictinus micropterus Kormilev, 1964b: 38 Notapictinus notatus Kormilev, 1974¢c: 239 Notapictinus ornatus Kormilev, 1966d: 21 Notapictinus paramaculatus Kormilev, 1960e: 214 Notapictinus parvulus Kormilev, 1960a: 37 Notapictinus piliger Kormilev, 1982a: 483 Notapictinus platyceps Kormilev, 1966d: 20 Notapictinus plaumanni 1976a: 75 Notapictinus similis Kormilev and Heiss, 1979d: 107 Notapictinus subparallelus Kormilev, 1968k: 51 Notapictinus surinamensis Kormilev, 1974c: 239 Notapictinus terminalis Kormilev, 1964g: 159 Notapictinus testaceus Kormilev and Heiss, 1978d: 29 Notapictinus uruguayensis Kormilev, 1966d: 19 Notapictinus varelai Kormilev, 1975b: 221 Notapictinus venezuelanus Kormilev, LOE; 172 Notoplocaptera breviceps Kormilev, 1983a: 440 Notoplocaptera dollingi Kormilev, 1983a: 439 Notoplocaptera 1963a: 444 Kormilev, malaisei Kormilev, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Notoplocaptera mystica Kormilev, 1968g: 591 Notoplocoris guanabarensis Kormilev, 1982d: 274 Notoplocoris leptocerus Kormilev, 1963a: 452 and 454 Notoplocoris ovatus Kormilev, 1964b: 36 Notoplocoris robustus Kormilev, 1963a: 453 Notoplocoris triangulatus Kormiley, 19S 7e- 1 Notoplocoris usingeri Kormilev and Heiss, 1979d: 102 Odontonotus maai Kormilev, 19551: 179 Odontonotus oviventris Kormilev, 1957f: 265 Odontonotus sauteri Kormilev, 1955c: 35 Overlaetiella elongata Kormilev, 1983a: 448 Overlaetiella orientalis Kormilev, 1977a: 354 Paracalisiopsis aethiopicus Kormilev, 1963b: 607 Paracalisius granulatus Kormilev, 1974b: 296 Paracarventus woodwardi Kormilev, 1964c: 43 Parahesus truncatus Kormilev, 1960a: 47 Parapictinus brachypterus Kormilev, 1957e: 42 Parapictinus confusus Kormilev, 1983a: 444 Parapictinus ovatus Kormilev, 1956d: 293 Parartabanus tuberculatus Kormiley, 19724573 Pelecoris angustus Kormilev, 1971a: 98 Pictinellus brunneus Kormilev, 1968): 228 Pictinellus elongatus Kormilev, 1967g: 539 Pictinellus leai Kormilev, 1967g: 536 Pictinellus longiceps Kormilev, 1967g: 539 Pictinellus malayensis Kormilev, 1967g: 540 Pictinellus moturikiensis Kormilev, 1967g: 538 Pictinellus papuasicus Kormilev, 1967g: 534 Pictinellus thelmae Kormilev, 1966c: 306 VOLUME 97, NUMBER 3 Pictinus bechynei Kormilev, 1968k: 48 Pictinus beckeri Kormilev, 1959c: 22 Pictinus brachypterus Drake and Kor- milev, 1958a: 245 Pictinus carioca Kormilev, 1964f: 467 Pictinus derivatus Kormilev, 1959c: 25 Pictinus fictus Kormilev, 1959c: 29 Pictinus granuliferus Kormilev, 1959e: 318 Pictinus intermediarius Kormilev, 1953e: 241 Pictinus luteoincrustatus Kormilev, 1959c: 24 Pictinus maculatus Kormilev, 1959c: 26 Pictinus martinezi Kormilev, 1955a: 220 [genus name misspelled “Pictius” in species heading] Pictinus montrouzieri Kormilev, 1953e: 239 Pictinus nanus Kormilev, 1959c: 23 Pictinus pilosulus Kormilev, 1959c: 27 Pictinus plaumanni Kormilev, 1953e: 238 Pictinus rhombocarinatus Kormilev, 1964b: 37 Pictinus rutilus Kormilev, 1959c: 30 Pictinus sanmigueli Kormilev, 1959c: 24 Pictinus stali Kormilev, 1959: 27 Pictinus stolidus Kormilev, 1964g: 157 Pictinus venezuelanus Kormilev, 1968k: 49 Pictinus wittmeri Kormilev, 1959c: 28 **Pictius’ —see Pictinus “Placogenis” —see Placogenys Placogenys clarkei Kormilev, 1966d: 17 [genus name misspelled ‘‘P/acogen- ise Placogenys parva Kormilev, 1976e: 741 [genus name misspelled ‘‘Placogen- ise Placogenys constricta Kormilev, 1973d: 437 Probatoceps monteithi Kormilev, 1965b: 22 Proxius (Neoproxius) amazonicus Kor- milev, 1976e: Proxius (Nesoproxius) angulatus Korm1- lev, 1968g: 587 Proxius (Neoproxius) carioca Kormilev, 1982d: 270 541 Proxius (Nesoproxius) constrictus Kor- milev, 1978c: 54 Proxius (Neoproxius) costariquensis Kor- milev, 1982f: 38 Proxius (Nesoproxius) gracilis Kormilev, 1968g: 583 Proxius (Nesoproxius) hexagonalis Kor- milev, 1968g: 584 Proxius incaicus Kormilev, 1976e: 736 [new name for preoccupied Proxius peruvianus Kormilev, 1975a: 4] Proxius (Neoproxius) lindemannae Kor- milev, 1965d: 3 Proxius (Neoproxius) magdalenae Kor- milev, 1982f: 34 Proxius (Nesoproxius) malayensis Kor- milev, 1983a: 438 Proxius (Neoproxius) nicaraguensis Kor- milev, 1982f: 36 Proxius (Neoproxius) panamensis Kor- milev, 1982d: 272 Proxius (Neoproxius) peruvianus Kor- milev, 1966d: 9 Proxius (Neoproxius) peruvianus Kor- milev, 1975a: 4 [preoccupied, see Proxius (Neoproxius) incaicus Kor- milev, 1976e: 736] Proxius (Nesoproxius) punctulatus Kor- milev, 1968g: 585 Proxius (Nesoproxius) vietnamensis Kor- milev, 1968g: 584 Proxius (Nesoproxius) yoshimotoi Kor- milev, 1971b: 716 Pseudoartabanus brachypterus Kormilev, 197 ta: 133 Pseudopictinus dispar Kormilev, 1966f: 520 Rhombocoris distinctus Kormilev, 1965b: 18 Rhynchomirus brachypterus Kormilev, 1976e: 742 Rhysocoris cubanus Kormilev, 1976e: 740 Rhysocoris jamaicensis Kormilev, 1964d: 112 Rustem bellicosus Kormilev, 1957e: 41 Santaremis armata Kormilev, 1974c: 237 Santaremia robusta Kormilev, 1960a: 45 [“‘Schirrhocoris” —see Scirrhocoris] 542 Scironocoris armigerus Kormilev, 1957a: 402 Scironocoris baliensis Kormilev, 1972d: Dire Scironocoris malayensis Kormilev, 1983a: 454 Scironocoris obscurus Kormilev, 197 1a: 26 Scironocoris papuasicus Kormilev, 197 1a: aT Scirrhocoris australiensis Kormilev, 1965b: 27 Scirrhocoris mirabilis Kormilev, 1965b: 27 [genus name misspelled “Schir- rhocoris’’| Stelgidocoris villiersi Kormilev, 1974a: 67 Strigocoris bishopi Kormilev, 1972d: 574 Trigonaptera pandani Kormilev, 1968): 226 Trigonaptera scabrosa Kormilev, 1978c: Usingeria moralesi Kormilev, 1966a: 390 Usingerida angulata Kormilev, 1971a: 132 Usingerida longiceps Kormilev, 1977c: 262 Usingerida longirostris Kormilev, 1983a: 455 Usingerida montana Kormilev, 1974c: 236 Usingerida parva Kormilev, 1967f: 297 Usingerida similis Kormilev and Heiss, LOTSEM07 Usingerida walshi Kormilev, 1955h: 143 Usumbaraia ampliata Kormilev, 1956c: 254 Usumbaraia arnaudi Kormilev, 1979a: Bu Usumbaraia elongata Kormilev, 1956c: 255 Wuiessa brachyptera Kormilev, 1978e: 243 Wuiessa longicornis Kormilev and Heiss, I975¢: 103 Wuiessa minor Kormilev, 1971a: 131 Wuiessa wittmeri Kormilev and Heiss, 1975c: 104 Zimmermania angulata Kormilev, 1968g: 581 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Zimmermania bicaudata Kormilev, 1968g: 580 Zimmermania_ breviceps Kormilev, 1971b: 714 Zimmermania excavata Kormilev, 1968g: 578 Zimmermania gressitti Kormilev, 1968g: 582 Zimmermania magdalenae Kormilev, 1954f: 130 Zimmermania solomonensis Kormilev, 1968g: 578 Zimmermania usingeri Kormilev, 1967c: 475 Colobathristidae Carvalhoia carioca Kormilev, 1951a: 68 Colobasiastes proseni Kormilev, 1951f: 337 Diascopoea carvalhoi Kormilev, 195 1a: 78 Labradoria bridrollii Kormilev, 1951a: 80 Neocolobathristes bituberus Kormilev, 195la: 71 Neocolobathristes longispinus Kormilev, L9Sila: 73 Paraelopura tipuliformis Kormilev, 1953d: 291 Peruda monrosi Kormilev, 1949e: 378 Perudella denieri Kormilev, 1949a: 170 Phaenacantha saileri Kormilev, 1951la: 65 Symphylax handschini Kormilev, 1953d: 288 Symphylax walshi Kormilev, 1953d: 289 Trichocentrus amazonicus Kormiley, 1951£:77335 Trichocentrus guayanensis Kormiley, 195la: 74 Trichocentrus horvathi Kormilev, 1949e: 372 Trichocentrus vianai Kormilev, 1949e: 368 Coreidae Bathysolens nubilus form grisescens Kor- milev, 1939a: 4 Ceraleptus brevicornis Kormilev, 1939a: 25 Flavius tristriatus Kormilev, 1951h: 42 VOLUME 97, NUMBER 3 Hirilcus burmeisteri Kormilev, 1954a: 171 Hirilcus stali Kormilev, 1954a: 173 Hirilcus willineri Kormilev, 1954a: 169 Lycambes martinezi Kormilev, 1954a: 178 Marichisme boliviana Kormilev, 1954a: 163 Paralycambes misionensis Kormilev, 1954a: 181 Petalops bridarollii Kormilev, 1952d: 97 Petalops willineri Kormilev, 1952d: 94 Cyrtocoridae Cyphothyrea bridarollii Kormilev, 19551: 325 Cyrtocoris subobtusus Kormilev, 19551: 330 Elasmodemidae Elasmodema bosqi Kormilev, 1948a: 143 Lygaeidae Crophius coleopteroides Kormilev, 1950a: Do) Cymoninus argentinensis Kormilev, 1955f: 6 Karamania balcanica Kormilev, 1938a: 168 Micrymenus kuscheli Kormilev, 1952f: 12 Micrymenus rugosus Kormilev, 1957h: 387 Riggiella vianai Kormilev, 1949d: 6 Robinsonocoris tingitoides Kormilev, 1952f: 10 Megaridae Megaris vianai Kormilev, 1956a: 3 Miridae Phylus balcanicus Kormilev, 1939b: 197 Systellonotus skopljensis Kormilev, 1936a: 49 [manuscript name]; 1939b: 195 Ochteridae Ochterus foersteri Kormilev, 1952g: 155 Ochterus gressitti Kormilev, 1971d: 438 Ochterus jaczewskii Kormilev, 1971d: 439 Ochterus minor Kormilev, 1973a: 4 Ochterus piliferus Kormilev, 1973a: 6 Ochterus secundus Kormilev, 1971d: 441 Ochterus papuasicus Kormilev, 1972e: 585 543 Ochterus philippinensis Kormilev, 1971d: 436 Pentatomidae Adoxoplatys brasiliensis Kormilev, 1951c: 91 Adoxoplatys bridrolli Kormilev, 1949f: 316 Adoxoplatys gallardoi Kormilev, 1951c: 93 Adoxoplatys giaii Kormilev, 1950b: 343 Adoxoplatys panamensis Kormilev, 1951c: 90 Adoxoplatys willneri Kormilev, 1949f: 320 Carpocoris lunulatus nigricornis Kormi- lev, 1936a: 53 Daimonocoris bolivianus Kormilev, 1951h: 38 Glyphepomis setigera Kormilev, 1952h: 304 Harpagogaster willineri Kormilev, 19574: 50 Heteroscelis zischkai Kormilev, 195th: 40 Neoadoxoplatys haywardi Kormilev, 1956b: 5 Neoadoxoplatys saileri Kormilev, 1956b: 5 Neoderoploa willineri Kormilev, 1955e: 10 Neoleprosoma argentinensis Kormilev, 1952b: 214 Ornithosoma rivierei Kormilev, 1957d: 54 Placocoris albovenosus Kormilev, 1949c: 4 Prionotocoris suilari Kormilev, 1955e: 8 Ramosiana militaris Kormilev, 1950b: 340 Serdia bergi Kormilev, 1956b: 4 Serdia stali Kormilev, 1956b: 4 Serdia vianai Kormilev, 1956b: 3 Willinerinia decorata Kormilev, 1950d: 497 Phymatidae [“‘Agdistis” —see Agdistocoris] Agdistocoris pallida Kormilev, 1962e: 359 [genus name misspelled ‘A gdistis’’] 544 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Agdistocoris testacea Kormilev, 1962e: 359 Amblythyreus chapa Kormilev, 1962e: 353 and 356 imblythyreus izzardi Kormilev, 1962e: 353 Bakerinia acutangula Kormilev, 1962c: 12 Bakerinia granulata Kormilev, 1962c: 11 Carcinocoris ochraceus Kormilev, 1962e: 365 Chelocoris alatus Kormilev, 1988b: 69 Chelocoris bianchii Kormilev, 197 1c: 884 Chelocoris brancuccii Kormilev, 1988b: 13 Chelocoris spinosulus Kormilev, 1962e: 362 Chelocoris truncatus Kormilev, 1962e: 360 Chelocoris viethamensis Kormilev, 1990c: 298 Chelocoris wittmeri Kormilev, 1988b: 70 Cnizocoris abselluiferus Nonnaizab, Kor- milev, and Qi, in 1990a: 21 [manu- script name, changed to C. acellularis after manuscript was submitted to press—see that combination] Cnizocoris acellularis Nonnaizab, Kor- milev, and Qi, 1989c: 349 [published form of manuscript name C. absel- luiferus—see that combination] Cnizocoris aphyllus Nonnaizab, Kormi- lev, and Qi, 1990a: 21 [manuscript name, changed to C. unicellularis af- ter manuscript was submitted to press—see that combination] Cnizocoris bhutanensis Kormilev, 1988b: 64 Cnizocoris drakei Kormilev, 1957j: 64 Cnizocoris mongolicus Nonnaizab, Kor- milev and Qi, 1989c: 342 and 347 Cnizocoris sinensis Kormilev, 1957): 67 Cnizocoris unicellularis Nonnaizab, Kor- milev, and Qi, 1989c: 342 and 347 [published form for manuscript name C. aphyllus—see that combination] Diurocoris thailandicus Kormilev, 1984b: 631 Diurocoris truncatus Kormilev, 1962c: 13 Eurymnus insignis Kormilev, 1984b: 628 Glossopelta indica Kormilev, 1966g: 283 Kelainocoris farri Kormilev, 1963c: 223 Lophoscutus armatus Kormilev, 1986f: 301 Lophoscutus arnaudi Kormilev, 1988c: 142 Lophoscutus brasiliensis Kormilev and Doesburg, 1986b: 119 Lophoscutus chemsaki Kormilev, 1984b: 627 Lophoscutus confusus Kormilev, 1989d: 61 [was listed as a manuscript name “In press,” 1990a: 30] Lophoscutus dimorphus Kormilev, 1986f: 299 Lophoscutus froeschneri Kormilev, 1987a: 702 Lophoscutus geijskesi Kormilev and Doesburg, 1986b: 121 Lophoscutus haitiensis Kormilev, 1987a: 703 Lophoscutus hispaniolensis Kormilev and Doesburg, 1991a: 280 Lophoscutus maurus Kormilev, 1989a: 451 [was listed as a manuscript name “In press,” 1990a: 30] Lophoscutus michelbacheri Kormilev, 1984b: 628 Lophoscutus paracrassimanus Kormilev, 1988c: 144 Lophoscutus pugil Kormilev and Does- burg, 1992b: 417 Lophoscutus rideri Kormilev, 1988c: 144 Lophoscutus rileyorum Kormilev, 1990d: 221 [was listed as a manuscript name “In press,” 1990a: 32] Lophoscutus sagimani Kormilev and Doesburg, 1986b: 116 Lophoscutus schaffneri Kormilev, 1986b: 304 Lophoscutus virginensis Kormilev, 1986d: 306 Lophoscutus viridis Kormilev, 1984b: 626 Lophoscutus woolleyi Kormilev, 1988c: 141 Lophoscutus ypsilon Kormilev, 1990d: VOLUME 97, NUMBER 3 223 [was listed as a manuscript name “In press,” 1990a: 33] Macrocephalus argentinus Kormilev, 1950c: 581 Macrocephalus (Lophoscutus) aterrimus Kormilev, 1981lc: 216 Macrocephalus dissolutus Kormilev, 1984b: 625 Macrocephalus dollingi Kormilev, 1984b: 625 Macrocephalus (Lophoscutus) domini- canus Kormilev, 1963c: 224 Macrocephalus (Lophoscutus) drakei Kormilev, 1962a: 56 Macrocephalus (Lophoscutus) dudichi Kormilev, 1949b: 587 [new name for preoccupied M. obscurus Dudich, 1922: 172] Macrocephalus incisus maya Kormilev, 1962a: 52 Macrocephalus insignis Kormilev, 19571: 39 Macrocephalus (Lophoscutus) margaritis Kormilev, 1962a: 52 Macrocephalus (Lophoscutus) marmora- tus Kormilev, 1966g: 279 Macrocephalus notatus costariguensis Kormilev, 1962a: 51 Macrocephalus (Lophoscutus) paraspicu- losus Kormilev, 1953f: 57 Macrocephalus (Lophoscutus) prehensilis minor Kormilev, 1954c: 159 Macrocephalus (Lophoscutus) pulcher Kormilev, 1981lc: 218 Macrocephalus similis Kormilev, 1972b: 94 Macrocephalus (Lophoscutus) spiculiger Kormilev, 1968f: 243 Macrocephalus (Lophoscutus) subproduc- tus Kormilev, 1960a: 55 Macrocephalus testaceus Kormilev, 1963c: 223 Macrocephalus (Lophoscutus) wygodzin- skyi Kormilev, 1950c: 587 Neoanthylla bucki Kormilev, 1951g: 56 Neoanthylla peruviana Kormilev, 1964e: 138 Oxythyreus ruckesi Kormilev, 1962c: 3 545 Parabotha singularis Kormilev, 1984b: 629 Parabotha whiteheadi Kormilev, 1984b: 630 Paraphymata saileri Kormilev, 1962d: 464 Phymata albipennis Kormilev, 1965c: 207 Phymata americana obscura Kormilev, 1957b: 136 Phymata ampliata Kormilev, 1962d: 311 and 350 Phymata anchietai Kormilev, 1951b: 55 Phymata argentina montana Kormilev, 1951g: 98 Phymata barberi Kormilev, 1962d: 322 and 434 Phymata beieri Kormilev, 1962d: 310 and 333 Phymata bergi Kormilev, 1950: 586 Phymata bipunctata Kormilev, 195le: 174 Phymata birabeni Kormilev, 1950c: 582 Phymata bosgqi Kormilev, 1950c: 589 Phymata_ brailovskyi Kormilev, 1990f: 100 Phymata carioca Kormilev, 195le: 171 Phymata carinata cordobensis Kormilev, 1951g: 70 Phymata carinata pampeana Kormiley, 1951g: 71 Phymata carinata uruguayensis Korm1- lev. FLO Siew 2 Phymata carneipes subangulata Korm1- lev, 1962d: 439 Phymata carvalhoi Kormilev, 1950c: 588 Phymata centralis Kormilev, 1962d: 351 Phymata chilensis brasiliensis Kormilev, 1962d: 449 Phymata chinai Kormilev, 1962e: 350 Phymata colombiana Kormilev, 1962d: 429 Phymata delpontei Kormilev, 1950c: 583 Phymata denieri Kormilev, 1950c: 583 Phymata drakei Kormilev, 1957b: 131 Phymata elongata Kormilev, 1963c: 218 [preoccupied, see Phymata prolata Froeschner and Kormilev, 1990a: 67] 546 Ph mata fasciata panamensis Kormilev, 1962d: 411 Phymata foersteri Kormilev, 1951b: 53 [species name misspelled ‘‘foer- stieri’’] Phymata foerstieri [sic] Kormilev, 1951b: 53 [correct of species name species name is foersteri] Phymata fortificata paranensis Kormilev, 1951g: 88 Phymata fortificata patagonica Kormi- lev, 1951g: 89 Phymata granulosa evansi Kormilev, 1962d: 399 Phymata granulosa texasana Kormilev, 1957b: 134 Phymata haywardi Kormilev, 1950c: 585 Phymata husseyi Kormilev, 195le: 176 Phymata incisa Kormilev, 195le: 172 Phymata inconspicua Kormilev, 1962d: 431 Phymata jamaicensis Kormilev, 1963c: 216 Phymata luteomarginata Kormilev, 1957b: 130 Phymata maculata Kormilev, 1957b: 127 Phymata malaisei Kormilev, 1962d: 377 Phymata mansosotoi Kormilev, 1952e: 125 Phymata mayri Kormilev, 1962d: 393 Phymata minuta Kormilev, 1962d: 432 Phymata monrosi Kormilev, 1950c: 585 Phymata nasuta Kormilev, 1965c: 205 Phymata orfilai Kormilev, 1950c: 584 Phymata pacifica hainesi Kormilev, 1962d: 424 Phymata pallida Kormilev, 1957b: 128 Phymata paradoxa Kormilev, 1952e: 126 Phymata pilifera Kormilev, 1963c: 220 Phymata prolata Froeschner and Kor- milev, 1990a: 57 [new name for pre- occupied Phymata elongata Kormi- lev, 1963c: 218] Phymata rhynocerata Kormilev, 1957b: 124 Phymata roqueensis halaszfyi Kormilev, 1962d: 343 Phymata saileri Kormilev, 1957b: 133 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Phymata sanjuanensis Kormilev, 1950c: 530 Phymata severini dyeri Kormilev, 1962d: 389 Phymata simulans explanata Kormilev, 1962d: 346 Phymata simulans recifensis Kormilevy, 1962d: 348 Phymata simulans surinamensis Kormi- lev, 1962d: 347 Phymata simulans trinidadensis Kormi- lev, 1957b: 1125 Phymata subarmata Kormilev, 1953f: 59 Phymata superba Kormilev, 1962d: 452 Phymata torresi Kormilev, 1950c: 588 Phymata turnbowi Kormilev, 1983b: 139 Phymata variegata Kormilev, 1963c: 219 Phymata venezuelana Kormilev, 1950c: 589 Phymata vianai Kormilev, 1950c: 587 Phymata vicina parvula Kormilev, 1957b: 126 Phymata wygodzinskyi Kormilev, 1950c: 586 Phymatocoris iviei Kormilev and Does- burg, 1991a: 280 Themonocoris aethiopicus Kormilev and Doesburg, 1986b: 114 Piesmatidae Thaicoris sedlaceki Kormilev, 1969b: 646 Rhopalidae Maccevethus lineola macedonia Kormi- lev, 1936a: 54 Thaumastocoridae Discocoris vianai Kormilev, 1955g: 10 Thyreocoridae Galgupha denieri Kormilev, 1956a: 10 Galgupha durionei Kormilev, 1956a: 8 Galgupha fritzi Kormilev, 1956a: 11 Galgupha haywardi Kormilev, 1956b: 6 Galgupha torresi Kormilev, 1956a: 7 Galgupha vianai Kormilev, 1956b: 6 Pericrepis bergi Kormilev, 1956b: 7 Pericrepis bosqui Kormilev, 1956b: 7 Thyreocoris balcanicus ohridanus Kor- milev, 1936a: 53 Tingidae Baeotingis vianai Kormilev, 1955b: 65 VOLUME 97, NUMBER 3 Vianaididae Vianaida coleopterata Kormilev, 1955j: 472 Order Neuroptera Genus-group Names Mantispidae Paramantispa Williner and Kormilev, 1959d: 10 Species-group Names Mantispidae Drepanicus gayi schajovskoyi Williner and Kormilev, 1959d: 17 547 ACKNOWLEDGMENTS For very helpful reviews of this manu- script I am indebted to Nicholas A. Kor- milev (who modestly declined coauthorship of it), 2930 54th Street S., Gulfport, Florida; to Ernest Heiss, Entomology Research Group, Tiroler Landesmuseum, Innsbruck, Austria; to Thomas J. Henry, U.S. Depart- ment of Agriculture, Systematic Entomol- ogy Laboratory, National Museum of Nat- ural History, Washington, D.C.; and to A. G. Wheeler, Jr., Pennsylvania Department of Agriculture, Harrisburg, Pennsylvania. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 548-560 LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF ENTIBULLA CALIFORNICA (COQUILLETT) (DIPTERA: TEPHRITIDAE) ON CHRYSOTHAMNUS NAUSEOSUS (PALLAS) BRITTON IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN, DAviD H. HEADRICK, AND JEFFREY A. TEERINK Department of Entomology, University of California, Riverside, California 92521. Abstract. — Valentibulla californica (Coquillett) is univoltine and nearly monophagous on Chrysothamnus nauseosus (Pallas) Britton (Asteraceae). The larvae develop in short, subsurface mines mostly formed basally in current season’s branches, and are not easily detected on infested plants. They cause limited swellings, not readily detectable galls. The egg, second and third instar larva, and puparium are described and figured. The egg, described for the first time for this genus, is unique among nonfrugivorous, California tephritids studied to date because of its ellipsoidal-fusiform shape, acutely tapered pos- terior, and its prominent, anterior pedicel bearing aeropyles of two distinct diameters. The third instar characteristically is shagreened, i.e. only the integument of the gnatho- cephalon, the anterior third of the prothorax, and a small area surrounding the posterior spiracular plates are not covered by minute, dome-shaped verrucae. The puparium also maintains the shagreened surface. Premating, mating, and postmating behaviors are de- scribed; they are few, and similar to the closely related genus, Aciurina. Hymenopterous parasitoids of V. californica include three species of chalcidoids reared as primary, solitary, endoparasitoids from isolated puparia: Eupelmus sp. (Eupelmidae), Eurytoma sp. (Eur- ytomidae), and Halticoptera sp. (Pteromalidae). Key Words: Insecta, Valentibulla, Chrysothamnus, biology, taxonomy of immature stages, mating behavior, parasitoids. This study of Valentibulla californica (Coquillett) was undertaken after prolonged searches by RDG for its “galls,” as de- scribed and pictured by Wangberg (1978), failed repeatedly during the 1980’s at var- ious locations in southern California where adults otherwise were swept. Our subse- quent findings reported herein address sev- eral inconsistencies in the developmental biologies heretofore reported for V. califor- nica and its congeners, and add new knowl- edge on this tephritid otherwise largely known from States outside its type state- locality and namesake. MATERIALS AND METHODS Most field studies were conducted with V. californica on Chrysothamnus nauseosus (Pallas) Britton (Asteraceae) at a Mojave Desert (high-elevation desert) location in the easternmost foothills of the San Bernardino Mountains, 2 km NW of Pioneertown at 1280-m elevation, SW San Bernardino Co., during 1991-93. Samples of branches from plants that from previous dissections had been found to contain larvae and puparia were collected on different dates, months and seasons. Branch samples were trans- VOLUME 97, NUMBER 3 ported in cold-chests in an air-conditioned vehicle to the laboratory and stored under refrigeration for subsequent dissection, photography, description, and measure- ment. All larvae and 20 puparia dissected from these branches and eggs obtained from other excised branches used in oviposition studies were preserved in 70% EtOH for scanning electron microscopy (SEM). All other puparia were placed in separate glass rearing vials stoppered with absorbant cot- ton and held in humidity chambers at room temperature for adult and parasitoid emer- gence. Specimens for SEM later were hy- drated to distilled water in a decreasing se- ries of acidulated EtOH. They were osmi- cated for 24 h, dehydrated through an in- creasing series of acidulated EtOH, critically point dried, mounted on stubs, sputter- coated with a gold-palladium alloy, and studied with a JEOL JSM C-35 SEM in the Department of Nematology, University of California, Riverside. Most adults reared from isolated puparia were individually caged in 850-ml, clear- plastic, screened-top cages with a cotton wick and basal water reservoir and provisioned with a strip of paper toweling impregnated with yeast hydrolyzate and sucrose. These cagings were used for longevity studies and Oviposition tests in the insectary of the De- partment of Entomology, University of Cal- ifornia, Riverside, at 25 + 1°C, and 14/10 (L/D) photoperiod. Virgin male and female flies obtained from emergence vials were paired in clear-plastic petri dishes provi- sioned with a flattened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1991) for direct ob- servations, videorecording, and still-pho- tography of their general behavior, court- ship, and copulation. Six pairs were held together for at least 14 d and observations were made as opportunity allowed through- out each day. Field observations on adult behavior also were made on C. nauseosus at Cajon Junction at 1010-m elevation, San Bernardino Nat. Forest (N section), SW San 549 Bernardino Co., from 8.iv.1992 to 12.v.1992 for ca. 4 h per day for 2 days per week. Plant names used in this paper follow Munz (1974); tephritid names follow Foote et al. (1993) except for Aciurina trixa Curran (Dodson and George 1986). Nomenclature used to describe the immature stages fol- lows Goeden and Headrick (1990, 1991a, b, 1992), Headrick and Goeden (1990, 1991, 1993), and the telegraphic format of Goe- den et al. (1993). Means + SE are used throughout this paper. Voucher specimens of reared adults of V. californica and its par- asitoids reside in the research collections of RDG; preserved specimens of larvae and puparia are stored in separate collections of immature Tephritidae maintained by DHH and JAT. RESULTS AND DISCUSSION Taxonomy Coquillett (1894) first described V. cali- fornica (as a Trypeta). Foote et al. (1993) reviewed the taxonomy and other literature on this species and described and discussed recognition features for the adults. They also provided the key to U.S. and Canadian spe- cies used to confirm the identity of our flies. Wangberg (1978) briefly described and pro- vided line drawings of one third-instar larva and one puparium serving to illustrate both V. californica and V. steyskali Foote and labelled “‘Valentibulla spp.,” but without distinguishing these species. Egg.—Egg body smooth, shiny, white, elongate ellipsoidal-fusiform (Fig. 1 A); ped- icel peg-like, 0.02 mm long (Fig. 1B); 17 eggs dissected from branches in oviposition cages averaged 0.50 + 0.006 (range, 0.41- 0.52) mm in length, 0.15 + 0.002 (range, 0.13-0.17) mm in width. This is the first egg described for any Val- entibulla species. The egg of V. californica differs in shape from previously studied, nonfrugivorous tephritids (Goeden and Headrick 1992; Goeden et al. 1993, 1994a, 166.6U i5KU X240 gao2 Fig. 1. Egg of V. californica: (A) habitus, pedicel at left; (B) detail of pedicel, showing aeropyles. b; Green et al. 1993: Headrick and Goeden 1990) in that it is more acutely tapered pos- teriorly opposite the pedicel. Tephritis ari- zonaensis Quisenberry has a similar egg body shape, but does not taper as dramatically at the posterior end and the pedicel is not as prominent (Goeden et al. 1993). The egg of Trupanea californica Malloch is similar in length and slightly wider, however the shape is elongate-ellipsoidal and the pedicel is smaller (Headrick and Goeden 1991). The egg of Acuirina thoracica Curran, which 1s in the same subtribe as V. californica (Foote et al. 1993), is shaped very differently and bears distinctly raised polygonal reticula- tion on its surface (Headrick and Goeden 1993). The pedicel of V. californica also dif- fers from all previously examined eggs in that the aeropyle openings are of two dis- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tinct diameters (Fig. 1B). Tephritis bac- charis (Coquillett) has a similar shaped ped- icel twice as long as V. californica (Goeden and Headrick 1991b). Third instar.— Third instar creamy white, shagreened, elongate-ellipsoidal, bluntly rounded apically and posteriorly (Fig. 2A); gnathocephalon conical, smooth with very few rugose pads; pad margins laterad of the mouth lumen partly contiguous and not dis- tinctly raised (Fig. 2B-1); paired dorsal sen- sory organs dorsomediad of anterior sen- sory lobes each consisting of a single, dome- shaped papilla (Fig. 2B-2, 2C-1); anterior sensory lobes bear the lateral sensory organ (Fig. 2C-2), terminal sensory organ (Fig. 2C- 3), and a papillate sensillum dorsad of the lateral sensory organ (Fig. 2C-4); stomal sense organs lie ventrad of anterior sensory lobes, near the mouth lumen (Fig. 2C-5): lateral (Fig. 2B-3) and ventrolateral sensilla present (Fig. 2B-4), each consisting of a smooth, verruciform sensillum with a cen- tral pore; mouth hooks bidentate, teeth stout, conical, rounded apically (Fig. 2B-5, 2D-1); median oral lobe laterally compressed, ta- pers anteriorly between mouth hooks (Fig. 2B-6), attached basally to labial lobe (Fig. 2D-2); labial lobe bears two pore sensilla (Fig. 2D-3); prothorax smooth anteriorly, covered with minute, dome-shaped verru- cae posteriorly; anterior thoracic spiracles located dorsolaterally on posterior margin of prothorax, each consisting of four, obo- vate papillae (Fig. 2E); meso- and metatho- rax covered entirely by minute, dome- shaped verrucae; lateral spiracular complex on abdominal segments composed of an open spiracle (Fig. 2F-1), and a single ver- ruciform sensillum (Fig. 2F-2); caudal seg- ment bears posterior spiracular plates (Fig. 2G); plates bear three, oval rimae ca. 0.02 mm long (Fig. 2G-1), and four, short, mul- tibranched interspiracular processes, with three to four branches each, the longest measuring 0.006 mm (Fig. 2G-2); stelex sensilla surround margin of caudal segment in four-dorsal, six-ventral arrangement; VOLUME 97, NUMBER 3 caudal segment additionally bears a pair of compound sensilla ventrad of the spiracular plates consisting of a tuberculate chemo- sensillum (Fig. 2H-1), and a stelex sensillum (Fig. 2H-2). Valentibulla californica is only the second species examined by us that is shagreened. Tomoplagia cressoni Aczél, which belongs to a different tribe, also is shagreened (Goe- den and Headrick 199 1a). In V. californica, only the integument of the gnathocephalon, the anterior third of the prothorax, and a small area surrounding the posterior spi- racular plates are not covered by minute, dome-shaped verrucae. Aciurina thoracica, a closely related species (Foote et al. 1993), bears irregular patches of acanthae, but the intersegmental areas are smooth (Headrick and Goeden 1993). Valentibulla californica also lacks the rows of minute acanthae that circumscribe the intersegmental areas of several other tephritid species (Goeden and Headrick 1991b, 1992, Goeden et al. 1993, 1994b, Headrick and Goeden 1991). Foote et al. (1993) placed Valentibulla in a new subtribe Eurostina along with Aci- urina and Eurosta. Valentibulla californica larvae do resemble 4. thoracica larvae in many morphological characters (Headrick and Goeden 1993). The gnathocephalon 1s similar to A. thoracica, being smooth with few rugose pads. The median oral lobes are similar, laterally compressed and tapered apically. The gnathocephalon sensilla are very similar in structure and placement; however, there is one noteworthy differ- ence. The anterior sensory lobe in V. cali- fornica \acks a pit sensory organ. This 1s the second example of a tephritid with its an- terior sensory lobe not bearing the pit sen- sory organ. Zomoplagia cressoni, which is also shagreened, lacks the pit sensory organ (Goeden and Headrick 1991a). Pit sensory organs otherwise are typically similar in size, shape, and placement among tephritid spe- cies (Carroll and Wharton 1989, Goeden and Headrick 1990, 1991b, 1992, Goeden et al. 1993, 1994a, b, Headrick and Goeden 551 1990, 1993). The mouth hooks also differ, being bidentate in V. californica and tri- dentate in A. thoracica (Headrick and Goe- den 1993). Wangberg (1978) reported Val- entibulla spp. to have tridentate mouth hooks. The abdominal lateral spiracular complex consists of a spiracle and a single, domed-shaped papillae in V. californica, similar to those of 4. thoracica (Headrick and Goeden 1993), Trupanea californica (Headrick and Goeden 1991), 7. nigricornis (Coquillett), 7. bisetosa (Coquillett) (Knio and Goeden, unpublished), N. viridescens Quisenberry (Goeden and Headrick 1992), Procecidochares flavipes Aldrich (Goeden et al. 1994a), and P. stonei Blanc and Foote (Green et al. 1993). Second instar.—Second instar creamy white, shagreened, ovoid in shape, rounded anteriorly and posteriorly (Fig. 3A); gnath- ocephalon conical, laterally flattened, smooth with a few small rugose pads laterad of the anterior sensory lobes and stomal sense organs (Fig. 3B-1), and a few petals dorsad of the mouth lumen (Fig. 3B-2); paired dorsal sensory organs located dor- somedially to the anterior sensory lobes, consisting of a single dome-shaped papilla (Fig. 3C-1); anterior sensory lobes separated by a medial depression (Fig. 3C), flattened, bearing the terminal sensory organ (Fig. 3C- 2), lateral sensory organ (Fig. 3C-3), and a papillate sensillum dorsomediad of the lat- eral sensory organ (Fig. 3C-4); stomal sense organs lie ventrad of anterior sensory lobes, near the mouth lumen (Fig. 3C-5); mouth hooks bidentate, teeth conical, rounded api- cally (Fig. 3B-3); median oral lobe laterally flattened, tapered anteriorly between mouth hooks (Fig. 3B-4); prothorax smooth ante- riorly, covered with minute, dome-shaped verrucae posteriorly; anterior thoracic spir- acles located dorsolaterally on posterior margin of prothorax, each consisting of four obovate papillae (Fig. 3D); lateral spiracular complex not observed; caudal segment bears the spiracular plates, plates bear three oval rimae ca. 0.007 mm long, and four, thorn- 552 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 15SKV X32 8661 1006. 4U Fig. 2. Third instar larva of V. californica: (A) habitus, anterior to left; (B) gnathocephalon, left anterolateral view, | —rugose pads, 2—dorsal sensory organ, 3 —lateral sensillum, 4— ventrolateral sensillum, 5— mouth hooks, 6—median oral lobe; (C) left anterior sensory lobe, 1 —dorsal sensory organ, 2—lateral sensory organ, 3—terminal sensory organ, 4—papillate sensillum, 5—stomal sense organ; (D) gnathocephalon, ventral view, 1—mouth hooks, 2—median oral lobe, 3—labial sensilla; (E) anterior thoracic spiracles; (F) lateral spiracular complex, VOLUME 97, NUMBER 3 553 OG19 1SKY %1166 se 16. GU Fig. 3. Second instar larva of V. californica: (A) habitus, anterior to left; (B) gnathocephalon, anterolateral view, 1—rugose pads, 2—dorsal petals, 3—mouth hooks, 4—median oral lobe; (C) anterior sensory lobe, 1 — dorsal sensory organ, 2—terminal sensory organ, 3—lateral sensory organ, 4—papillate sensillum, 5—stomal sense organ; (D) anterior thoracic spiracles. like interspiracular processes, longest mea- suring 0.002 mm; caudal sensilla could not be seen on this specimen. The features of the second instar of V. californica are fairly well-developed mor- phologically. The gnathocephalon closely resembles that of the third instar. The ru- gose pads are more widely separated. The petals are present dorsal to the mouth lu- men. The mouth hooks are similar in shape to the third instar and bidentate. The me- dian oral lobe is also similar in shape to that of the third instar. The pit sensory organ was not observed on the anterior sensory lobe of the second instar. The lateral and ventrolateral sensilla were also not observed in the second instar. The lateral spiracular complex could not be seen in the sole spec- imen examined. The caudal segment, while not pictured, resembled the caudal segment in the third instar. Puparium.—Puparium creamy white —— second abdominal segment, | —spiracle, 2—verruciform sensillum; (G) caudal segment, spiracular plate, 1— rima, 2—interspiracular process; (H) caudal segment, compound sensillum, | —tuberculate chemosensillum, 2— stelex sensillum. Fig. 4. when newly formed, darkening to brown, or black with age; elongate-ellipsoidal; minute, dome-shaped verrucae remain distinct (Fig. 4A): 69 puparia averaged 3.14 + 0.05 (range, 2.3-3.9) mm in length, 1.28 + 0.02 (range, 0.90-1.59) mm in width; anterior end bears the invagination scar (Fig. 4B-1); anterior thoracic spiracles dorsolaterad of the in- vagination scar (Fig. 4B-2); posterior spi- racular plates bear three oval rimae ca. 0.02 mm in length (Fig. 4C-1), and four, thorn- like interspiracular processes, the longest measuring 0.01 mm (Fig. 4C-2); compound sensilla, i.e. a tuberculate chemosensillum (Fig. 4D-1) and a stelex sensillum (Fig. 4D- 2), ventrad of the spiracular plates were re- tained. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Puparium of V. californica: (A) habitus, anterior to left; (B) anterior end, 1 —invagination scar, 2— anterior thoracic spiracles; (C) caudal segment, posterior spiracular plate, |—rima, 2—interspiracular process; (D) caudal segment, compound sensillum, | —tuberculate sensillum, 2—stelex sensillum. Wangberg (1978) reported that an un- stated number of puparia of Valentibulla spp. averaged 3.74 (range, 3.2—4.0) mm in length, considerably longer than our mean and lower limit. Distribution and hosts The known distribution of V. californica includes the western third of the U.S. as mapped by Foote et al. (1993), who also noted that it had been collected in Baja Cal- ifornia Norte. Its known hosts are Chryso- thamnus nauseosus (Novak et al. 1967, Wasbauer 1972, Wangberg 1978, Foote et al. 1993) and C. parryi (Gray) Greene (Wangberg 1978), a high-altitude species in southern California (Munz 1974) which we VOLUME 97, NUMBER 3 have not examined for the presence of V. californica. Biology Egg.— The egg is inserted pedicel-last for up to two-thirds its length into a current or preceding year’s branch and within or near a prominent axillary bud. The axillary bud was used as a point of purchase on the smooth stems by the ovipositing female in insectary cagings, where as many as eight eggs were inserted into one axil. However, based upon dissections of field samples, an average of only 1.4 + 0.1 (range, 1-4) eggs (n = 80) were inserted singly into or near individual buds by one or more females. Larva.—Upon eclosion, the first instar tunnels apically (17%) or basally (83%) (n = 72) into the branch above or below the axillary bud, which subsequently may or may not be killed by larval feeding. The tunnel increases in diameter as the larva grows and deeply scores the woody pith (Fig. 5A), sometimes reaching the opposite epi- dermis (Fig. 5C), and thus killing the distal part of a thin branch. In most cases, how- ever, the branch above the larva continues its growth apparently unharmed; accord- ingly, 45 larval tunnels were found upon dissection to begin an average of 10.1 + 0.9 (range, 1.7—24.5) cm below the apical mer- istems in live branches. The initial, narrow- est part of the larval mine became packed with frass for a distance of 2.3 + 0.1 (range, 0.8-8.0) mm (n = 78) (Fig. 5C). The re- mainder of the tunnel was open and formed the cell within which the larva fed, presum- ably on sap conducted to and expressed from the surrounding woody tissues; these exca- vations are not much larger than the fully grown larvae and puparia (Fig. 5C, D). This type of larval development closely resem- bles that of Tephritis arizonaensis in its branch-tip mines on the woody shrub, Bac- charis sarothroides Gray (Goeden et al. 1993). It also resembles Aciurina thoracica, the larvae of which feed within small branch- tip galls on B. sarothroides (Headrick and 555 Goeden 1993), because of the small exca- vations formed by both species. However, unlike A. thoracica and contrary to Wang- berg (1978), we could not bring ourselves to call the feeding sites of V. californica ‘galls.’ Instead, like Foote et al. (1993), we prefer the description, slight “‘swelling,” or better yet, as Dodson (1987b) described the “gall” of V. dodsoni Foote “‘a rather incon- spicuous swelling” of a branch, because this is the only external sign of the larva or pu- parium within (Fig. 5A, B, C, D, E). Neither do our observations agree with the descrip- tion for V. californica by Wangberg (1978, p. 481) that its ““Galls are usually polythal- mous and contain 2-6. . . larvae. Each larva is housed in a cavity that is separated from the others by a tissue wall.’’ Further dis- cussion of this matter follows in the next subsection. Before pupariation, the fully grown larva of V. californica excavates a circular, epi- dermal window above one end of its cell (Fig. 5G). The larva then pupariates facing the window either basally (36%) or apically (63%, n = 78) along the branch. Pupa.—Eighty-seven full-size cells con- taining puparia (Fig. 5D, E) measured 4.6 + 0.1 (range, 2.6—7.1) mm in length and 1.3 + 0.02 (range, 0.4—1.7) mm in width. Thirty of these cells were found within branches that averaged 2.2 + 0.2 (range, 0.8—4.4) mm in diameter. The branch diameters just above (n = 23) and just below (n = 30) these cells differed from the aforementioned stem diameters at the middle of these cells by only 0.4 + 0.06 (range, 0.0-1.0) mm and 0.3 + 0.05 (range, 0.0-1.1) mm, respec- tively. Masked by normal variations in thickness of uninfested branches and stems of C. nauseosus at our study site, and by the conspicuous galls caused by other insects, e.g. Aciurina trixa Curran (Wangberg 1981, Dodson and George 1986, Dodson 1987b) and several species of Cecidomylidae (Gagné 1989), the slight swellings symptomatic of V. californica are, indeed, “‘inconspicuous.” We further suggest that the gall of Valenti- 556 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Life stages of V. californica: (A) young larva in branch of Chrysothamnus nauseosus; (B) branch where fully grown larva in (C) was found; (D) saggital section of branch with intact puparium in cell facing exit hole still covered by epidermal window; (E) two puparia in cells within same branch; (F) puparium of V. californica formed beneath gall of Aciurina trixa; (G) exit hole in branch; (H) female adult; (1) mating pair. Line = 1 mm. VOLUME 97, NUMBER 3 bulla spp. pictured by Wangberg (1978) and his description of it quoted above, instead probably refer to a misidentified cecido- myliid gall. This interpretation also may ex- plain differences in the galls described for V. dodsoni by Dodson (1987a, b) and for Valentibulla spp. by Wangberg (1978). Moreover, V. californica larvae and puparia readily were found in juxtaposition to galls of Cecidomyiidae in branches during the present study, and during our laboratory dissections occasionally were located be- neath or near galls of A. trixa (Fig. 5F, un- published data). Another ambiguity is the report by Novak et al. (1967) that V. steys- kali Foote [as munda (Coquillett)] larvae form “large, polythalmous galls’ on stems of C. nauseosus in Idaho. These same au- thors reported V. californica from flower heads of C. nauseosus in Idaho, where Wangberg (1978) conducted his study. Nev- ertheless, after 3 years of study, we remain unable to approach individual plants of C. nauseosus known from past dissections to harbor V. californica at our main study site and with any measure of assurance select branches infested with this tephritid. This alone attests to the inconspicuousness of immature V. californica in situ in southern California. Adult.— Egress of the adult (Fig. 5H) en- tails breaking through the epidermal win- dow covering each exit hole (Fig. 5G). New- ly emerged females (Fig. SH) have imma- ture ovaries (n = 3), but within 2 weeks, the numerous, small, irregularly rounded, fat globules found in the haemocoel disappear and are converted into full-size ova. In in- sectary cagings, isolated, unmated, non-ovi- positing females (Fig. 5H) lived 45 + 2.8 (range, 37-56) days; isolated, unmated males lived 49 + 1.6 (range, 42-55) days. Wing displays.— Adults of V. californica held their wings parted with the posterior wing margins along the abdominal pleura, supinated 45°-60° relative to the substrate, and with the wing blade bent ventrally at the subcostal break in the costal margin. 557 Both sexes displayed synchronous and asynchronous supinations with their wing blades in the bent position and with ancil- lary rotations as described for Aciurina thor- acica (Headrick and Goeden 1993). Both sexes also displayed abdominal flexures during asynchronous supination, also de- scribed for A. thoracica, Paroxynagenalis (Thomson) (Goeden et al. 1994b), and Tru- panea californica (Headrick and Goeden 1991). Mounting, copulatory induction behav- ior, and copulation.—Males of V. califor- nica did not display aggregation behaviors such as abdominal pleural distention, wing displays, territoriality, and male-male com- bat as described for and commonly ob- served in other non-frugivorous tephritids (cf. Headrick and Goeden 1990, 1991, 1993, Goeden and Headrick 1991b, 1992b, Goe- den et al. 1994b, Headrick et al. 1994). Nor did they initiate courtship displays involv- ing movement of one or more body parts, also commonly observed in non-frugivo- rous tephritids in either laboratory or field observations (cf. Headrick and Goeden 1990, 1991, 1993, Goeden and Headrick 1991, 1992, Goeden et al. 1993). Males of V. californica oriented toward and stalked females from behind, then abruptly jumped onto their dorsa, as described for V. dodsoni and A. trixa males (Dodson 1987, Headrick and Goeden, unpublished data). As also ob- served for A. trixa, V. californica females immediately began rapid walking around the arenas after being mounted, as the males struggled to hold onto them with their front legs. Because mounting by males was so abrupt, they often landed on females in awkward positions that required re-posi- tioning themselves to successfully gain in- tromission. This readjustment also was ob- served with A. thoracica males (Headrick and Goeden 1993). After mounting and suc- cessfully holding onto the female, the male then curled the apex of his abdomen ven- trally and placed his epandrium at the apex of her oviscape. His front legs grasped the 558 widest portion of her abdomen and his mid- die i hind legs rested on the substrate. fhe mounted female used her hind legs to push vigorously at the male. The male ini- tiated copulatory induction behavior (CIB) immediately after positioning himself on the female. CIB involved the mounted male drumming vigorously on the venter of the abdomen of the female with his hind legs, unlike A. thoracica males who used their middle legs for CIB (Headrick and Goeden 1993). The receptive female exserted her aculeus which pushed the male backward to accommodate its length, because the male kept his abdomen curled ventrally. Inser- tion of the aedeagus through the ventral flap into the cloacal opening followed aculeus exsertion and the female began to slowly retract her aculeus as the male moved for- ward on the female to assume the copula- tory posture (Fig. 51). During copulation the female intermittently exerted pressure on her aculeus which in turn stimulated the male to begin agitation wing displays. The agitated male lofted his wings, as described for P. genalis (Goeden et al. 1994b), 45° above the long axis of his body, while slight- ly parted and the wing blades were supi- nated to 90°. Then, the wings were syn- chronously lowered ca. 10° and raised again without vibration once every half second. The male returned his wings to the typical resting position along his pleura when the agitation stimulus ceased. After ca. 1 h in copula, the female began to push at the male with her hind legs. This activity signaled the termination of copu- lation. Disengagement lasted ca. 1 min. The male lifted his abdomen and moved back- ward on the female; his hind legs were on the substrate and his middle legs were around her ovipositor. He continued to raise his abdomen, disengaged his surstyli from the aculeus, and pulled his aedeagus from the ventral flap while he remained mounted on the female. This disengagement was sim- ilar to that described for P. genalis males (Goeden etal. 1994b). After disengagement, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the male moved forward on the female and with the aid of his hind legs recoiled the aedeagus. Males initiated CIB behavior in laboratory cagings after ca. 1 min by grasp- ing the ovipositor and drumming it vigor- ously with the middle legs. Copulations were commonly repeated by the same individu- als in less than 1 h; however, no more than two copulations were observed in a single day. After disengagement, females began turning rapidly in circles and resumed buf- feting the mounted males with their legs and wings. This latter behavior continued until the male either lost his grip and fell off the female or she again exserted her aculeus. After a second copulation, males fully dis- mounted from females, which typically dis- played aggression by synchronously extend- ing their wings and lunging at males. The only copulation timed from start to finish lasted 1 h. Field observations.—Adults of V. cali- fornica were observed on C. nauseosus plants along with adults of Aciurina trixa and Pro- cecidochares minuta Snow at the Cajon Junction study site. Adults of A. trixa were prevalent at the start of field observations on 8.iv.1992, but diminished in numbers as densities of P. minuta increased to 12.v.1992. A total of only eight adults of V. californica was observed during all 4 weeks. Males were observed singly on host plants usually in the interior of the crowns and at rest on stems; all three females were ob- served while ovipositing. No encounters be- tween the sexes were observed. Females Oviposited into leaf axils apparently utiliz- ing the same ovipositional niche as A. trixa and P. minuta females (Headrick and Goe- den unpublished data); however, their ovi- position behavior was different from these two species. Valentibulla californica fe- males did not climb to the tops of branches before ovipositing in the leaf axils as ob- served for both A. trixa and P. blantoni (Headrick and Goeden, unpublished data), but rather explored for oviposition sites on the lower portions of branches. VOLUME 97, NUMBER 3 Wangberg (1978) reported that adults of Valentibulla. spp. on C. nauseosus in Idaho were active on host plants throughout the day and displayed their wings toward con- generic individuals. He further described fe- males as vying for the attention of males, which then pursued the females by follow- ing behind them. The male then continued to approach the female and if she stood still, he moved behind her and tapped her with his front tarsi. This initial tracking and con- tact lasted from 1-2 s to 1-2 h. Receptive females remained motionless and males then mounted them. Wangberg (1978) noted that males bent their abdomens ventrally in “‘an effort to penetrate (the female) with his gen- italia.” He also reported that copulations lasted 60-90 min, but did not note any dis- engagement behavior or repeated copula- tions by males. Finally, Wangberg (1978) reported that females oviposited under the epidermis of stems rather than into leaf axils as observed in the present study. Seasonal history.—Valentibulla califor- nica 1s univoltine and nondiapausing on C. nauseosus in southern California. Adults emerge in late winter and early spring (Feb- ruary—April) and mating and oviposition commence shortly thereafter; both latter be- haviors continue for about |'2 months. The eggs hatch in about a week and the larvae feed and grow slowly through late-spring, summer, and fall (April-November) with pupariation occurring in winter (Decem- ber—March), as the cycle resumes. Natural enemies.— Three species of chal- cidoid Hymenoptera were reared as pri- mary, solitary endoparasitoids from indi- vidual puparia of V. californica dissected from C. nauseosus: Eupelmus sp. (Eupel- midae), Eurytoma sp. (Eurytomidae), and Halticoptera sp. (Pteromalidae). Wangberg (1978) reported an Eupelmus sp. and Hal- ticoptera sp. as associates of Valentibulla spp. galls, but also Platygaster sp. (Hyme- noptera: Platygasteridae), known parasit- oids of Cecidomyiidae (Clausen 1962) and never reared by us from any Tephritidae in 559 southern California to date (unpublished data). ACKNOWLEDGMENTS We thank A. C. Sanders, Curator of the Herbarium, Department of Botany and Plant Sciences, University of California, Riverside, for identification of plants. Par- asitoids were identified by comparison with specimens identified by J. LaSalle, J. Luh- man, and C. M. Yoshimoto during their visits or while otherwise associated with the Department of Entomology, University of California, Riverside. We also are grateful to F. L. Blanc and G. Dodson for their help- ful comments on earlier drafts of this paper. LITERATURE CITED Carroll, L. E.and R. A. Wharton. 1989. Morphology of the immature stages of Anastrepha ludens (Dip- tera: Tephritidae). Annals of the Entomological Society of America 82: 201-214. Clausen, C. P. 1962. Entomophagous Insects. Mc- Graw-Hill Book Co., Inc., New York and London. Reprint Edition, Hafner Pub. Co., New York. Coquillett, D. W. 1894. New North American Try- petidae. Canadian Entomologist 26: 71-75. Dodson, G. 1987a. Biological observations on Aci- urina trixa and Valentibulla dodsoni (Diptera: Te- phritidae) in New Mexico. Annals of the Ento- mological Society of America 80: 494-500. 1987b. Host-plant records and life history notes on New Mexico Tephritidae (Diptera). Pro- ceedings of the Entomological Society of Wash- ington 89: 607-615. Dodson, G. and S. B. George. 1986. Examination of two morphs of gall-forming Aciurina (Diptera: Te- phritidae). Ecological and genetic evidence for spe- cies. Biological Journal of the Linnean Society 29: 63-79). Foote, R. H., F. L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York. Gagné, R. J. 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ith- aca. Goeden, R. D. and D. H. Headrick. 1990. Notes on the biology and immature stages of Stenopa affinis Quisenberry (Diptera: Tephritidae). Proceedings of the Entomological Society of Washington 92: 641-648. . 199la. Notes on the biology, hosts, and im- 560 mature stages of Tomoplagia cressoni Aczél in southern California. Proceedings of the Entomo- logical Society of Washington 93: 549-558. —. 1991b. Life history and descriptions of im- mature stages of Tephritis baccharis (Coquillett) on Baccharis salicifolia (Ruiz & Pavon) Persoon in southern California (Diptera: Tephritidae). Pan- Pacific Entomologist 67: 86-98. 1992. Life history and descriptions of im- mature stages of Neaspilota viridescens Quisen- berry (Diptera: Tephritidae) on native Asteraceae in southern California. Proceedings of the Ento- mological Society of Washington 94: 59-77. Goeden, R. D., D. H. Headrick, and J. A. Teerink. 1993. Life history and descriptions of immature stages of Tephritis arizonaensis Quisenberry (Dip- tera: Tephritidae) on Baccharis sarothroides Gray in southern California. Proceedings of the Ento- mological Society of Washington 95: 210-222. 1994a. Life History and descriptions of im- mature stages of Procecidochares flavipes Aldrich (Diptera: Tephritidae) on Brickellia spp. in south- ern California. Proceedings of the Entomological Society of Washington 96: 288-300. 1994b. Life history and description of im- mature stages of Paroxyna genalis (Thomson) (Diptera: Tephritidae) on native Asteraceae in southern California. Proceedings of the Entomo- logical Society of Washington 96: 612-629. Green, J. F., D. H. Headrick, and R. D. Goeden. 1993. Life history and description of immature stages of Procecidochares stonei Blanc & Foote on Viguiera spp. in southern California. (Diptera: Tephriti- dae). Pan-Pacific Entomologist 69: 18-32. Headrick, D. H. and R. D. Goeden. 1990. Descrip- tion of the immature stages of Paracantha gentilis (Diptera: Tephritidae). Annals of the Entomolog- ical Society of America 83: 220-229. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1991. Life history of Trupanea californica Malloch (Diptera: Tephritidae) on Gnaphalium spp. in southern California. Proceedings of the En- tomological Society of Washington 93: 559-570. 1993. Life history and description of im- mature stages of Aciurina thoracica (Diptera: Te- phritidae) on Baccharis sarothroides in southern California. Annals of the Entomological Society of America 86: 68-79. Headrick, D. H., R. D. Goeden, and J. A. Teerink. 1995. Life history and description of immature stages of Euaresta stigmatica (Diptera: Tephriti- dae) on native ragweeds (Ambrosia spp.) in south- ern California. Annals of the Entomological So- ciety of America 88: 55-71. Munz, P. A. 1974. A Flora of Southern California. University of California Press, Berkeley. Novak, J. A., W. B. Stoltzfus, E. J. Allen, and B. A. Foote. 1967. New host records for North Amer- ican fruit flies (Diptera: Tephritidae). Proceedings of the Entomological Society of Washington 69: 146-148. Wangberg, J. K. 1978. Biology of gall-formers of the genus Valentibulla (Diptera: Tephritidae) on rab- bitbrush in Idaho. Journal of the Kansas Ento- mological Society 51: 472-483. . 1981. Gall-forming habits of Aciurina species (Diptera: Tephritidae) on rabbitbrush (Composi- tae: Chrysothamnus spp.) in Idaho. Journal of the Kansas Entomological Society 54: 711-732. Wasbauer, M. W. 1972. An annotated host catalog of the fruit flies of America north of Mexico (Dip- tera: Tephritidae). California Department of Ag- riculture, Bureau of Entomology, Occasional Pa- pers 19: 1-72. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 561-568 A BASAL NEW SPECIES OF OLBIOGASTER (DIPTERA: ANISOPODIDAE) IN DOMINICAN AMBER, AND ITS SYSTEMATIC PLACEMENT Davip A. GRIMALDI AND DALTON DE SouZA AMORIM (DAG) Entomology Department, American Museum of Natural History, Central Park West at 79th St., New York, New York 10024-5192; (DSA) Departamento de Biologia, Faculdade de Filosofia, Ciéncias e Letras de Ribeirao Preto/USP, Av. Bandeirantes 3900, 14040-901 Ribeirao Preto, SAo Paulo, Brazil. Abstract.—A distinctively new species of O/biogaster is described from Oligo-Miocene amber of the Dominican Republic. It is based on a single, virtually complete female and named Olbiogaster perezi. Phylogenetic analysis of external morphological characters indicates it is probably the sister group to all of the living species of the genus, or at least among a group of phylogenetically basal species. The four living species of Olbiogaster described from Caribbean islands belong to clades that are more recently derived than is the fossil. Key Words: The family Anisopodidae, in the broad sense, is a small family of relatively prim- itive flies that is of keen phylogenetic inter- est. Only about 100 species are described for the family, which is probably no more than half of the actual number of species. The larvae occur under bark of decaying trees, rotting vegetation, and associated with fungi. Major, synthetic papers on the sys- tematics of the family are by Amorim and Tozoni (1994) and Tozoni (1989). The for- mer is a phylogenetic classification of the world genera of Anisopodidae, s.1.; the latter is a phylogenetic study of the species of O/- biogaster, from which we drew heavily for characters to study. This is the second paper on Dominican amber Anisopodidae. The first one, by Gri- maldi (1991), described four species: Va- leseguya disjuncta, Mycetobia antillea, My- cetobia cryptambra, and Mesochria neotro- pica. The last three species Amorim and Tozoni (1994) are placing in a new genus, Anisopodidae, amber, O/biogaster, cladistics Neomesochria, with all the Neotropical spe- cies formerly put into Mycetobia as well as two New Caledonian species. Valeseguya disjuncta is of great interest because it 1s a relict: its closest living relative is Valese- guya rieki from Australia, the only other member of this aberrant genus. Amorim and Tozoni (1994) placed Valeseguya in a nar- rowly defined family, the Mycetobiidae (in- cluding also Mycetobia, Mesochria, and Neomesochria). The placement of Valese- guya with mycetobiines is currently being tested using DNA sequences from the am- ber fossils (DeSalle and Grimaldi, in prep.). Valeseguya disjuncta is curiously much more common than the other mycetobiines in Dominican amber (even though the genus is now restricted to Australia). Also, the ex- tinct species is represented by a large series of males and females; the one living in Aus- tralia is represented by a few poorly pre- served males. The finding of an O/biogaster in Dominican amber now adds a major new 562 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON component to this extinct fauna of aniso- podids. METHODS AND MATERIALS We are indebted to J. Manuel Perez for providing the specimen to us for study. Ex- act provenance within the Dominican Re- public is uncertain but derives from the group of central mines in the mountains surrounding Santiago. Despite repeated ci- tations of Dominican amber being Eocene in age (ca. 40 myo), there is very little basis for that dating. Grimaldi (1995) reviewed the stratigraphic evidence. At present it is most prudent to assume a Miocene-Oligo- cene age (ca. 25 million years old). The spec- imen was prepared and photographed as given in Grimaldi (1991, 1993). SYSTEMATICS Genus Olbiogaster Osten-Sacken, 1886 Tozoni (1989) and Amorim and Tozoni (1994) provided diagnostic characters of the genus. These included several aspects of male sternite 10 (bifid apically, with a pair of lateral projections, and slender at the base but apically enlarged) and the gonostyles (slender, with setae on the outer face). The most obvious feature that distinguishes this genus is vein M, weaker than the other pos- terior veins. The genus Olbiogaster is restricted to the New World, with all but two species being Neotropical. Tozoni (1989) indicated that there were 7 undescribed species from Bra- zil. Only 4 species are described from the Caribbean: antillarum Lane & D’Andretta (U.S. Virgin Islands), fu/va Edwards (Puer- to Rico), danista Stone and evansi Stone (island of Dominica, Lesser Antilles). Olbiogaster perezi Grimaldi and Amorim, NEw SPECIES Fig. 1 Diagnosis.—Head and thorax with dis- tinctive pattern of dark brown areas (see description). Wing with stigma surrounding basal half of vein R,,;. Discal cell fusiform, ends not truncate, with medial veins branching directly off apical point. Vein M, lost distad of discal cell. Tergites and ster- nites mostly blackish-brown. Description. — Body coloration: With ex- tensive dark brown areas, including: clypeus and antennal bases; gena, postgena; labium; antenna (except the scape and pedicel); postpronotal lobe; notum, posterior half of scutellum; anepisternum and _ katepister- num; anterior and posterior basalares; por- tion of the anatergite; katatergite; coxae and trochanters (mid coxa less so); hind femora and tibiae; all tergites and sternites. Head: Antenna with 14 cylindrical seg- ments; pedicel very short. Eye large, com- pletely bare. Palp with 5 segments; basal 2 almost circular, third segment longest and widest; 2 apical segments equal to length of segment 3 but half the thickness. Clypeus protruding, extended to apex of palp seg- ment 3. Labellum large, laterally flattened. Gena, postgena, and labium with numer- ous, fine, whitish setae. Occiput with 6 stiff, black setae. Thorax: Notum covered with fine, whit- ish, decumbent acrostichal setulae. Post- pronotal lobe large, with ca. 30 fine, whitish setae. Anepisternum heavily sclerotized, with patch of 20 fine, whitish setae on dorsal lobe; rest of anepisternum bare. Katepister- num completely bare. Anterior and poste- rior basalare heavily sclerotized. Anepi- meron lightly sclerotized, with small patch of ca. 10 fine, whitish setae on postero-dor- sal portion. Katatergite sclerotized, bare. Legs: Fore and midlegs yellow, apices of tibiae and tarsi brown. All legs with setulae irregularly arranged (not in longitudinal rows); tibiae and tarsi with 3—4 longitudinal rows of stiff, stout, black setae. Comb on apex of hind tibia not visible: lost with tarsi at surface of amber. Wings: Membrane with microtrichia only, arranged irregularly, not in rows. Wing veins dark; radial veins darkest. Costal vein VOLUME 97, NUMBER 3 Fig. 1. extended to slightly past R,,;. Sc and R, meeting C very close together. Dark stigmal area surrounding apical half of R,,3;. Ry; extended to tip of wing. Crossvein r-m very short, connected to middle of discal cell (d); cell d is fusiform in shape, ends not truncate. Veins M, and M, arising from apical point of cell d, vein M, absent distad of discal cell. 563 Paexe eee Xx ; A Aj l SE BLS ee REET, Habitus of holotype of O/biogaster perezi, n. sp. Crossvein cu-m very short, proximal to r-m, branch of CuA, and CuA, almost perpen- dicular to longitudinal axes of veins. Vein A, incomplete, A, obscured. Halter light yellow. Abdomen: Tergites and sternites dark brown; sternites large, covering pleural area. Apical sternite light, elongate, with deep 564 V-shaped notch in middle. Cerci long, nar- TOW, Gark. Holotype: Female, AMNH-DR11-657, in a clear, light yellow piece of amber, 10 x 13 mm; exact provenance within the Do- minican Republic unknown. The surface of the piece on the fly’s right side has been ground and polished flat. The specimen 1s virtually complete, except for loss of the mid and hind tarsi. Etymology: Patronym, for J. Manuel Perez, an amber dealer in Orlando, Florida, who made this specimen available for study. RELATIONSHIPS Tozoni (1989) presented a phylogenetic analysis of the species of Olbiogaster, in- cluding 21 described and 7 new species (the new species are actually described in Tozoni [1994], with an additional 6 species). Her phylogeny of the genus included four major subgroups widely distributed in the Neo- tropical region. The genus is defined here in the sense of Amorim and Tozoni (1994), diagnosed above. Features used in her anal- ysis were derived mostly from male ter- minalia and body coloration. Sexual di- morphism in sclerite colors exists, and not all species are known by both sexes, so sep- arate cladograms had to be developed for males and females. In both males and fe- males there occurred transformation series where the color of sclerites changed from brown to ocher or yellow. Olbiogaster has most sclerites brown, ex- cept part of the scutellum, epimeron II and capitulum of the halter. It is thus clear that O. perezi is not a recently derived species in the genus. The basal levels of Tozoni’s (1989) phylogeny for the recent species is completely resolved into five clades. Char- acter incongruence was evaluated using the ACCTRAN and DELTRAN procedures, which allowed for reversals and multiple origins of characters, respectively (Figs. 2, 3). The reversals in the cladogram of Fig. 2 do not imply reacquisition of complex structures, so this shorter cladogram is the preferred explanation. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON We present below Tozoni’s (1989) list of the features that can be examined in the fossil species (e.g. external female), and which refer to the level of generality of this discussion; the character matrix is present- ed in Table 1. Note that O/biogaster mex- icana is known from Mexico, O. danista from Dominica, O. erythrohydra and O. ta- moioi are from southeastern Brazil; the re- maining species, in the marinonii group, oc- cur throughout the Neotropical Region (ex- cept Chile and southern Argentina). Ple- siomorphic states are given first for each character. Use of a superscript cross (*) de- notes a monophyletic group plus its sister group(s). This is a convention established by Amorim and used elsewhere (e.g., Amo- rim and Tozoni, 1994). 1. Scutum entirely brown/scutum yellow or ochre. The plesiomorphic condition of this char- acter in Olbiogaster is found only in O. per- ezi. This is one of the features that indicates O. perezi is the probable sister group of all the remaining species of the genus. Most species of Eogaster, Austrogaster, and OI- biogaster have a brown scutum, leaving no doubt about the polarity of the character. 2. Scutellum entirely brown/scutellum entirely yellow or ochre. 3. Scutellum entirely brown/scutellum yellow below. The polarization of these characters 1s also unquestionable. An entirely brown scutel- lum is known to occur only in O. danista, from Dominica. Olbiogaster perezi is ple- siomorphic for the color in the dorsal part of the scutellum, but has an apomorphic color for the ventral surface of the scutel- lum. Character two also indicates that O. perezi would be the sister group of the re- mainder of the genus. 4. Anepisternum entirely brown/anespi- sternum entirely yellow or ochre. The plesiomorphic condition of this char- acter is found in O. perezi and O. erythro- VOLUME 97, NUMBER 3 mexicana O. erythrohydra O. tamoioi O. marinoniit+ O. danista O. 1, 7, lla, 12a length = 23 steps Figs2.)3: 565 3 3 an 3 =, “to =. - oes 3 5 Fa ~ * fap ined a ey * om ai = na . pee F = 5 Sy = S SoS S 8,9, 10, 12b 1,6, 7, lla, a length = 27 steps Basal relationships of Olbiogaster as proposed by Tozoni (1989), with two interpretations for the transformation series. 2, ACCTRAN procedure for all characters, allowing for character reversals (shaded characters). 3, DELTRAN procedure for all characters, allowing for multiple origins of characters (characters placed above in the cladogram). hydra. Some species belonging to the mono- phyletic marinonii*+ group have brown maculae on the anepisternum, but character optimization shows that the ground plan of this group should be apomorphic for these characters (species with maculae being cases of reversals). An ACCTRAN optimization of this character results in two steps, a DEL- TRAN approach produces four. However, neither approach affects the position of O. perezi in the cladogram, since the mono- phyly of erythrohydra + tamoioi is well sup- ported and unambiguous. 5. Katepisternum brown/katepisternum entirely yellow or ochre. The plesiomorphic condition of this char- acter is found in O. perezi, as well as in O. mexicana and O. erythrohydra. As in the preceeding character, some species of the marinonii* group also have brown maculae on the katepisternum, but the group is apo- morphic in its ground plan. On this basis Table |. Olbiogaster. Character matrix for the basal groups of Character 1 1 ae 1 1 12345 67890 2345 Taxon Outgroups 00000 00000 00000 O perezi 00100 10000 00000 1 mexicana 11010 01000 10000 1 erythrohydra 11000 11111 21002 1 tamoiol LOM Te 210021 danista gr. 10011 11111 21000 0 marinoniil gr. HOM Ea Ord 566 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2 oS} SE 3 ot > 1 i] ha} Se = SS .. = S's 3 S Se = Soe = tm B poe aS) mB ‘~ & & = 5-3 rE bs] 9 sey t BES 8 25 iS} 3 a & § 3 5 rss = Soy S ES ia vxuY § S079 Sr ‘Si 7 -O7S19) 9 1,2,4, 7, lla, a, 16 length = 29 steps length = 24 steps Figs. 4-7. = 3 "3 oe x S00 3 oe x Lay ee 5 S88 S oy IN hss) ee * iS) S-2- BS = Se ee eS iS Q AS a4 2 my ef { 22k s = 8 wig S =~ a 8 iS} See SES iS sSSuaq § SSS Se08 S "SGc'S gs 1,2, 4,6, 7, lla, 12a 1,2, 4,6, 7, lla, 12a, 16 6 1 length = 31 steps length = 35 steps Four topologies for the relationships of O. perezi with remaining species of the genus. 4, Characters 1, 2, 4, 7, lla, and 12a are taken as true synapomorphies, so O. perezi would be the sister group of the rest of the genus. 5, Characters 5, 6, 8, 9, 10, and 12b are considered true synapomorphies, so O. mexicana becomes the sister group to the rest of the genus, with O. perezi as the next plesiomorphic sister group. 6, Assumption of character 16 having multiple origins and the apomorphic condition shared by O. perezi and mexicana interpreted as a true synapomorphy. 7, Reversal of character 2 interpreted as a true synapomorphy, which links danista and perezi, and many plesiomorphic conditions in the latter species thus becoming reversals. the fossil species is excluded from the mar- inonil* group. 6. Epimeron II entirely yellow/epimeron II entirely brown. Olbiogaster perezi has epimeron II en- tirely yellow (sometimes referred to in the Anisopodidae as the anepimeron). Olbio- gaster mexicana, on the other hand, has the sclerite entirely brown (based on the pub- lished description) and some species of the marinonii* group also have epimeron II maculated brown. The fact that O. perezi shares the apomorphic condition with the remaining species of the genus except O. mexicana indicates that it could belong to the danista** group (Figs. 5-7), a topology different from that suggested by characters 1, 2, 4, 7, and 12a (Figs. 4-6). 7. Mediotergite entirely brown/medioter- gite entirely yellow or ochre. Olbiogaster perezi is the only species in the genus to have the mediotergite entirely brown. This would indicate a sister-group relationship between O. perezi and the rest of the genus. 8. Coxa I entirely brown/coxa I entirely yellow or ochre. 9. Coxa II entirely brown/coxa II entirely yellow or ochre. 10. Coxa III entirely brown/coxa III en- tirely yellow or ochre. Olbiogaster mexicana is the only species of the genus sharing with O. perezi the ple- siomorphic condition of this character. Al- though the two species cannot be linked on the basis of this symplesiomorphy, it cor- roborates that O. perezi does not belong to the danista group. 11. Femur III entirely brown/a. with distal 4 brown / b. entirely yellow or ochre. The presence of an entirely brown femur III in Olbiogaster is exclusive to O. perezi. At this level of the analysis, the most apo- morphic condition, of femur III being en- tirely yellow, is found only in the species of the danista* group, and occuring homopla- siously in the marinonii* group. The first step corroborates the hypothesis of O. perezi being the sister group to the remaining O/- biogaster. VOLUME 97, NUMBER 3 12. Tergites I-IV entirely brown/a. with distal 4% yellow / b. with distal ' yel- low. 13. Tergites I-IV entirely brown/with a median yellow line on distal 7. 14. Tergites I-IV entirely brown/a. with a yellow band laterally on distal 7. / b. with a complete yellow band. 15. Tergite V entirely brown/with a median yellow line on distal 7. A complex transformation series 1s pro- posed by Tozoni (1989) for the coloration of the abdominal tergites in Olbiogaster, varying from brown to yellow. These mod- ifications include independent changes in different areas of the sclerite (distally, lat- erally, and mesally), and with homoplasious origins of some of these changes. O/biogas- ter perezi is the only known species with the tergites entirely brown. Olbiogaster mexi- cana has only the distal fifth of tergites I- IV yellow and other species have a larger portion of the tergites yellow. This, once more, confirms the basal position of O. per- ezi in the genus. 16. Capitulum of halter brown/yellow. Olbiogaster has the capitulum yellow, as do other basal species of the genus with the exception of the O. danista and some spe- cies belonging to the marinonii* group. If the apomorphic condition of the character is attributed to the ground plan of the genus, the plesiomorphic condition of this char- acter in O. danista would be a reversal. Of the list above, O. perezi is apomorphic only for characters 3, 6, and 16. Character 3 is an autapomorphy. Character 6 is shared with all Olbiogaster except O. mexicana and character 16 is shared with most species of the genus, but is plesiomorphic in O. dan- ista. The plesiomorphic conditions of char- acters 13 and 14 exclude O. perezi from the marinonii group. The topology of the most parsimonious cladogram has O. perezi as the sister group of all remaining known re- cent Olbiogaster (Fig. 4). The apomorphic 567 features of O. perezi, however, allow for some alternative hypotheses. O/biogaster perezi could be the sister group of all recent species of the genus except O. mexicana, which in this case would be the most basal species of the genus (Fig. 5). This hypothesis would be obtained if we consider the yellow epimeron II of O. perezi (character 6) ho- mologous to the same feature seen in other species of the genus. Olbiogaster perezi is theoretically the sister species of O. mexi- cana, if we consider the yellow halter ca- pitulum to be homologous among them (Fig. 6). Finally, O. perezi could be the sister group of O. danista if we accept character 6 as having a single origin in the group and the brown scutellum (character 2) of O. perezi and O. danista being homologous (Fig. 7). Still other topologies present even less par- simonious character evolution. The length of the cladogram in which O. perezi is the sister group to the remaining species of the genus has 24 steps, five steps shorter than the second most parsimonious cladogram. The other two cladograms have 32 and 35 steps, respectively. A sister-group relationship of Ol/biogaster perezi to the other species of the genus adds a basal stem to this monophyletic group. The autapomorphies shown by O. perezi— particularly the virtual absence of vein M,— indicate that the fossil is itself a derived clade arising mostly closely from the stem group, but not an ancestral member of the genus. LITERATURE CITED Amorim, D. S. and S. H. S. Tozoni. 1994. Phylo- genetic and biogeographic analysis of the Aniso- podoidea (Diptera, Bibionomorpha), with an area cladogram for intercontinental relationships. Re- vista brasileira de Entomologia 38(3/4): 517-543. Grimaldi, D. A. 1991. Mycetobiine woodgnats (Dip- tera: Anisopodidae) from the Oligo-Miocene am- ber of the Dominican Republic, and Old World affinities. American Museum Novitates 3014, 24 pp. . 1993. The care and study of fossiliferous am- ber. Curator 36: 31-49. 1995. On the age of Dominican amber. /n PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 568 kK. B. Anderson and J. C. Crelling, eds., Amber, . 1994. Revision of the genus Olbiogaster Os- esinites, and Fossil Resins. American Chemical ten-Sacken (Diptera: Anisopodoidea: Olbiogastri- society Symposium Volume (Washington, D.C., dae). I. Description of 13 new species, taxonomical Sug. 1994). notes, and key to the Neotropical species. Revista Tozoni, S. H.S. 1989. Analise filogenética do género nordest Biologia 7 (In press.) Olbiogaster (Diptera: Anisopodidae). M.Sc. thesis, Universidade Federal da Paraiba, Joao Pessoa. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 569-581 THE NORTH AMERICAN SPECIES OF SYSTASIS WALKER (HYMENOPTERA: PTEROMALIDAE) S. L. HEYDON Bohart Museum, Department of Entomology, University of California, Davis, Cali- fornia 95616. Abstract.—The nearctic species of Systasis Walker are reviewed. These include two holarctic species, S. encyrtoides Walker and S. parvula Walker, and three new nearctic species—S. aquila, n. sp., S. hansoni, n. sp., and S. tena, n. sp. A summary of the biology and distribution for the genus and for each species is presented. Key Words: Systasis Walker, 1834, and Semiotellus Westwood, 1839, were placed together in a new tribe, the Systasini (Ormocerinae), by Bouéek (1988). Systasini is defined on the basis of reduction in the number of antennal segments to 12 with at least the two basal flagellar segments anelliform (Figs. 4, 6, 8- 11), reduction of the number of teeth on each mandible to 3 (Fig. 2), and the uniquely structured clypeus—quadrate, nearly flat, with the anterior margin nearly straight across, and with deep anterior tentorial pits in the dorsolateral corners (Figs 1, 2). Two other unusual characters present in most species of these two genera and which need more study of their distribution are 1) the tendency of the mesepisternum to be non- metallic, varying from almost black to dull yellowish brown depending on the species, and 2) the presence of a narrow membra- nous fovea basally on the dorsolateral cor- ner of the gaster. Systasis and Semiotellus are very similar genera, and if the morpho- logical variation found in the Nearctic spe- cies described herein is taken into account, Systasis species are distinguished from those of Semiotellus only in having 1) the spec- ulum of the upper surface of the wing ex- Systasis, Semiotellus, Pteromalidae, Ormocerinae, Cecidomyiidae, gall, blueberry, biology, new species tending down to the stigmal vein, making the line of strong setae on the ventral side of the wing membrane just posterior to the marginal vein easily visible and 2) the free anterior margin of the clypeus nearly straight across (Fig. 2). Semiotellus species have the dorsal surface of the fore wing setose pos- terior to the marginal vein and the free an- terior margin of the clypeus truncate. It is probable that these two genera will someday be regarded as subgenera of Systasis once a worldwide survey of the species is com- pleted. Systasis was originally created for the spe- cies S. encyrtoides Walker and S. tenuicornis Walker. Current authors recognize eight ad- ditional palearctic species—S. acuta (Fon- scolombe), S. annulipes (Walker), S. an- gustula Graham, S. basiflava Graham, S. ephedrae Dzhanokmen, S. /ongula Bouéek, S. parvula Thomson, and S. tenuicornis Walker; four Oriental species—S. cenchri- vora Farooqi & Menon, S. dalbergiae Mani, S. dasyneurae Mani (key to these three spe- cies in Farooqi and Menon 1972), and S. vischnu (Motschulsky); 21 Australian spe- cies—S. australiensis (Girault), S. cecido- myiae (Ashmead), S. cecili Girault, S. cer- Figs. 1-6. nus (Walker), S. dice (Walker), S. doddi (Gi- rault), S. euctemon (Walker), S. flindersiae Girault, S. graminis (Cameron), S. grotiusi Girault, S. henrici Girault, S. horridula (Gi- rault), S. insularis Dodd & Girault, S. keatsi Girault, S. lelex (Walker), S. merula (Walk- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SD 5@U 8813 1+4,Systasis parvula, female. 1, Head, anterior view. 2, Clypeus. 3. Whole body. 4. Antenna. 5- 6, Systasis aquila, female. 5, Whole body. 6, Antenna. er), S. persimilis (Dodd & Girault), S. punc- tativertex Girault, S. seposita Girault, S. varipes Girault, and S. viridis (Girault); and three Ethiopian species—S. afra Masi, S. coerulea Delucchi, and S. guierae (Risbec). There are no described species from the VOLUME 97, NUMBER 3 Neotropical Region, and based on collec- tions I have examined, they do not seem to occur there. The latest catalog of nearctic Pteromali- dae (Burks 1979) lists two species for North America—S. encyrtoides and S. diplosidis Eckel. The latter species was transferred to Melancistrus Graham (Heydon and Bouéek 1992). The palearctic species S. parvula Thomson is reported from the New World for the first time herein, and three new nearctic species are described—S. aquila, n. sp., S. hansoni, n. sp., and S. tena, n. sp. Terminology in this paper generally fol- lows that of Graham (1969), except that genal concavity is used instead of genal hol- low and club is used instead of clava. In addition, the gastral tergites are numbered T1-T7 beginning with the first tergite after the petiole. The following abbreviations are used: the median ocellar diameter is MOD, the ocellar-ocular distance is OOL, the pos- terior ocellar distance is POL, the lateral ocellar distance is LOL, the multiporous plate sensilla are MPP sensilla, the lower ocular line is LOcL, and the antennal fu- nicular segments are Fl through F6. The units of measurement given in the descrip- tions can be converted to millimeters by multiplying by 0.02. The acronyms for the museums from which material was bor- rowed are listed in the acknowledgments section. Systasis Walker Systasis Walker, 1834: 288, 296 (Type spe- cies: Systasis encyrtoides Walker; West- wood 1840: 70 designation). Thomson, 1876: 193, 203. Ashmead, 1904: 274, 275. Schmiedeknecht, 1909: 275, 276, 280. Girault, 1915: 86. Nikol’skaya, 1952: 240. Boucéek, 1955: 324-327. Peck, 1963: 644. Peck et al., 1964: 33. Graham, 1969: 257- 263. Bouéek 1976: 11. Dzhanokmen, 1978: 102, 112. Boucek, 1988: 221, 310- 312, 386. Mani, 1989: 539, 540. Boucek and Rasplus, 1991: 106. Paruriella Girault, 1913a: 308; 1913b: 107 (Type species: Paruriella australiensis Gi- 571 rault; original designation). Bouéek, 1988: 310-311 (synonymy). Guieralia Risbec, 1951: 253 (Type species: Guieralia guierae Risbec; monotypic ge- nus). Boucéek, 1976: 11 (synonymy). Bouéek (1955) described the subgenus Systasina for the palearctic species S. an- nulipes (as S. clavicornis Bouéek) because this species has a short, clavate antennae and slender body in both sexes, and appar- ently a four-segmented club in the males. Systasina was more completely character- ized by Graham (1969), but because Sys- tasina contained only one species, it was difficult to differentiate subgeneric from species characters. The discovery of S. (Sys- tasina) tena, n. sp., and S. (Systasina) han- soni, n. sp., Shows that subgeneric charac- ters for Systasina are possibly limited to the weakly sculptured body, the elongate gaster and clavate flagellum in females, and the short scape in males. (The female body and antennal proportions for S. tena are un- known, however.) Other differences men- tioned by Bouéek (1955) and Graham (1969) are no longer valid. As in male S. (Systasis) spp., male S. tena and S. hansoni have the combined length of the flagellum and ped- icel as long as or longer than the head width, and at least in S. tena, the flagellum parallel- sided with the basal four funicular segments elongate. In male S. tena, the club has a small ventral patch of micropilosity and is as long as the distal three funicular seg- ments; in male S. hansoni, the club has an enlarged ventral patch of micropilosity sur- rounded by a well developed sulcus and so appears four-segmented. Male S. tena and both sexes of S. hansoni have the basal cell open posteriorly. Biology. The hosts of most Systasis spe- cies are probably either gall-formers or in- quilines, although some Systasis species may consume plant material as gall formers in their own right. Several of the Australian Systasis species have been reared — Systasis cecidomyiae from Didymomyia tiliacea (Bremi) (= Cecidomyia frauenfeldti) (Ceci- oie) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON domyiidae), S. flindersiae from galls on F] rsia (Rutaceae), S. graminis from ds of Panicum sp. (Poaceae), and S. lelex rom horn galls on iron-bark eucalyptus and galls on spotted gum (all records from Bou- éek 1988). The palearctic species Systasis angustula has been recovered from Kiefferia pimpinellae (F. Loew) (Cecidomyiidae) and Putoniella marsupialis (F. Loew) (Cecido- myiidae). Rearing records of S. encyrtoides from the Old World include Cecidomytiidae in the genera Contarinia Rondani, Dasy- neura Saunders, and Stenodiplosis Reuter (Anonymous 1965, Anonymous 1966, Bohm 1954, Graham 1969, Manniger 1940, Rubin 1965, Selivanova 1948); Agromyzi- dae in seeds (Graham 1969); two Tortrici- dae (Anonymous 1963 and Anonymous 1965, but see Graham 1969); and Apion compactum Desbrochers des Loges (Cur- culionidae) from broom (Hoffman 1958: 1561). Two of the Oriental species have also been reared from Cecidomyiidae. Systasis dasyneurae is recorded from Dasyneura lini Barnes (Ahmad 1939) and S. dalbergiae from two species on Dalbergia sissoo Rox- burgh (Fabaceae)—Contarinia dalbergiae Mani (Mani 1942, Bouéek et a/. 1978, Mani 1989, and possibly Gangwar and Prasad 1984) and Erosomyia indica Grover (Grov- er 1986 and Grover 1987). Unidentified In- dian Systasis species are also reported, one from the Noorda moringae Tams (Pyrali- dae) (Cherian and Basheer 1939), and four others from the Cecidomyiidae species As- phondylia sesami Felt (Mathur and Verma 1973), Contarinia sp. on Jasminum sambac Ait. (Oleaceae) (David, et al. 1990), Dasy- neura amaramanjarae on mango (Anacar- diaceae) (Grover 1986), and Phyllodiplosis sp. on Zizyphus jujuba Tamk. (Rhamna- ceae) (Tiwari 1975). The Ethiopian species, Systasis guierae, was reared from a gall on Guiera senegalensis Lam. (Combretaceae) (Risbec 1951). Guiera senegalensis is an Af- rican shrub on which walnut-sized galls, caused by Vuilletia howardi (Vuillet) (Phlaeothripidae), form on young shoots (Bradley 1969). These galls are home to a number of inquilines in the orders Thysa- noptera and Lepidoptera (Bradley 1969, Pitkin and Mound 1973), so the exact host needs clarification. The type series for the new nearctic species, S. aquila, was collect- ed in association with bud galls on Vaccin- ium corymbosum L. (Ericaceae), and Lyonia ligustrina (L.) DC. (Ericaceae). Systasis cen- chivora is phytophagous on the seeds of Cenchrus ciliaris Linnaeus and C. setigerus Vahl. (Poaceae) (Wadhi and Verma 1970 and Farooqgi and Menon 1972). As far as is known, those Systasis species on cecido- mylids are always external parasites on the larval stages of their hosts. Key To NEARCTIC SysT4SIS WALKER 1. Body sculpturing weak, vertex and metapleu- ron nearly smooth. Male with scape not quite reaching median ocellus in anterior view .... 2 — Body distinctly alveolate, vertex and meta- pleuron with distinct raised sculpture (Figs. 3, 5). Male with scape reaching at least to median ocellus Male with mesepisternum and tibiae metallic blue. Terminal segment of club with large patch of micropilosity (Fig. 10). Fore wing with about 15 or more admarginal setae .... hansoni, n. sp. — Male with mesepisternum and tibiae weakly metallic. Terminal segment of club with min- ute patch of micropilosity (Fig. 11). Fore wing with about 10 admarginal setae. (Female un- known) tena, Nn. sp. 3. Body color olive green to black. Scape pale and nonmetallic. Female with eye height 2.0 or more times malar distance; male eye height 2.2 or more times malar distance ........ aquila, n. sp. — Body color green or blue. Scape dark brown or black, with distinct metallic reflections or wholly metallic. Female with eye height 2.0 or less times malar distance; male with eye height 2.2 or less times malar distance 4. Female body length 1.4-1.7 mm. Antenna with scape reaching the median ocellus; Fl half length of pedicel; club length longer than com- bined length of F3-5 (Figs. 4, 8). Male antenna with scape reaching vertex; combined length of pedicel and flagellum 1.0-1.1 x head width; funicular setae sparse, setal length equal to that of a funicular segment ........ parvula Thomson — Female body length 1.8-—2.5 mm. Antenna with scape extending to or above vertex; Fl about to VOLUME 97, NUMBER 3 as long as pedicel; club length shorter than combined length of F3—S (Fig. 9). Male antenna with scape extending above vertex; combined length of pedicel and flagellum 1.2—1.4= head length; funicular setae fairly dense, setal length half that of a funicular segment 5a age ee it ae Ree Net encyrtoides Walker Systasis (Systasis) aquila Heydon, New SPECIES Figs. 5-6 Diagnosis.— Systasis aquila may be rec- ognized by the following combination of characters: 1.* Body color greenish black or bluish black. 2.* Scape pale, nonmetallic. 3. Body sculpture coarse. 4. Female body length 1.7-—2.5 mm; male body length 1.3- 2.0 mm. 5. Female eye height 2.0—2.2 x ma- lar distance; male eye height 2.2—3.0 x ma- lar distance. 6. Female scape reaching me- dian ocellus, length ~%4 eye height (range 0.72-0.82); male scape reaching just above median ocellus, length ~% eye height (range 0.64-0.72). 7. Male with combined length of pedicel and flagellum 1.2 head width. 8. Female antenna with F1 about as long as pedicel, flagellum parallel-sided, width of F5 1.0-1.1< width of Fl (Fig. 6). 9. Male funicular setae length '2—*4 the length of the funicular segments. 10. Plicae cariniform, extending 7 of way to anterior margin of propodeum. 11. Female gaster length 1.4—- 1.8x width, 0.86-0.95x the combined length of head and mesosoma (Fig. 5). Description.— Holotype, female. Color: Body greenish black with yellow reflections on face, pleural region, T5—7; dorsellum, propodeum, T1 dark green. Antenna with scape yellow brown; flagellum black. Legs with coxa, trochanter, femur greenish black; fore tibia and tarsi yellow-brown, tibia slightly darker over mesal '; middle, hind tibiae greenish black (yellow-brown at tips); middle and hind tarsi brownish yellow with * Asterisk indicates unique character state among nearctic species. 573 pretarsi brown. Fore wing veins translucent brown. Sculpture: Head, mesosoma minutely al- veolate (Fig. 5); anterior aspect of head (ex- cept for strip from anterior margin of clyp- eus to median ocellus) with distinct pilifer- ous punctures; sculpture of mesopleural re- gion relatively low; middle lobe of mesoscutum and scutellum with scattered piliferous punctures; gaster alveolate, sculp- ture only slightly raised above body surface. Structure: Body length 2.5 mm. Head pentagonal in anterior view, width 1.2 x height (42:36), 2.1 length (42:20); eye height 1.3 x length (20:15), 2.2 x malar dis- tance (20:9), length 5.0 x temple length (15: 3); torulus 1.5 x own diameter above LOcL; ratio of MOD, OOL, POL, LOL as 3.5:5.0: 13.5:6.0. Antenna (Fig. 6) with length of pedicel plus flagellum 1.0 x head width (43: 42); scape strongly flattened, length 3.3 x greatest width (15.0:4.5); relative lengths of scape, pedicel, anelli, F1—5, club as 15.0: 329212575:075.0:4:5:5:0:5.0:12 5s widthssot F1, F5, clubas 4.0:4.0:4.5; 3-5 MPP sensilla in single irregular row visible on each seg- ment. Mesosoma length 1.3 width (54: 41); dorsellum just shorter mesally than lat- erally; propodeum with median carina, pos- terior '2 of plicae, petiolar foramen strongly carinate; callus with six erect mesal and one reclinate lateral setae; spiracle ovate, par- tially beneath the metanotum. Fore wing length 2.2 x width (110:50); relative lengths of submarginal, marginal, postmarginal, stigmal veins as 38.0:25.0:12.5:10.0; costal cell with single dorsal seta, with complete and partial ventral rows; basal cell bare; bas- al vein with row of four setae; admarginal setae in more than one row. Gaster chor- date, its length 1.4 width (60:42): hind margin of T1 straight; hypopygium extend- ing 0.37 length of gaster (22:60); lateral membranous areas on T1 large, extending posteriorly beyond basal fovea. Allotype, male: Similar to holotype ex- cept generally paler—head, pleural region, gaster dark green; dorsum of mesosoma dark 574 blue; and flagellum brownish black. Body length 2.0 mm. Antenna with combined length of pedicel and flagellum 1.2 head vidth (47.5:38.0); relative lengths of scape, pedicel, anelli, Fl-5, club as 13.0:5.0:1.0: 73525.5752525-525.0212-5> widthsoieF ik E>. club as 4:4:4; scape flattened, length 2.6 x maximum width (13:5); 2-4 MPP sensilla in single irregular row visible per segment; setae dense, semierect, length 2 funicular segment length. Gaster ovate, length 1.2 x width (38:33). Variation. — Female body length 1.7 to 2.5 mm; male body length 1.3 to 2.0 mm. The color of the dorsum of the male mesosoma is sometimes greenish black. The costal cell may have no dorsal setae and only one ven- tral setal row. In about half the specimens, there is only a single row of admarginal se- tae. Etymology.—The species name, derived from the Latin word aquilus, meaning dark colored, refers to the dark body color of this species relative to others in the genus. Type material.—The holotype, allotype, and 8 female and 3 male paratypes were collected associated with bud galls on Vac- cinium corymbosum L. from IV-V.1983 at Chatsworth, New Jersey (United States), by H. P. Boyd (USNM). Four other paratypes were collected as follows (CNCI, UCDC, USNM): United States. West Virginia: Cooper’s Rock State Forest, 2.5 miles w. Bruceton Mills, 5.VI.1964, 1 6, 6.V1I.1964, 1 6. Unknown locality: 1 2; on Lyonia li- gustrina (L.) DC. bud gall, 1 °. Biology.—The host(s) of Systasis aquila are unknown but this species has been reared in association with bud galls on Vaccinium corymbosum L. (Ericaceae), and Lyonia li- gustrina (L.) DC. (Ericaceae). Systasis (Systasis) encyrtoides Walker Fig. 9 Systasis encyrtoides Walker, 1834: 296. Haliday, 1841-1842: v, pl. B, fig. 1. Gra- ham: 1969: 259. 261. Boucek,.1977: 56. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hormocerus impletus Walker, 1872: 96. Graham, 1969: 261 (synonymy). Systasis longicornis Thomson, 1876: 204. Graham, 1969: 261 (synonymy). The synonymy, proposed by Graham (1969), of Hormocerus impletus Walker and Systasis longicornis Thomson with S. en- cyrtoides Walker are accepted without con- firmation by the author. Tridymus puncta- tus Ratzeburg may also be a synonym of this species (Reinhard 1857 and Graham 1969), but the types are lost. Diagnosis: Systasis encyrtoides can be recognized by the following combination of characters: 1. Body color blue or green. 2. Scape dark, weakly metallic. 3. Body sculp- ture coarse. 4. Female body length 1.8-2.5 mm; male body length 1.0-1.6 mm. 5. Fe- male eye height 1.6-2.0 malar distance; male eye height 2.0-2.2 x malar distance. 6.* Female scape extending above vertex, length ~% eye height (range 0.73-0.82); male scape extending above vertex, length about 0.7 x eye height. 7.* Males with com- bined length of pedicel and flagellum 1.2- 1.4x head width. 8. Female antenna with the length of F1 slightly shorter or equal to that of the pedicel, flagellum parallel sided (Fig. 9). 9. Male funicular setae length '2 length of funicular segments. 10. Plicae ridgelike, with a smooth crest extending ~73 of the way to the anterior margin of the propodeum. 11. Female gaster length 1.1- 1.7x width, 0.80-0.96x the combined length of the head and mesosoma. Material examined.—I examined 14 fe- males and | male from the Nearctic Region collected as follows (CNCI, UCDC, USNM): Canada. ALBERTA: Elkwater Lake, 19.VI1I.1956, 1 9. BRITISH COLUMBIA: 15 miles e. Hope, 12.VII.1973, 1 2; Robe- son, 12.VI-1950) 1-2, 21-253VEI94 7. lee 22.V1.1947, 1 2; Squamish, Diamond Head Trail. 7.VAML1953.0 1 2.5283 Vil 1953 alee: MANITOBA: Churchill, 13.VIII.1952, 1 2. NORTHWEST TERRITORIES: Reindeer Depot, Mackinzie Delta, 7.VII.1948, 1 9; VOLUME 97, NUMBER 3 Figs. 7-11. toides, female, antenna. 10, Systasis hansoni, male, antenna. 11. Systasis tena, male, antenna. Yellowknife, 5.VI.1953, 1 2. United States. ALASKA: McKinley Hotel, 5.VII.1954, 1 2. COLORADO: Chicago Cr., Clear Creek Co., 2.VIII.1961, 1 °, along Fish Creek, 6 kms. bake George, 7.VITE1992. 1 9. MARYLAND: Adelphi, 21.V.1988, 1 2. OREGON: Saddleback Mountain, Lincoln Co., 3.1X.1960, 1 2. Systasis encyrtoides is common throughout Europe (Bouéek 1977). Biology.—The host(s) of Systasis encyr- toides in the Nearctic Region are unknown, but this species has been reared from galls of cecidomyiids or agromyzids in seeds in Europe. Old World hosts include the agro- Sie rn ee a ee ee 279 -- v2 ie) Be -- a A cod 7,Systasis hansoni, female, whole body. 8, Systasis parvula, female, antenna. 9. Systasis encyr- myzid Phytomyza isais Hering in seeds of Odontites verna (Bell.) Dum. (Scrophulari- aceae) (Graham 1969), and the cecido- myiids Cecidomyia crataegi Winn. (Ghigi 1901), Dasyneura affinis Kieffer (B6hm 1954), D. epilobii (F. Loew) on Chama- enerion angustifolium L. (Graham, 1969); Contarinia citri (Rubin 1965), C. ilicis Kief- fer (Anonymous 1965), C. medicaginis Kief- fer (Anonymous 1961, Anonymous 1966, Bonnemaison 1968, Ferron 1964, Kralovic 1964, and Stavrakis and Lambrakopoulos 1971), C. pulchripes (Kieffer) on Sarotham- nus scoparius (L.) Wimmer (Parnell 1963); 576 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Stenodiplosis panici (Selivanova 1948); and pods of broom (Walker 1848 Ratzeburg 1852, Reinhard 1857, Graham 1969). There are palearctic records from the tortricids Conchylidea implicatans Wek. and Tortrix viridana L. (Anonymous 1963 and Anon- ymous 1965), but Graham (1969) suspects these reports may be in error. Other records include two species of weevils of the genus Apion that occur in pods of broom in France (Hoffman 1958), but they do not seem to parasitize this host in England (Parnell 1963). Graham (1969) suspects there may be more than one generation per year in Europe. Further details on the biology of S. encyrtoides are found in Ghigi (1901), Par- nell (1963), Ferron (1964), and Tudor et al. (1978). Systasis (Systasis) hansoni Heydon, NEw SPECIES Figs. 7, 10 Diagnosis. — Systasis hansoni may be rec- ognized by the following combination of characters: |. Body color metallic blue or blue-green. 2. Scape blue, metallic. 3. Body sculpture weak. 4. Female body length 2.3- 2.6 mm, known male body length 1.8 mm. 5. Female eye height 2.8-3.0 x malar dis- tance; male eye height 2.9 x malar distance. 6. Female scape not extending to median ocellus, length between '2 and *% eye height (range 0.57-64). 7. Male with combined length of pedicel and flagellum 1.0 head width. 8. Female antenna with Fl much shorter than pedicel, flagellum clavate, width of F5 1.8x width of Fl. 9. Male funicular setae length about equal to length of the funicular segments. 10. Plicae weakly de- veloped as low broad folds in propodeum, smooth crest lacking. 11. * Female gaster length 2.1-3.6x width, * 1.4-1.6x com- bined length of the head and mesosoma (Fig. D): Description.— Holotype, female. Color: Body blue except dorsum of mesosoma, gaster with yellowish green reflections, pro- notum just anterior to spiracle yellowish green. Antenna with scape, pedicel blue; re- mainder brownish black. Legs blue except fore and middle knees and tibial tips brown- ish yellow, hind knees and tarsi brown. Wing veins brown. Sculpture: Clypeus finely granulate; re- mainder of head weakly alveolate but an- terior aspect of head with weak piliferous punctures; mesoscutum, scutellum alveo- late; frenum, dorsellum, propodeum weakly alveolate; gastral terga weakly alveolate. Structure: Body length 2.5 mm. Head nearly circular in anterior view, width 1.2 x height (26.0:22.5), 1.9 length (26:14); eye height 1.3 x length (14:11), 2.8 x malar dis- tance (14:5), length 3.7 x temple length (11: 3); torulus | x own diameter above LOcL; ratio of MOD, OOL, POL, LOL as 3.0:3.0: 7.0:3.5. Antenna with length of pedicel plus flagellum 0.83 head width (21.5:26.0); scape flattened, length 2.7 its greatest width (8:3); relative lengths of scape, ped- icel, anelli, F1—5, club as 8.0:4.0:1.0:1.0:1.5: 1.5-2.0:2.0:7.5; widths of Fl, F5, club as 2.0:3.5:4.5; 1-2 MPP sensilla in a single row visible per segment. Mesosoma length 1.6 x width (41:26); dorsellum shorter mesally than laterally; propodeum with median ca- rina, plicae, petiolar foramen carinate; cal- lus with | lateral reclinate seta, 6 mesal erect setae; spiracle round, on the anterior margin of the pronotum. Fore wing length 2.5 x width (86:35); relative lengths of submar- ginal, marginal, postmarginal, stigmal veins as 29.0:19.0:13.0:7.5; costal cell with 5 dor- sal setae, ventrally with | complete row and partial second row in apical '; basal cell bare, with | or 2 setae on cubital vein; basal vein with narrow band of setae down its length; admarginal setae in single row of 16 setae on left fore wing, 15 setae on right. Gaster lanceolate (Fig. 7); length 2.6 x width (75:29), 1.5 x combined length of head and mesosoma (75.0:51.5); hind margin of T1 straight; hypopygium extending 0.29 x gas- tral length (22:75); lateral membranous ar- eas on T1 short. Allotype, male: Similar to holotype ex- cept posterior 2 of T1, T2—4 dull dark pur- ple. Body length 1.8 mm. Antenna (Fig. 10) VOLUME 97, NUMBER 3 with combined length of pedicel plus fla- gellum 1.0 head width (25:25): relative lengths of scape, pedicel, anelli, F1—5, club as1O1025292h.0:2102275:2:5:2.5:2.5:8:5: widths of Fl, F5, club as 2:2:4; 1-2 MPP sensilla in single row visible per segment; setae rec- linate, length about equal to length of a fu- nicular segment. Gaster ovate, length 3.1 x width (40:13). Variation.— The body length of females varies from 2.3—2.5 mm. All the specimens in the type series were critical point dried so the body lengths and relative lengths of the gaster reported herein may be somewhat exaggerated. The two females with a gaster length 1.6 x the combined length of the head and mesosoma have the gaster appearing noticeably expanded. The females with a gaster length 1.3-1.4 x the combined length of the head and mesosoma have a very nat- ural appearance, and living or air-dried specimens probably have their gaster of about these proportions. The color patterns of the females are generally similar to the holotype except one is noticeably more vi- olet and two are more green. The setae of the dorsal surface of the basal cell are re- stricted to a single row down the basal vein except in the holotype and allotype where there is a partial second row. Etymology.—This species is named in honor of Paul Hanson through whose col- lecting efforts the Oregon chalcidoid fauna is becoming better known. Type material.—The holotype (OSUO), allotype (OSUO), and 3 female paratypes were collected on 24 June 1983, on the bluff at Pike Creek, Oregon (United States), by Paul Hanson. Three additional paratypes were collected | July 1986, 2 milese. Sisters, Oregon. Biology. — Nothing is known of the host(s) of S. hansoni. Systasis (Systasis) parvula Thomson Figs. 1-4, 8 Systasis parvula Thomson 1876: 205. Gra- ham 1969: 259, 260, 263. ST Diagnosis: Systasis parvula can be rec- ognized by the following combination of characters: 1. The body color blue or green. 2. The scape metallic green or blue. 3. Body sculpture coarse (Fig. 3). 4. Female body length 1.4-1.7 mm; male body length 1.1- 1.5 mm. 5. Female eye height 1.8—1.9 x ma- lar distance; male eye height 2.0x malar distance. 6. Female scape reaching median ocellus, length ~7 eye height; male scape reaching to vertex, length ~*4 eye height. 7. Males with combined length of pedicel and flagellum 1.1 >. : . 8¢ PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Scanning electron micrographs of P. sericans eggs. (A) lateral view of anterior end, showing compact granular ridges; (B) lateral view of middle portion, showing flat connected ridges; (C) lateral view of posterior end, showing strengthened and compact ridges; (D) anterolateral view, magnification of aligned minute granules on front end; (E) lateral view, showing toe-like granules of circular structure on the middle portion; (F) pos- terolateral view, showing tooth-like granules on a circular structure. Magnifications are based on scale bar in A: A = 0.1 mm, B = 0.1 mm, C = 0.1 mm, D = 25 um, E = 23 wm, F = 15 um. portion of the egg, and are parallel and con- nected by cross-ridges (Fig. 3B). High mag- nification shows that the granules of the flat ridges are toe-like (Fig. 3E). At the anterior end they become higher and compact (Figs. 3A, D). At the posterior end they are high- est, and form a circular structure (7-30 wm in diameter; Fig. 3C). The granules on the thin ridges of the posterior end are tooth- like (Fig. 3F). The surface of the exochorion between the thin ridges 1s covered by scat- tered enlarged granules (Figs. 3A-C) and compact aeropyles (Figs. 3A-F). The en- larged granules are 10-15 um in diameter and the aeropyles are |.5—2 um in diameter. P. hepaticus Erichson.—Size: length, 0.79- 0.88 (0.83 + 0.03) mm; width, 0.39-0.50 (0.44 + 0.03) mm. The egg is extended an- teriorly into a small projection (Figs. 1E, 4A) and lacks a process toward the posterior end (Figs. 1E, 4B). The exochorion is cov- ered by compact minute granules and scat- tered irregular enlarged granules. These en- larged granules are alternated with pits, forming cone-shaped structures resembling volcanoes (Figs. 1E, 4A, B) which are evenly distributed over the entire surface (Fig. 1 E). The minute granules on the enlarged ones are short and scattered (Fig. 4C), but tooth- like and aligned into lines between the cone- shaped structures (Fig. 4D). The granules form no precise patterns. VOLUME 97, NUMBER 3 1 tN +>” fe Fig. 4. Scanning electron micrographs of P. hepaticus eggs. (A) anterolateral view; (b) posterolateral view, showing volcano-like structure; (C) lateral view, showing minute granules on the enlarged ones; (D) lateral view, showing minute granules on the egg surface between the enlarged ones. Magnifications are based on scale bar in A: A = 0.10 mm, B = 0.10 mm, C = 20 um, D = 20 pm. KEY TO THE EGGs OF DUNG-INHABITING PHILONTHUS IN NORTHCENTRAL FLORIDA Exochorion with strong, longitudinal ridges (E185: GWAC) ie ast Seti hs e E SR VY 2 Exochorion without strong longitudinal ridges (E1gs:4lIDSE)) cco; SES Ee ae Ao PLL 4 With 4—5 ridges; exochorion between ridges covered with aeropyles and scattered tubercles (Rigs) aeons ventralis (Gravenhorst) With 7-8 ridges; exochorion between ridges covered by tubercles or granular teeth ....... 3 . All ridges well-developed (continuous); tuber- cles present on the exochorion between the midgesi(hign ZA), see. longicornis Stephens Not all ridges well developed (discontinuous); granular teeth present on the exochorion be- tween the ridges (Fig. 2B) flavolimbatus Erichson . Exochorion with weak and thin granular ridges 588 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and enlarged granules over the entire surface (Figs, 3A—©@)3 22 eee sericans (Gravenhorst) Exochorion with compact minute granules and cattered enlarged granules which are alternat- ed with pits, forming cone-shaped structures over the entire surface (Figs. 1E, 4A, B) .... ee here ownh oe hepaticus Erichson Egg size.— Mean egg length was found to vary significantly between species (F = 167.08; df = 4,45; P < 0.01). Duncan’s mul- tiple range test determined that the eggs of P. longicornis and P. ventralis were statis- tically similar to each other yet significantly larger (longer) than those of P. flavolimba- tus, P. sericans and P. hepaticus. Egg of P. flavolimbatus, P. sericans and P. hepaticus did not differ significantly in size. DISCUSSION Alcohol fixation and desiccation were tried using staphylinid eggs of several spe- cies from three subfamilies. Most eggs of Philonthus and those of Neohypnus pusillus (Sachse) (Staphylininae) did not collapse, but almost all the eggs of Aleochara notula and Atheta sp. (Aleocharinae) and Platystethus americanus Erichson and Anotylus insig- nitus (Gravenhorst) (Oxytelinae) collapsed. This may be because the egg chorion of Phi- lonthus spp. and N. pusillus is thicker than that of the other species and is sculptured over the entire outer surface. The sculpture forms supportive structure for the egg shells and may help the eggs resist external pres- sure and substances from decaying dung. The egg chorion of A. notula, Atheta sp. and P. americanus, however, is thin and smooth, without any surface sculpture. Egg size is not a reliable character for sep- arating eggs of these five species of Philon- thus, because the size is similar for P. /on- gicornis and P. ventralis, and for P. flavo- limbatus, P. sericans and P. hepaticus. Mi- crosculpture is the only egg character that can be used for species determination of Philonthus studied here. These include pat- terns and presence of ridges, tubercles, pro- cesses and projections, granular teeth, gran- ules, cone-shaped structures and aeropyles. The egg of P. cruentatus Gmelin was de- scribed by Hunter et al. (1989). Its shape and ridge characteristics are similar to those of P. longicornis. Further detailed study is needed for separating these two species be- cause they have overlapping distribution (Blume 1985). Tiny holes evenly cover the egg surface of P. ventralis and P. sericans, and open on the tubercles of the egg ridges of P. /ongi- cornis and P. flavolimbatus. These holes were named aeropyles by Hinton (1970, 1981a, b), because they are used for absorbing ox- ygen from the ambient air. Aeropyles are common on the eggs of terrestrial insects (Hinton 1970). Philonthus eggs were most abundant in field-collected dung of about 7 d old. ACKNOWLEDGMENTS We thank W. Carpenter and H. L. Crom- roy (Department of Entomology and Nem- atology, University of Florida) for their technical advice and assistance with SEM, and G. T. Fincher (USDA-ARS, College Station, Texas) for the gift of one of his emergence boxes for extracting dung-inhab- iting insects. We also thank M. C. Thomas (Division of Plant Industry, Gainesville, Florida) and J. P. Parkman (Department of Entomology and Nematology, University of Florida) for careful review of this manu- script. This is Florida Agricultural Experi- ment Station, Journal Series Number R-03853. LITERATURE CITED Blume, R.R. 1985. Achecklist, distributional record, and annotated bibliography of the insects associ- ated with bovine droppings on pastures in Amer- ica north of Mexico. Southwestern Entomologist 9: 1-55. Endris, R. G., D. G. Young, and P. V. Perkins. 1987. Ultrastructural comparison of egg surface mor- phology of five Lutzomyia species (Diptera: Psy- chodidae). Journal of Medical Entomology 24: 412- 415. Fincher, G. T. and J. W. Summerlin. 1994. Predation on the horn fly by three exotic species of Philon- thus. Journal of Agricultural Entomology 11: 45- 48. VOLUME 97, NUMBER 3 Frank, J.H. 1968. Notes on the biology of Philonthus decorus (Grav.) (Col., Staphylinidae). Entomolo- gist’s Monthly Magazine 103: 273-277. 1986. A preliminary checklist of the Staph- ylinidae (Coleoptera) of Florida. Florida Ento- mologist 363-382. Harris, R. L. and L. M. Oliver. 1979. Predation of Philonthus flavolimbatus on the horn fly. Envi- ronmental Entomology 8: 259-260. Harris, R. L. and R. R. Blume. 1986. Beneficial in- sects inhabiting bovine droppings in the United Stages, pp. 10-15. Jn R.S. Patterson & D. A. Rutz, eds., Biological Control of Muscoid Flies. Ento- mological Society of America Miscellaneous Pub- lications No. 61. Hinton, H.E. 1970. Insect eggshells. Scientific Amer- ican 223: 84-91. 198la. Biology of Insect Eggs, Vol 1. Per- gamon Press, Oxford. 1981b. Biology of Insect Eggs. Vol 2. Per- gamon Press, Oxford. Hu, G. Y. and J. H. Frank. 1995. New distributional records for Platystethus (Coleoptera: Staphylini- dae: Oxytelinae) with notes on the biology of P. americanus. Florida Entomologist 78: 137-144. Hunter, J. S. III, G. T. Fincher, and D. E. Bay. 1986. Methods for rearing Philonthus spp. associated with cattle droppings. Journal of Entomological Science 21: 83-86. Hunter, J. S. IJ, D. E. Bay, and G. T. Fincher. 1989. Laboratory and field observations on the life his- tory and habits of Philonthus cruentatus and Phi- lonthus flavolimbatus. Southwestern Entomologist 14: 41-47. Laurence, B. R. 1956. The larval inhabitants of cow pats. Journal of Animal Ecology 23: 234-260. Macqueen, A. and B. P. Beirne. 1975. Influence of other insects on production of horn fly, Haema- tobia irritans (Diptera: Muscidae), from cattle dung in South Central British Columbia. Canadian En- tomologist 107: 1255-1264. Mank, H.G. 1923. The biology of the Staphylinidae. 589 Annals of the Entomological Society of America 16: 220-37. Roth, J. P. 1982. Predation on the horn fly, Hae- matobia irritans (L.). Southwestern Entomologist 7: 26-30. SAS Institute. 1990. User’s Guide: Statistics. Ver. 6, 4th ed. SAS Institute Inc., Cary, North Carolina. Sanders, D. P. and R. C. Dobson. 1966. The insect complex associated with bovine manure in Indi- ana. Annals of the Entomological Society of Amer- ica 59: 955-59. Tawfik, M. F. S., K. T. Awadallah, E. D. Ammar, and S. M. Abul-Ela. 1976a. The life history of Phi- lonthus misor Tott. (Coleoptera: Staphylinidae). Bulletin de la Societe Entomologique d’Egypte 60: 345-355. 1976b. The life history of the staphylinid Philonthus turbidus Er. (Coleoptera: Staphylini- dae). Bulletin de la Societe d’Entomologique Egypte 60: 357-366. 1976c. On the bionomics of Philonthus lon- gicornis Steph. (Coleoptera: Staphylinidae). Bul- letin de la Societe Entomologique Egypte 60: 379- BSi- Thomas, G. D. and C. E. Morgan. 1972. Field-mor- tality studies of the immature stages of the horn fly in Missouri. Environmental Entomology 1: 453- 459. Valiela, I. 1969. An experimental study of the mor- tality factors of larval Musca autumnalis DeGeer. Ecological Monographs 39: 119-225. Wingo, C. W., G. D. Thomas, G. N. Clark and C. E. Morgan. 1974. Succession and abundance of in- sects in pasture manure: relationship to face fly survival. Annals of the Entomological Society of America 67: 386-390. Woodruff, R. E. 1973. The scarab beetles of Florida. Pt. 1. Arthropods of Florida and Neighboring Land Areas. Vol. 8. Florida Department of Agriculture and Consumer Services, Division of Plant Indus- try, Gainesville. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 590-595 NEW AMISEGINE WASPS FROM SOUTHEAST ASIA (HYMENOPTERA: CHRYSIDIDAE) Lynn S. KIMSEY Department of Entomology, University of California, Davis, 95616. Abstract. —Six new species of Amiseginae (Chrysididae) are described from southeast Asia, including Atoposega decorata (New Caledonia), Bupon thailandicus (Thailand), Bu- pon bicornutus (Vietnam), Magdalium orchidense (Taiwan); Magdalium lucidum (Malaya), and Perissosega flavipes (Thailand). Key Words: Additional collecting with flight inter- cept, malaise, and pan traps in southeast Asia has demonstrated how little we know about the fauna of amisegine chrysidids in this region. Recently studied material that was collected in this fashion, and deposited in the Canadian National Insect Collection and Bishop Museum, clearly indicates how our knowledge of these genera is still in a primitive state. Generic distributions are clearly more widespread than recorded in Kimsey and Bohart (1991). Bupon was previously known from a single Malaysian species. The new species described below are from Thailand and Vietnam. Magdalium is another genus known from one Malaysian species, and has now been collected from Orchid Island, Tai- wan, along with a second species from Ma- laya. Baeosega, known from Sri Lanka, is recorded below from southern Japan. Per- issosega, previously described from Sr Lan- ka, 1s recorded below from Thailand. Fi- nally, an additional, spectacularly blue-col- ored species of Atoposega is described below from New Caledonia. The other three spe- Amiseginae, Chrysididae, Atoposega, Bupon, Magdalium, Perissosega cies in this genus are from Malaysia and the Philippines. Specimens described below were ob- tained from and/or deposited in the follow- ing collections. Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, J. M. Carpenter, S. R. Shaw, (CAMBRIDGE); Canadian National Insect Collection, Ottawa, Ontario, L. Masner (OTTAWA); Bohart Museum of Entomol- ogy, University of California, Davis, S. L. Heydon (DAVIS); Bernice P. Bishop Mu- seum, Honolulu, Hawaii, G. Nishida (HO- NOLULWU), and U. S. National Museum, Washington, D.C. (K. V. Krombein). Re- positories given below are indicated by the city name of the collection in capital letters enclosed in parentheses as above. Atoposega decorata Kimsey, NEW SPECIES Fig. 1 Description. — Female (holotype): Body length 5 mm. Face (Fig. 1): scapal basin clearly delimited and evenly cross-ridged, malar space 2.8 midocellus diameters long; head slightly wider than long; midocellus VOLUME 97, NUMBER 3 591 2. Perissosega flavipes ‘le if Magdalium lucidium 6. Magdalium % orchidense thailandicus Figs. 1-7. 1-3, 5-7, Front view of face. 4, Dorsal view of male head. 592 } midocellus diameters from ocular mar- >in: subantennal distance 2.8 midocellus di- armeters long; flagellomere I length 2.9x breadth; flagellomere II 0.7 as long as broad; pronotum slightly shorter than me- dian length of scutum; vertex and pronotum with punctures sunken between irregular, polished, longitudinal welts, pronotum and scutum with punctures sunken between coarse longitudinal ridges; pronotal pos- terolateral lobe polished and nearly im- punctate; mesopleuron with large nearly contiguous punctures, scrobe not apparent; scutellum with dense, deep, contiguous punctures; metanotal length 0.8 that of scutellum; forewing densely setose except for medial band traversing wing from stig- ma to posterior margin, wing darkly and stained and with dark setae, except pale me- dially and around apex extending in along stained remnant of Rs, including setae at base; hindfemur with minute punctures; propodeal angle digitate and pale apically; terga highly polished with few small widely scattered punctures, without striae or “scratches” laterally. Head and thorax gen- erally metallic blue, scutum, metanotum and propodeum more purple, scutellum greener; legs metallic blue on coxae, femora medially and mid- and hindtibiae; forefemur entirely blue except apex pale yellow, foretibia pale yellow with blue tints; tarsi brown; remain- der pale yellow to cream-colored; tergum I brown with blue tints, posterolateral cor- ners pale yellow to cream-colored; remain- der of abdomen brown with blue tints. Male. Unknown. Type material.— Holotype, female, NEW CALEDONIA: Ciu, near Mt. Canala, 300 m, E. O. Wilson (CAMBRIDGE). Etymology.— The name refers to the dec- orative coloration of this species. Discussion.—This is the most brilliantly colored species of Atoposega. The bright blues and purples will readily distinguish decorata its congeners, simulans Kimsey, rieki (Krombein) and /ineata (Krombein). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON It has the smooth forefemur, terga without fine “scratches”, patterned wings and large size of simulans and lineata, and metallic coloration more typical of rieki. Bupon bicornutus Kimsey, NEw SPECIES Figs. 3, 4 Description.— Male (holotype): Body length 4 mm. Face (Fig. 3); scapal basin highly polished, and deeply concave below strongly projecting bilobate brow; brow produced into two large flattened, and dor- sally concave projections; vertex convex; head and thorax covered with coarse con- tiguous punctures; eye encircled by obso- lescent carina, clypeal apex broadly round- ed; subantennal distance 1.5 midocellus di- ameters long; malar space 3 midocellus di- ameters, with vertical sulcus; hindocellus 1 diameter from ocular margin; midocellus 2.2 midocellus diameters from ocular mar- gin; flagellomere I 2.6 as long as broad and arched; flagellomere II 1.3 x as long as broad; occipital carina complete; pronotum as long as scutum, with obscure oblong de- pression posteromedially and small pit on lateral lobe; mesopleuron without scrobal sulcus, propodeal posterior surface coarsely and densely rugose, lateral angle short and blunt; terga relatively sharp-edged laterally, with dense, small punctures, 0.5—1 puncture diameter apart or less. Body black; with blu- ish highlights on terga; antennae dark brown; legs dark brown with pale joints, and fore- and midtarsi pale; wings brown tinted. Female unknown. Paratypes differ in length from the type varying from 4-5 mm. Type material.—Holotype, male, VIET- NAM: Fyan, 900-1000 m, 11 July—9 Aug. 1961, N. R. Spencer (HONOLULU). Para- types: | male, Mt. Lang Bian, 1500-2000 m, 19 May-8 June 1961, N. R. Spencer (HONOLULU), | male, 17 km s Dilinh, 1300 m, 6-13 Oct. 1960, C. M. Yoshimoto (DAVIS). VOLUME 97, NUMBER 3 Etymology.—The species name refers to the strongly projecting bilobate frontal ca- rina. Discussion.—The most distinctive fea- tures of this species are the exaggerated frontal carina, which is produced and mod- ified into two excavated, subtruncate lobes, and the strongly concave and smooth scapal basin. Otherwise bicornutus resembles thai- landicus in most respects, with the same general coloration and density of puncta- tion. Bupon thailandicus Kimsey, NEw SPECIES Fig. 5 Description.— Male (holotype): Body length 4 mm. Face (Fig. 5); scapal basin deeply concave and sunken below strongly projecting carinate brow, with coarse cross- ridges; vertex convex; head and thorax cov- ered with coarse contiguous punctures; eye encircled by obsolescent carina, clypeal apex broadly rounded; subantennal distance 1.2 midocellus diameters long; malar space 2.5 midocellus diameters long, with vertical sulcus; hindocellus 0.8 diameter from oc- ular margin; midocellus 2.5 midocellus di- ameters from ocular margin; flagellomere I 3x as long as broad and arched; flagello- mere II 1.3x as long as broad; occipital carina complete; pronotum 0.5 x combined lengths of scutum, scutellum and metano- tum, with oblong pit posteromedially and small pit on lateral lobe; mesopleuron with- out scrobal sulcus, propodeal posterior sur- face coarsely and densely rugose, lateral an- gle short and blunt; terga relatively sharp- edged laterally, with dense, small punctures, 0.5 puncture diameter apart or less. Body black with bluish tints on abdominal dor- sum; antennae dark brown; legs dark brown with pale joints, and fore- and midtarsi pale; wings brown tinted. Female unknown. Paratypes differ somewhat from the type in size, ranging from 3—4 mm. Type material.— Holotype, male, THAI- 593 LAND: 70 km sw Chiang Mai, 800 m, Doi Inthanon National Park, Vachiratharu Falls, Cerocarpus forest, April-May 1990, B. V. Brown (OTTAWA). Paratypes: 8 males, same data as type (HONOLULU, DAVIS, WASHINGTON); one male: THAILAND: Korat, Pakchong, Musi, Vil. 9, Khlong Yai, 25 May 1969, G. R. Balmer (HONOLU- LU). Etymology.— This species named after its country of collection, Thailand. Discussion.—The two other species de- scribed in this genus are pashoanus Kimsey and bicornutus. Bupon thailandicus can be distinguished from pashoanus by the dark legs (yellow in pashoanus), convex vertex, and arched flagellomere I, and from bicor- nutus by the simple frontal carina. Magdalium lucidum Kimsey, NEw SPeEcIES Fig. 7 Description.— Female (holotype): Body length 4 mm. Face (Fig. 7); scapal basin highly polished and impunctate; malar space 3 midocellus diameters long, with vertical sulcus; subantennal distance slightly longer than 1 midocellus diameter; flagellomere I 1.6x as long as broad; flagellomeres II-III 0.9 x as long as broad; occipital carina pres- ent dorsally; midocellus 2 diameters from ocular margin; vertex and frons with small, widely separated punctures, 2-5 puncture diameters apart; hindocellus 0.8 diameter from ocular margin; pronotum only slightly longer than scutum, with thin sulcus ex- tending anterioriorly along medial line and pit on lateral lobe; mesopleuron with sub- alar fossa, short oblique mesopleural carina and scrobal sulcus, scrobal sulcus parallel- sided and about 6 x as wide as long; thoracic punctures 0.5—1.0 puncture diameter apart dorsally and 1-2 puncture diameters later- ally; notauli deep and straight; metanotum as long as scutellum; propodeum with short blunt posterolateral corner; tergal punctures 0.5—1.0 puncture diameter apart, except for impunctate medial line. Head, thorax and 594 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON odomen shiny black, except pale oblique medial line traversing terga; antenna yellow, xcept dorsum of apical nine flagellomeres dark brown; legs entirely yellow, except hindfemur brown medially on inner and outer surfaces; wing membrane brown tint- ed. Body with short erect pale setae. Male. Unknown. Type material.—Holotype, female, MA- LAYA: Sentine Tea Estate, vii—viii 1985, W. Budenberg (OTTAWA). Etymology.—The species name refers to the highly polished, sparsely punctate, “shining” integument. Discussion.—This species differs from cuneifacialis and orchidense below by the shorter pronotum, entirely pale legs, me- tallic dorsal tints and flagellum with erect bristling red setae. Magdalium lucidum is a slender species with the pronotum consid- erably longer than the scutum. Magdalium orchidense Kimsey, NEw SPECIES Fig. 6 Description.— Male (holotype): Body length 4.5 mm. Face (Fig. 6); scapal basin with polished medial stripe and coarse cross- ridges laterally, punctures 0.5—1 puncture diameter apart; malar space 3 midocellus diameters long, with vertical sulcus; suban- tennal distance 0.3 midocellus diameter; flagellomere I 2.5 as long as broad; fla- gellomere II length 1.8 x breadth; occipital carina present dorsally; midocellus 2.3 mid- ocellus diameters from ocular margin; ver- tex with impunctate medial welt 1.5 mi- docellus diameters long; hindocellus 1 di- ameter from ocular margin; pronotum only slightly longer than scutum, with thin sulcus extending anteriorly along medial line, with pit on lateral lobe; mesopleuron with sub- alar fossa, short oblique mesopleural carina and scrobal sulcus long, narrow and paral- lel-sided; notal punctures nearly contiguous to | puncture diameter apart, larger and fur- ther apart on head and pronotum than scu- tum; notauli deep and straight posteriorly; metanotum as long as scutellum; propo- deum with short blunt posterolateral cor- ner; tergum II punctures sparse and 2-4 puncture diameters apart, except for im- punctate medial stripe; terga IHII-V with posterior band of punctures. Head, thorax and abdomen black with greenish bronze tints on vertex and thoracic nota; scape and pedicel red; flagellum black; legs pale brown. Body with erect brown setae. Female: Same as male, except malar space 4 midocellus diameters long; flagel- lomere I 1.9 x as long as broad, flagellomere II 0.8 x as long as broad; scape and pedicel red, flagellomere I pale greyish; rest of fla- gellum black; legs reddish brown; wings brown stained, and propodeal corners prominent and acute. Type material.—Holotype, male, TAI- WAN: Orchid Is. (Batel Tobago), 5-9 July 1991, C. K. Starr (OTTAWA). Paratypes: 1 female, 5 males, same data as type (OT- TAWA, DAVIS). Etymology.—This species is named after the site of collection, Orchid Island. Discussion.— This species has a shorter pronotum than other species where the pro- notum is considerably longer than the scu- tum, entirely pale legs, metallic dorsal tints and flagellum with erect bristling red setae, which will immediately distinguish it from cuneifacialis and lucidum. These are much more slender species with the pronotum considerably longer than the scutum. Perissosega flavipes Kimsey, NEw SPECIES Fig. 2 Description. — Female (holotype): Body length 4.5 mm. Face (Fig. 2); scapal basin with coarse cross-ridges laterally, punctures 1-2 puncture diameters apart; malar space 3.5 midocellus diameter long, with vertical sulcus; subantennal distance 0.5 midocellus diameter; flagellomere I 2.4x as long as broad; flagellomere II length 0.7 x breadth; occipital carina present dorsally; midocellus 2 midocellus diameters from ocular margin; hindocellus | diameter from ocular margin; VOLUME 97, NUMBER 3 pronotum only slightly longer than scutum, with thin sulcus extending anteriorly along medial line, with pit on lateral lobe; meso- pleuron with subalar fossa, omaulus pres- ent; scrobal sulcus long, narrow and paral- lel-sided, about 7 x as long as wide; punc- tures on vertex and notum 0.5-1 puncture diameter apart; notauli deep and straight posteriorly; metanotum three-fourths as long as scutellum; propodeum with long slender posterolateral spines; tergum II punctures small and 0.5—1.5 puncture diameters apart, except for impunctate medial stripe; terga IIJ—V with posterior impunctate. Head, tho- rax and abdomen black; coxae, except ba- sally, and rest of legs bright yellow; scapal apex yellow, rest of scape brown; pedicel and flagellomere I yellow, remaining flagel- lomeres dark brown; tegula and basal wing veins yellow, apical wing veins dark brown; wing membrane brown tinted. 595 Type material.— Holotype, male, THAI- LAND, 180 km ne Bangkok, Khao Ysi Natl. Park, 780 m, 11-18 April 1990, malaise trap, B. V. Brown (OTTAWA). Etymology.—The name flavipes, refers to the bright yellow legs. Discussion.—This is the second species placed in Perissosega. It differs from vena- blei Krombein in a number of major fea- tures, including the presence of a scrobal sulcus and omaulus, larger size (4.5 mm versus 3 mm), and the wings are evenly brown tinted instead of maculate. In addi- tion, the legs are bright yellow instead of black or brown as in venablei. LITERATURE CITED Kimsey, L. S.and R. M. Bohart. 1991 (1990). Chry- sidid Wasps of the World. Oxford Univ. Press. 652 pp. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 596-602 A REVIEW OF THE GENUS REMENUS RICKER (PLECOPTERA: PERLODIDAE), WITH THE DESCRIPTION OF TWO NEW SPECIES Boris C. KONDRATIEFF AND CHARLES H. NELSON (BCK) Department of Entomology, Colorado State University, Fort Collins, Colorado 80523; (CHN) Department of Biological and Environmental Sciences, 615 McCallie Av- enue, The University of Tennessee, Chattanooga, Tennessee 37403. Abstract. —The formerly monotypic Nearctic genus Remenus is revised to include three species: R. bilobatus (Needham and Claassen), Appalachian in distribution; R. kirchneri n. sp., from Virginia; R. duffieldi n. sp., from Georgia. The male and female are described for each species, and the descriptions are supported by illustrations and SEM photomi- crographs. Key Words: Since Ricker’s (1952) classic review of the Perlodinae, the Nearctic genus Remenus has been considered monotypic. Stark and Szczytko (1984) assigned Remenus to the Diploperlini based on typical turtle-shaped eggs and the male seventh sternum which is produced into a distinct lobe. The only species, R. bilobatus (Needham and Claas- sen), has been reported along the Appala- chians from Connecticut to South Carolina and Georgia (Stark, Szezytko and Baumann 1986). During the preparation of a chapter on the Perlodinae of Eastern North America for an upcoming publication, it became ap- parent that more than one form was in- volved. Morphology and terminology follow Stark and Szczytko (1984). Abbreviations for de- positories of specimens are: Cornell Uni- versity Insect Collection, Ithaca, New York (CUIC); C. P. Gillette Museum of Arthro- pod Diversity, Colorado State University (CSU); National Museum of Natural His- tory (USNM); Charles H. Nelson Collection (CHN); Bill P. Stark Collection, Missis- sippi College, Clinton (BPS); and Virginia Plecoptera, stoneflies, Perlodidae, Remenus, Nearctic Polytechnic Institute and State University (VPI). Remenus bilobatus (Needham and Claassen) Fig. 1-3, 8a, 8b, 11, 12 Perla bilobata Needham and Claassen, 1925: 95. Holotype male: New York: Old Forge, 7-19 July 1905 (CUIC, #1153), exam- ined. Description.— Male: Length of forewing 8-9 mm; length of body 7-9 mm. General body color in alcohol light brown (pale yel- low-brown in life). Head and pronotum with light brown pattern (Fig. 1). Wings hyaline; veins light brown. Male genitalia: Tenth tergum weakly cleft for 3 of its length; hemitergal lobes short, clothed with short setae and sensilla basi- conia (Fig. 3). Ninth tergum with short setae and sensilla basiconia (Fig. 3). Paraprocts reduced. Lateral stylets absent. Epiproct covered basally with dense yellow spinulae, enclosed by a golden setae-lined pocket, ter- minating in a threadlike lash (Figs. 2, 3). VOLUME 97, NUMBER 3 Female: Length of forewing 10-11 mm; length of body 9-10 mm. General color and external morphology similar to the male. Subgenital plate broadly rounded to more narrowly rounded, produced 7% the length of the 9th sternum, with basal lateral crease (Figs. 8a, 8b). Egg: Turtle shaped. Chorion relatively smooth with keel extending from lid to pos- terior 7; lid with irregular follicle cell im- pressions (Figs. 11, 12). Diagnosis.—Males of R. bilobatus are easily distinguished from both R. kirchneri n. sp. and R. duffieldin. sp. by the threadlike lash of the epiproct apex (Figs. 2, 3). The length of the lash varies, reaching to the seventh or eighth tergum when fully ex- tended. Most preserved specimens have only a short length of the lash exposed as figured by Hitchcock (1974, Fig. 309) or partially extended as illustrated by Kondratieff and Voshell (1982, Figs. 26-27). Additionally, the fragile terminal lash 1s occasionally bro- ken and missing from the golden setae-cov- ered basal portion of the epiproct in pre- served specimens. Females can usually be distinguished by the large subgenital plate with an incomplete basal transverse crease (Fig. 8a, 8b). Remarks.—Needham and Claassen (1925) originally described Perla bilobata from “many specimens” from Old Forge, New York and Black Mountain, North Car- olina. The Black Mountain specimens are typical R. bilobatus. Apparently this small perlodine ranges from New England to Al- abama and Tennessee along the Appala- chians and into the Piedmont Plateau Phys- iographic province. Remenus bilobatus oc- curs in small to medium-sized cool streams. Adults are active from May to July, but never commonly collected. Material examined.—ALABAMA: Cle- burne Co., small stream above lake, Cheaha State Park, 14 May 1988, B. Kondratieff and R. F. Kirchner, 3 males, 3 females (CSU). CONNECTICUT: Tolland Co. Storrs, 18 June 1954, J. A. Slater, 1 male, Figs le Remenus bilobatus, head and pronotum. 1 female (USNM); brooklet crossing Rt. 48, Killingworth, 18 June 1965, S. W. Hitch- cock, 4 males (USNM); Madison, near Rt. 80, 18 June 1965, S. W. Hitchcock, 1 male (USNM). MARYLAND: Fredrick Co., Big Hunting Creek, Thurmont, 4 June 1987, R. M. Duffield, 1 male (CHN); same but 22 June 1988, 1 female (CHN). NORTH CAR- OLINA: Black Mountain, July—Aug. 1912, Beutenmuller, 2 males (previously cleared), 1 female (CUIC, Paratypes #1153); Hay- wood Co., Sterling Creek, Rt. 23 (Rd 1397), GSMNP, 10 July 1983, B. Kondratieff and R. F. Kirchner, 2 females (VPI); Macon Co., Jarrett Creek, Arrowwood Glade, Nanta- hala National Forest, 25 May 1993, B. Kon- dratieff and R. F. Kirchner, | male (CSU); Transylvania Co., South Fork of Mills Riv- er, FR 1206 off Rt. 276, 8 July 1981, B: Kondratieff and R. F. Kirchner, 2 males, 3 females (VPI). PENNSYLVANIA: Adams Co., near Fairfield, 18 June 1950, D. G. Shappiro, 2 males (USNM). SOUTH CAR- OLINA: Oconee Co., Townes Creek, Road 710, Sumter National Forest, 24 May 1993, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fy H{ Ky 4 I: Figs. 2-7. Qyxhl ay us eZ4 BSAA <8 S Remenus, male terminalia. 2. R. bilobatus, lateral (epiproct fully extended). 3. R. bilobatus, dorsal. 4. R. kirchneri, lateral. 5. R. kirchneri, dorsal. 6. R. duffieldi, lateral. 7. R. duffieldi, dorsal. Kondratieff and Kirchner, 1 female (CSU); Pickens Co., Wildcat Creek, 9 km NW Clemson, 24 May 1981, B. P. Stark et al., 6 males (BPS); TENNESSEE: Polk Co., GoForth Creek, Rt 64, Cherokee National Forest, 3 June 1993, C. H. Nelson, 1 male (CHN); VIRGINIA: Bland Co., Wolf Creek, Rt. 61, 10 June 1978, B. Kondratieff, 1 male (VPI); Montgomery Co., Toms Creek, Rt. 655, 29 May 1978, B. Kondratieff, 1 male (reared) (VPI); small spring flowing into Craigs Creek, 2.7 km off Rt. 460 on Rt. 621, 17 June 1980, B. Kondratieff, 1 male (VPI); Tazewell Co., East Fork Cove Creek, Rt. 662, 12 June 1983, B. Kondratieff and R. F. Kirchner; i male: (VPI): VOLUME 97, NUMBER 3 599 ee ) / | (lf? | 1, VW/ - \ ~ } le 4 X (Z SAS ~ ( \ \\ ; (AMAL | \ / vl Lt Dy i j il at ( Se aM \ Cat Wty tl fT 1) hy \ i \ i / / yy \ ie ) ii 7) 1) 1) DK ) tilegeres Mr | ptt PLE 7 Ai wk iy! 1] j eh ea f . 1 | , i Na Ma ioe ; j i En aD \ rf | VA Stn 1 at ALL) YEN WA AND TS eit arse SI open | SS tt anise YO EN ANN TT AZ atin a WSS Sy HI INC ee NT Ae A A) ARS LE TY TEV hoped OSs eet Vij, HIT] 4 pote e 7, Y S CRN yh I ei ante yl fh ee jp gle, = ey Bei itl Le dh hag ee Sets ey ow! es) , ate acy en hy fh ALYY: \ . : RG peer fre hin Zs 87, Lf ea ae ee yet / Wot, Yl OV j ATUL ty aa clipe eat \ eat MA! 1 YOY 2 j Hy] DA with ROMS ie ! I) / 4 Wie a i ee : 8 b val | ) hy OS WS } /] Wd SSA i Ne =a Oe / oi ay, \\ | 4, Qe NN 4 / 1 y = y = | . 9 ae y | y, / | yMif gead Z [! ee aad Ved oy ZY pela opal pie Wilts L\ 1 l | Cali) E seleelnes // Pe ay ee ene { 4 | hg < Si ea fe 4 Ree pb Sty Laaien ds RENE Ss d [ | / Say iy iy / Lah); Y ry SRSA J NILA SONS ay 1 eke Wap) WSS 7 \ ‘ Gi, ALS A | j SSS See! (oe Ee a MM Ly le i SSS B \ ? WA eI) J SIS ie nS : no ‘ WG fy \ \ lige SSS \ Ww cb Tt Lae LL We al j / 6 LIE sc> x Mer nk I | La ; : Mie 7 10 Figs. 8-10. Remenus, female subgenital plate. 8a. R. bilobatus, Maryland. 8b. R. bilobatus, South Carolina. 9. R. kirchneri. 10. R. duffieldi. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON eous kirchneri Kondratieff and Nelson, NEw SPECIES Figs. 4, 5, 9 Description.— Male: Length of forewing 8—9 mm; length of body 8-9 mm. General body color in alcohol light brown (pale green- brown in life). Head and pronotum with light brown pattern similar to Fig. 1. Wings hyaline; veins light brown. Male genitalia: Hemitergal lobes short, clothed with short setae and few sensilla basiconia (Fig. 5). Ninth tergum with long setae and few scattered sensilla basiconia (Fig. 5). Paraprocts reduced. Lateral stylets absent. Epiproct in dorsal view, pear-shaped (Fig. 5), covered with fine appressed spi- nulae; in lateral view, subparallel, slightly constricted at apex (Fig. 4). Female: Length of forewing 10-11 mm; length of body 9-10 mm. General color and external morphology similar to the male. Subgenital plate broadly rounded, produced 7; length of 9th sternum, with basolateral margins parallel for at least 2 of plate length (Fig. 9). Egg: Unknown. Types. —HOLOTYPE male: Virginia, Patrick Co., Little Rock Castle Creek, Rock Castle Gorge National Recreation Area, 24 May 1990, B. Kondratieff and R. F. Kirch- ner. PARATYPES: Same as holotype, 1 male, 1 female; Floyd Co., spring-fed stream entering Little River, Rt. 686, 8 June 1978, B. Kondratieff, 1 male (VPI); small spring seep, Rt. 221, Roadside Park, 4 mile S of Floyd, 6 July 1980, B. Kondratieff, 2 males (CHN); small spring-fed stream, 6 miles E of Floyd, Rt. 221, 28 June 1981, B. Kon- dratieff, 1 male, 1 female (VPI); Patrick Co., spring-fed tribs. of Little Rock Castle Creek, Rt. 605, 10 May 1983, B. Kondratieff, 1 male (VPI); small spring-fed stream into Tallant Reservoir, Dan River, 2 August 1982, B. Kondratieff, 1 female (VPI). The holotype will be deposited in the USNM. Paratypes will be deposited in CSU, CHN, and VPI collections. Etymology.—The authors take great pleasure in naming this species for our friend, Ralph F. Kirchner, Huntington, West Virginia, in recognition of his many contri- butions to the study of North American stoneflies. Diagnosis.—Males of R. kirchneri are easily distinguished from R. bilobatus by the absence of a terminal epiproctal lash, and from R. duffieldi n. sp. by the pear-shaped epiproct (Fig. 5). The smaller subgenital plate with parallel basolateral margins (Fig. 9) distinguishes the females from the other two species. Remarks.—This species is only known from small headwater spring-fed streams or seeps of the lower Blue Ridge physiographic province of Virginia. At the type locality, another apparently endemic stonefly species of this area, Sweltsa voshelli Kondratieffand Kirchner is found. Other common stoneflies at the type locality are Pe/toperla tarteri Stark and Kondratieff and Tallaperla maria (Needham and Smith). Remenus duffieldi Nelson and Kondratieff, NEW SPECIES Figs..6, 710; 135 14 Description.— Male: Length of forewing 8-9 mm; length of body 8-9 mm. General body color in alcohol light brown (pale green in life). Head and pronotum with light brown pattern similar to Fig. 1. Wings hyaline; veins light brown. Male genitalia: Hemitergal lobes short, clothed with short setae and few sensilla basiconia (Fig. 7). Ninth tergum with long setae (Fig. 7). Paraprocts reduced. Lateral stylets absent. Epiproct elongate, slightly ex- panded apically (Figs. 6, 7) with long hair- like spinulae on apex, with dorsal sclerite long, narrow, reaching apex (Fig. 6). Female: Length of forewing 10-11 mm; length of body 9-10 mm. General color and external morphology similar to male. Subgenital plate broadly rounded, produced to posterior margin of 9th sternum (Fig. 10). Egg: Turtle-shaped. Chorion with weak follicle cell impressions and dorsal keel ex- VOLUME 97, NUMBER 3 Figs. 11-14. Remenus eggs. 11. R. bilobatus, dorsal (203 x). 12. R. bilobatus, ventral (230 x). 13. R. duffieldi, dorsal (186). 14. R. duffieldi, ventral (178 x). tending from lid to near middle; lid covered with irregular follicle cell impressions (Figs. 13, 14). Types.—HOLOTYPE male: Georgia: Towns Co., Soapstone Creek, Rt 180 near junction Owl Creek Road, Chattahoochee National Forest, 8 June 1994, C. H., C. P., B. A. Nelson. PARATYPES: Same as ho- lotype, 2 males (CHN); Union Co., Slaugh- ter Creek, Rt. 180, Chattahoochee National Forest, 30 May 1994, C. H. Nelson, | fe- male (CHN); Rock Creek, junction Rt 69, 1.5 miles W High Tower Gap, Chattahoo- chee National Forest, 29 May 1990, R. M. Duffield, 2 males (CHN); Soapstone Creek, Junction 180, Chattahoochee National For- est, 26 May 1990, R. M. Duffield, 1 male (CHN); White Co., Andrews Creek, Rt 17/ 75, Andrews Cove Campground, Chatta- hoochee National Forest, 8 June 1994, C. H. Nelson, 1 male, 3 females, 1 nymph (CHN). The holotype will be deposited in the USNM. Paratypes will be deposited in CSU and CHN collections. Etymology.—This species 1s named for Dr. Richard Dufheld, Howard University, Washington, D.C. who first collected spec- imens of this interesting new species. Diagnosis. — Males of R. duffieldi are eas- ily distinguished from both R. bilobatus and R. kirchneri by the elongate almost club- like epiproct (Figs. 6, 7). Females of R. duf- fieldi can be separated from both R. bilob- atus and R. kirchneri by the broadly round- ed subgenital plate reaching the posterior margin of ninth sternum (Fig. 10). Remarks.—The nymph of R. duffieldi is similar to the nymph of R. bilobatus as de- scribed and figured by Stewart and Stark (1988). Like R. bilobatus, R. duffieldiis found in small to mid-sized montane Appalachian streams. Other stonefly species collected with this new species were Pteronarcys scotti PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ick Tallaperla cornelia (Needham and Smith), Acroneuria abnormis (Newman), » bulbosus (Frison), Alloperla usa Ricker, Sweltsa lateralis (Banks), Amphi- nemura wui (Claassen), Leuctra biloba Claassen, and Leuctra alexanderi Hanson. With the inclusion of the two new species, R. duffieldi and R. kirchneri, adult males of the genus Remenus are defined by the fol- lowing combination of characters: (1) tenth tergum cleft, ' of its length producing short hemitergal lobes, (2) well-developed ventral lobes on sterna seven and eight and (3) east- ern Nearctic in distribution. Females of the genus are difficult to distinguish from other Perlodidae, without associated males, es- pecially Jsoperla, but the combination of the dorsal head pattern, small size (body length 9-10 mm), and subgenital plate form are usually diagnostic. KEY TO REMENUS SPECIES 1. Epiproct of male terminating in a threadlike lash (Figs. 2, 3); subgenital plate of female broadly to narrowly rounded, with basal trans- verse crease incomplete (Figs. 8a, 8b) — Epiproct without terminal lash (Figs. 4, 5 and 6, 7); apex of subgenital plate broadly rounded, without basal crease (Figs. 9, 10) ............ 2 2. Epiproct in dorsal view, pear-shaped (Fig. 5); subgenital plate not reaching posterior margin of 9th sternum, with sides subparallel (Fig. 9) Cy Nee A PRRs ond eT ee ent Sra R. kirchneri — Epiproct, in dorsal view elongate, sides almost parallel (Fig. 7); subgenital plate, reaching pos- terior margin of 9th sternum, broadly rounded (Fig. 10) R. duffieldi ACKNOWLEDGMENTS We thank E. Richard Hoebeke, Cornell University for the loan of type material of R. bilobatus. We are also grateful to Oliver S. Flint, Jr., National Museum of Natural History, Washington, D.C.; Bill P. Stark, Mississippi College, Clinton, Mississippi; and J. Reese Voshell, Virginia Polytechnic Institute and State University, Blacksburg, Virginia for the loan of valuable material. Bill P. Stark, and Richard W. Baumann, Brigham Young University are thanked for reviewing the manuscript. Alison Ander- son-Williams provided the illustrations. Dr. Robert E. Lee, Department of Anatomy and Neurobiology, Colorado State University, graciously provided the SEM photomicro- graphs. LITERATURE CITED Hitchcock, S. W. 1974. Guide to the insects of Con- necticut. Part VII. The Plecoptera or stoneflies of Connecticut. Bulletin of the State Geological and Natural History Survey of Connecticut 107: 1- 262. Kondratieff, B. C. and J. R. Voshell, Jr. 1982. The Perlodinae of Virginia, USA (Plecoptera: Perlod- idae). Proceedings of the Entomological Society Washington 84: 761-774. Needham, J. G. and P. W. Claassen. 1925. A mono- graph of the Plecoptera or stoneflies of America North of Mexico. Thomas Say Foundation 2: l- 397. Ricker, W. E. 1952. Systematic studies of Plecoptera. Indiana University Publications Science Series 18: 1-200. Stewart, K. W. and B. P. Stark. 1988. Nymphs of North American Stonefly Genera (Plecoptera). Thomas Say Foundation, Entomological Society America 12. 460 pp. Stark, B. P., S. W. Szczytko, and R. W. Baumann. 1986. North American stoneflies (Plecoptera): Systematics, distribution and taxonomic refer- ences. Great Basin Naturalist 46: 383-397. Stark, B. P. and S. W. Szczytko. 1984. Egg mor- phology and classification of Perlodinae (Plecop- tera: Perlodidae). Annales de Limnologie 20: 99- 104. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 603-625 TAXONOMIC REVIEW OF APPLE-FEEDING SPECIES OF PHYLLONORYCTER HUBNER (LEPIDOPTERA, GRACILLARIIDAE) IN NORTH AMERICA JEAN-FRANCOIS LANDRY AND DAvip L. WAGNER (JFL) Agriculture Canada, Centre for Land and Biological Resources Research, Central Experimental Farm, Ottawa, Ontario K1A 0C6, Canada; (DLW) Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06269. Abstract. — Four species of Phyllonorycter whose larvae mine apple leaves are recognized in North America: P. blancardella (F.), P. crataegella (Clemens), P. elmaella Doganlar and Mutuura, and P. mespilella (Hiibner). The former two species are widely distributed in the eastern United States and southeastern Canada. We confirm the presence of P. blancardella in the Pacific Northwest, earliest records being from Oregon in 1985, Wash- ington in 1986, and British Columbia in 1987. Phyllonorycter mespilella occurs over much of western North America, from California north to British Columbia, and east to Utah and New Mexico. Phyllonorycter elmaella is known from Oregon, Washington, and British Columbia. Lithocolletis malimalifoliella Braun is synonymized under P. crataegella. Rec- ords of Phyllonorycter sorbi (Frey) in North America and of P. scudderella (Frey and Boll) on apple are regarded as erroneous. Phyllonorycter deceptusella (Chambers), also wrongly mentioned as feeding on apple is here regarded as a nomen dubium. We provide keys based on external features and on genitalia of adults, as well as diagnoses and notes on the geographical distribution and biology of each species. Key Words: Lepidoptera, taxonomy, biology, distribution, apple, tentiform leaf miner, North America, Gracillariidae, Phyllonorycter With nearly 80 described species (Davis 1983) and dozens of as yet undescribed en- tities, the genus Phyllonorycter is one of North America’s most taxonomically di- verse genera of leafmining Lepidoptera. The larvae feed on a variety of woody peren- nials, usually forming tentiform mines on the underside of leaves (Needham et al. 1928). Eight species have been reported to feed on apple leaves in North America: P. blancardella (F.), P. crataegella (Clemens), P. malimalifoliella (Braun), P. sorbi (Frey), P. mespilella (Hiibner), P. elmaella Dogan- lar and Mutuura, P. deceptusella (Cham- bers), and P. scudderella (Frey and Boll) (Stultz 1964, Pottinger and LeRoux 1971: 6-9, Doganlar and Mutuura 1980, Weires et al. 1980). Phyllonorycter blancardella, P. crataegella, and P. elmaella have often been reported as orchard pests. A fourth species, of European origin, P. mespilella, occurs abundantly on apples and related rosaceous plants, yet its presence in western North America has been all but overlooked. Our studies indicate that P. mespilella has been established in the western United States for at least 50 years, and that recent literature treating P. elmaella (e.g. Weires and For- shey 1978, Orphart 1982, Hoyt 1983, Bar- rett and Jorgensen 1986, Barrett and Brun- ner 1990, Jones 1991) likely refers to P. mespilella. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON right valva sternum 8 (dashed outline) right costal tegumen left valva process vinculum (dashed) left costal process transtilla Q) Fig. 1. Male genitalia of Phyllonorycter species, unspread, showing the position of tegumen (stippled) and sternum VIII (dashed); aedeagus omitted. Pottinger and LeRoux (1971) compre- hensively reviewed literature on the apple- feeding Phyllonorycter in North America up to 1964. They pointed out that there was much confusion, particularly among eco- nomic entomologists, as to what species they actually worked with, and that virtually all papers lacked reliable determinations. All the apple-feeding species are superficially very similar to one another. There are small differences in coloration and pattern among species, but these are often muddled by pro- nounced intraspecific, especially brood-re- lated, variation. Stultz (1964) provided the first North American work in which male genital characteristics of P. blancardella and P. crataegella were compared and illustrat- ed. Pottinger and LeRoux (1971) distin- VOLUME 97, NUMBER 3 guished and illustrated the male genitalia of P. blancardella, P. crataegella, and P. mes- pilella. Neither work compared the female genitalia of these species. Doganlar and Mu- tuura (1980) provided illustrations of both the male and female genitalia of P. e/maella but they did not compare them to other apple-feeding Phyllonorycter. In order to clear the taxonomic confusion surrounding the apple-feeding species of Phyllonorycter in North America, we pre- sent diagnoses of adults, keys for separating them based on genital features and on adult coloration, and summaries of biological and distributional data for the species. We re- gard as valid four species for which we have examined specimens reared unequivocally from larvae mining apple (Malus spp.) leaves: P. blancardella, P. crataegella, P. el- maella, and P. mespilella. The first and the last are introduced from Europe; the other two are believed indigenous species, which transferred to apple from Crataegus and other native Rosaceae. Of the remaining species reported in the literature, one is a junior synonym [P. malimalifoliella (Braun, 1908a) = P. crataegella (Clemens, 1859), new synonymy], two are thought to have been reported in error [P. sorbi (Frey) and P. scudderella (Frey and Boll)], and the sta- tus of one remains uncertain [P. deceptusella (Chambers)]—it most resembles cherry feeding members of the genus. MATERIALS AND METHODS Specimens were examined from the fol- lowing collections: ANSP Academy of Natural Sciences of Philadelphia, 19th & Parkway, Logan Square, Philadelphia, PA 19103 (D. Azuma). Canadian National Collection, Agriculture Canada, Centre for Land and Biological Resources Research, C.E.F., Ottawa, Ontar- 10 KIA 0C6 (J.-F. Landry). Gerfried Deschka Collection (pri- CNCI DESC 605 vate), Resselstrasse 18, A-4400 Steyr, Austria. Lyman Entomological Museum, Macdonald College, 21111 Lake- shore Drive, Ste-Anne-de-Belle- vue; OQuebece Hox 1C0M(C.-c. Hsiung). Museum of Comparative Zoolo- gy, Harvard University, Cam- bridge, Massachusetts 02138 (P. D. Perkins). Oregon State University, Corval- lis, Oregon 97331 (J. D. Lattin). Essig Museum, Wellman Hall, University of California, Berke- ley, California 94720 (J. A. Pow- ell). University of Connecticut, Storrs, Connecticut 06269 (D. L. Wag- ner). United States National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 (D. R. Davis). LEMC MCZC OSUO CISC UCON USNM We have studied the types of P. cratae- gella (Clemens), P. deceptusella (Cham- bers), P. elmaella Doganlar and Mutuura, and P. malimalifoliella (Braun) as well as specimens of European origin of P. blan- cardella, P. mespilella, P. sorbi, and P. cy- doniella (F.), and the Japanese apple-feed- ing P. ringoniella (Matsumura). Genitalia were dissected following the standard method described by Robinson (1976), stained with both Orange G (in 30% ethanol; enhances sclerites) and chlorazol black (in 70% ethanol; enhances mem- branes), and mounted in Euparal. In mount- ing most male genitalia, the tegumen and sternum VIII were left attached to the ab- domen. The rest of the genitalia (vinculum- valvae-aedeagus) were separated and mounted dorsal side up after severing the transtilla medially and prying the valvae apart, in order to expose dorsally the costal processes, thereby providing a diagnostic view of the genitalia (e.g. Figs. 2-4). Female PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON AE RV LV VIN Figs. 24. Male genitalia of Phyllonorycter species, dorsal aspect, with tegumen, sternum VIII, and setae of valvae omitted; transtilla cut medially with valvae and their processes spread flat; posterior end oriented towards top of page. Abbreviations are as follows: AE, aedeagus; LCP, left costal process; LV, left valva; RCP, right costal process; RV, right valva: TR, transtilla; VIN, vinculum. 2, mespilella (slide MIC 2071); 3, crataegella (slide JFL 785); 4, elmaella (slide JFL 860). VOLUME 97, NUMBER 3 genitalia were left within the abdomen and mounted ventral side up (removing the gen- italia can cause much distortion or damage). Records that we mention below have been confirmed by genital dissections and are de- posited in the collections indicated in pa- rentheses. Drawings were prepared with a drawing tube mounted on a Nikon Optiphot com- pound microscope, at magnifications of 100x or 200x. Drawings were prepared from single specimens whenever possible, but parts have been re-arranged slightly on some drawings to compensate for distor- tions present in slide preparations. In the illustrations, the posterior end is directed upward on the page. Male genitalia are shown in dorsal aspect, females in ventral aspect; hence in male illustrations, the ac- tual right side appears on the left. Positions of the costal processes are affected by prep- aration and therefore are not indicative of specific differences. Photographs of the adults were taken with a Nikon F3 camera fitted with a Leitz 63 mm objective, using Kodak Technical Pan film (set at 12 ASA) and processed in Kodak Technidol. GENERAL ASPECTS OF LiFe HisTORY The species in this group have closely similar life histories. The eggs, about 0.3 mm in length, are flat and oval, and laid on the lower leaf surface. All species have three sap-feeding and two tissue-feeding larval in- stars. The legless sap-feeding instars expand the mine to its full extent, separating the lower leaf surface from the overlying par- enchymic tissues. From below the mine ap- pears waxy white to greenish; the early in- star mine is not visible from above. The fourth and fifth tissue-feeding instars are legged with a fully developed spinneret that they use to lay down silk within the mine, that pulls the edges of the mine in- wards, drawing the mine into an elongate tentiform bubble. The lower leaf surface is 607 drawn into a set of four to numerous closely set creases. The larvae remove small patch- es of tissue up to the upper leaf surface, giving the upper side of the leaf a shot-hole or skeletonized appearance, and hence the common name for the group, the “‘spotted tentiform leafminers.” The size of the mines may vary depending on the host, e.g. on apple cultivars with thick leaves the mine may average considerably smaller. The fifth instar spins a sparse cocoon against the up- per side of the mine and pupates within a few days. The pupa is extruded through the lower leaf surface prior to eclosion of the adult. The species in this group are multivoltine with pupae overwintering within the mines. At least two broods and as many as five broods, are inferred. For example, in Cali- fornia, P. mespilella has up to five genera- tions in the Central Valley, three to four broods in coastal areas, and as few as two generations in higher elevation apple and pear orchards in the Sierra Nevada (L. Var- ela, in litt. 1993). Three broods are reported for both P. crataegella and P. blancardella in the northeastern United States (Maier 1985, Maier and Davis 1989). The presence of summer-diapausing sap-feeding instars is reported for second-generation individuals of P. blancardella (Laing et al. 1986) and probably occurs in other members of the group as well (Maier and Davis 1989). For more information on biology, consult Pot- tinger and LeRoux’s (1971) monographic treatment of P. blancardella; much con- tained therein is likely applicable to the oth- er apple-feeding species. Some populations of Phyllonorycter cra- taegella and P. blancardella have developed insecticide resistance in sprayed orchards in eastern North America (Maier 1983, Pree et al. 1986, Barrett and Brunner 1990). In California P. mespilella has developed re- sistance to both guthion and vygate (L. Var- ela, in litt.). Thus, accurate species identi- fication may be necessary if effective control is intended. We have acquired numerous collections of active mines throughout our study from apple trees growing in Arkansas, British Co- lumbia, California, Connecticut, New York, Ontario, Oregon, Utah, Vermont, and Washington, and in only one case has a col- lection yielded more than a single species. This is surprising in that both P. blancar- della and P. crataegella are broadly sym- patric over much of eastern North America and P. blancardella, P. elmaella, and P. mespilella over the Pacific Northwest. Weires et al. (1980) reported several sites in New York where both P. crataegella and P. blancardella co-occurred although the latter species predominated in most or- chards. Early collections from Nova Scotia (pre- 1950) were all P. crataegella, but beginning in 1957, P. blancardella began outbreaking in Nova Scotia (Stultz 1964). All recent col- lections from eastern Canada in the CNCI have been of P. blancardella. Stultz (1964), in his studies of apple-feeding Phyllonoryc- ter in Nova Scotia, was unable to locate a single population of P. crataegella on apple during the several years of his survey efforts. In Europe, P. blancardella tends to replace other species of Phyllonorycter once estab- lished in an area (G. Deschka, in litt. 1993). If competitive displacement 1s a common outcome among these apple-feeding spe- cies, the situation in the Pacific Northwest should prove especially interesting in that both P. blancardella and P. mespilella are introduced-the former presumably has been present little more than a decade. Indeed, P. elmaella may have been common on ap- ple prior to the arrival of these two moths, but presently we do not know of a single orchard where P. e/maella can be reliably obtained. Related to the above may be the fact that the apple-feeding species may respond to a common, or at least to common compo- nents of the female sex pheromone. A fe- male-produced sex attractant of P. blancar- della, identified by Roelofs et al. (1977) as PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (E10)-Dodeceny] acetate, has also been used to survey for adults of P. mespilella (Cos- sentine and Jensen 1992, Gries et al. 1993). However, it is scarcely, if at all, attractive to males of P. crataegella (Weires et al. 1980). In Europe Deschka (in litt. 1993) noted that related Rosaceae-feeding species of Phyllonorycter (but not all of them) have one and the same pheromone, and the males of two or more species may be attracted to a single female. REMARKS ON IDENTIFICATION External characters.—Considerable phe- notypic differences occur both within and between broods such that it is virtually im- possible to identify reliably all adults using only scaling characters. But because color characters have been used extensively in the past, we provide a discussion of features that have diagnostic value. Series rather than individuals should be examined whenever possible. Antennal and hindleg coloration applies to the scaling of the dorsal surfaces. The white scales of the mesoscutellum are easily abraded during the process of pin- ning. Individuals of the overwintering or spring brood tend to be larger and more darkly marked. In some specimens of P. blancar- della the orange forewing scales may be mostly replaced or overlaid with dark scales. The legs and antennae also tend to be more fuscous in the spring generation. Summer generation moths vary greatly in size, but average smaller (for each species, our mea- surements are based on the examination of more than 20 specimens, except where not- ed), black scaling is much reduced, and the orange ground color appears paler. Moths emerging in the fall display both spring (few) and summer (most) phenotypes. Genital characters.— Reliable identifica- tion of Phyllonorycter species feeding on po- moid and prunoid Rosaceae should be based on examination of the genitalia. Vouchers with their dissected abdomen and genitalia should always be preserved. Whenever pos- VOLUME 97, NUMBER 3 sible adult moths should be pinned, pref- erably on minutens and staged (double- mounted). They should never be glued on points, as this often renders removal of the abdomen for genital examination difficult or impossible. A simple and rapid method for mounting microlepidoptera is described in Landry and Landry (1994). Dry speci- mens from light traps or sticky specimens from pheromone traps should be relaxed and pinned, preserved in ethanol, or placed into small microvials mounted on pins. In the latter case a tiny plug of cotton within the microvial will reduce excessive move- ment. To dissect and examine the genitalia, the whole abdomen (on dry specimens, easily removed by applying from beneath gentle upward pressure with fine forceps) is soaked in 20% KOH (aqueous solution) for 2-3 hours at room temperature, or for 3—5 min- utes in a warm (but not boiling) water bath. The abdomen is then descaled in 30% eth- anol using very fine camel hair brushes to expose the genitalia; the digested abdominal content should be extruded through the an- terior (proximal) opening by gentle pushes with the brush. A less satisfactory result can be obtained by gently rolling away the ab- dominal contents and scales with the head of an insect pin. It is usually not necessary to separate the genitalia from the abdomen to view diagnostic characters. The genitalia may be then examined in alcohol, glycerine, or lactic acid, and stored in glycerine in a microvial kept with the specimen. Perma- nent slide mounts, such as in Euparal or Canada balsam, are desirable but not es- sential for routine identifications. For the preparator inexperienced with permanent slide mounts, glycerine preservation of dis- sected vouchers is preferable to poor per- manent slides. Before storage into glycerine the abdomen-genitalia should be briefly soaked in lactic acid or in 30% ethanol acid- ified to pH 4 with a few drops of acetic acid, to ensure that all KOH is neutralized, then rinsed in water. 609 Terms.—Terms for genitalia are shown in Figs. 1-3 and 9, those for forewing mac- ulation in Fig. 13. We refer to the sterigma as any sclerotized area that surrounds the female copulatory opening or ostium bur- sae. In Phyllonorycter species treated here, the sterigma derives from modification of the posterior margin of sternum VIII and the attached anterior apophyses. Externally males can be separated from females by examining either the frenulum or the apex of the abdomen. Males have a single frenular bristle, females have two. In males the valvae, tegumen, and sternum VIII, though covered by scales, are usually discernable; ventrally a longitudinal slit is visible where the valvae come together. In females the apex of the ovipositor usually protrudes from the abdomen, appearing as a setose lobe. KEY TO ADULTS BASED ON GENITAL CHARACTERS Kars Miales ey tore Sen rn oe an oe ee 2 ibs Bemalesieirn Sar trys testers ea 5 2a. Right costal process dilated, thicker than left process. Valvae asymmetrical, right valva broader than left valva (Fig. 2) ... P. mespilella 2b. Right costal process slender, comparatively as thin as left process. Valvae symmetrical or nearly so, right valva as wide as left valva (E1gsh3325) ee eta Aer mer ne nals 3 3a. Right costal process (excluding apical spine) subequal to or slightly longer than left costal Process: (h1gs4) Peer ne ree P. elmaella 3b. Right costal process (excluding apical spine) at least twice as long as left costal process (Figs. 3, 5-7) 4a. Spines of costal processes long and slender, setiform. Left costal process very short and stubby (Fig. 5), exceptionally in few speci- mens left process about one-third length of right process (Figs. 6-7) ........ P. blancardella 4b. Spines of costal processes short and stout, spi- niform. Left costal process about half length of right process'\(Fig: 3)) >. ...292. P. crataegella 5a. Posterior margin of sternum VIII markedly projected into truncate cone (Figs. 9-10) ... PES VR Bs Bee fb rope TAL Tae P. blancardella 5b. Posterior margin of sternum VIII not pro- jected or only slightly protruded (Figs. 8, 11- 1D) ais ek, CRS CE CTE 9 Rh | 6 610 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ostium bursae surrounded by ringlike thick- ening (Big) ge es P. mespilella 6b. Ostium bursae without sclerotized ring Ja. Posterior margin of sternum VIII concave (Fig. 11). Base of ovipositor distad of ostium bur- sae membranous, flat. Anterior apophyses sit- uated closer to middle of sternum VIII than to.its lateralsmareins. mo oaseece P. elmaella 7b. Posterior margin of sternum VIII straight or slightly convex. Base of ovipositor distad of ostium bursae sclerotized, medially elevated, laterally steeply sloped, with transverse cutic- ular creases just caudad of ostium bursae (Fig. 8). Anterior apophyses situated closer to lat- eral margins of sternum VIII than to its mid- dle=. Cee nro ee re tere P. crataegella KEY TO ADULTS BASED ON EXTERNAL CHARACTERS This key is provided for convenience, but results should always be checked by ex- amining genitalia. Some variant individuals will not key out. Diagnostic features may not be well rendered in our black and white photographs. Lastly, scale colors lighten ap- preciably as specimens age in collections. la. Forewing with basal patch that tends to bulge toward basal streak (e.g. Figs. 17, 20); me- soscutellum with prominent patch of white scales; basal streak broad, of 4-6 scale rows, often subequal or exceeding width of orange- scaled area between basal streak and costa (Figs? DO=18, 20) he. shee coe P. blancardella lb. Forewing with basal patch narrow, parallel to wing margin (few individuals in series with bulge); mesoscutellum with few or no white scales; basal streak narrow, of 3-4 scale rows, often subequal or much narrower than width of orange-scaled area between basal streak and COStas(HIgs#22—248)20—Si/)i ares ees ett 2 2a. Axis of first costal strigula running to dorsal (posterior) margin, less than twice as long as broad, often touching or confluent with first dorsal strigula (Figs. 27-29); fourth costal strigula often expanded toward wing apex and broader than third; ground color orange; first metatarsomere pale white or gray scaled (rare- ly with subapical dark band) ...... P. elmaella 2b. Axis of first costal strigula running to tornus or Outer margin, usually more than twice as long as broad, infrequently touching or con- fluent with first dorsal strigula (Figs. 22-24, 26, 30-37); fourth costal strigula reduced or absent; ground color coppery, fiery or red or- ange; first metatarsomere with conspicuous black subapical dark band or black scaling dorsad 3a. Antenna mostly pale to fuscous, terminal ar- ticles occasionally darkened; first metatarso- mere mostly pale, often with apical band of dark scales; forewing with apical row of la- melliform scales (before ciliary fringe) dark only in distal half, forming narrow iridescent blue-black band between outermost fasciae (Figs. 30-37); forewing ground color fiery or- ange/omred orange. eacty ee P. mespilella 3b. Antenna mostly dark (especially dorsal sur- face); first metatarsomere dorsally entirely dark (some specimens with light scales apically); forewing with apical row of lamelliform scales dark for at least two-thirds of their length, forming broad iridescent blue-black band be- tween outermost fasciae (Figs. 22-24); ground color with decided metallic or coppery luster eg ee eR Ube Nad | cde he xr! Ae Bt P. crataegella Phyllonorycter blancardella (F.) Figs. 5-7, 9-10, 14-21 Tinea blancardella Fabricius, 1781: 305. Lithocolletis blancardella (F.): Pottinger and LeRoux, 1971. Phyllonorycter blancardella (F.): Bradley, 1972: 10. Davis, 1983: 10. Emmet et al., 1985: 319. Lithocelletis concomitella Bankes, 246. Diagnosis.—In male genitalia (Fig. 5), P. blancardella is recognized by the markedly different costal processes, the right process being slender, with a slender apical spine that is about one-third the length of the pro- cess and extends nearly to the apex of the valva. The valvae are the narrowest of the species here treated. The left process is very short and stubby in most specimens (Fig. 5), but some variation occurs in its length, and exceptionally it can be up to nearly half the length of the right process (Figs. 6-7). Specimens with an unusually long left pro- cess resemble in that respect specimens of P. elmaella, but in the latter the right pro- cess is much shorter and the valvae are broader than in P. blancardella. The apical hooklike lobe of the aedeagus 1s barely sug- 1899: VOLUME 97, NUMBER 3 XA / i> VW Figs. 5-7. ©) Male genitalia of P. blancardella in dorsal aspect with tegumen, sternum VIII and setae of valvae 611 @ omitted; posterior end oriented towards top of page; 5, transtilla cut medially with valvae and their processes spread flat (slide MIC 2077); 6, redrawn from Pottinger and LeRoux (1971, p. 217, fig. 18-B), showing variation in length of left costal process; 7, id. (p. 220, fig. 21-F). gested in P. blancardella (Fig. 5), whereas it is distinct in P. elmaella (Fig. 4). In female genitalia (Figs. 9-10) P. blan- cardella is the most distinctive of the spe- cies. It is recognized by the markedly pro- jected sterigma which forms a truncate cone extended far beyond the posterior margin of sternum VIII. The length and width of the conical projection vary somewhat, but the overall aspect is unlike that of any of the other species. Forewing length: spring generation, 3.1— 4.5 mm (n = 20); summer generation, 3.0- 4.3 mm (n = 20). Antenna often fuscous in winter generation and pale with conspicu- ously darkened terminal articles in summer broods. Mesothoracic scutellum with dis- tinct patch of white scales, 3-6 rows wide. Forewing (Figs. 14—21) with white and black scales more or less equal in number to or- ange scales; black scaling especially prom- inent in spring brood (Figs. 18-20); white basal patch (anal macula) 2—5 scale rows wide, often bulging toward basal streak; first costal strigula usually at least twice as long as broad, axis usually intersecting tornus or outer margin, rarely confluent with first dor- sal strigula (Fig. 16); fourth costal strigula often subequal to third; basal streak broad, of 4-6 scale rows, often subequal or ex- ceeding width of orange-scaled area be- tween basal streak and costa, in some spec- imens, basal streak connected to first dorsal strigula (Fig. 20); outer row of lamelliform fringe scales with distal 2 darkened, form- ing narrow iridescent blue-black band be- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON yutermost strigulae. Hindleg: tibiae sly pale, or faintly darkened; first and od tarsomeres pale or with dark apical Distribution.—We have examined spec- imens of P. blancardella from South Car- olina northward to Nova Scotia and west- ward to Ontario and Illinois, as well as from Oregon, Washington, and the Vancouver District of British Columbia. Records from farther south in the eastern U.S. await ver- ification. Commonly known as the spotted tentiform leafminer, P. blancardella is pre- sumably introduced from Europe and is a pest of orchards in northeastern United States and southern Canada (Pottinger and LeRoux 1971, Weires et al. 1980, Maier and Davis 1989). The first reports of P. blancardella are quite old (Walsingham 1882: 202, Busck 1903: 190) and may be correct although we have not seen the adults upon which they are based. The oldest confirmed records for this species (determinations based on gen- ital dissections) date back to 1957 from Nova Scotia (Stultz 1964), but presumably the insect was established here well before this time. The oldest specimens of P. blan- cardella that we have examined were col- lected in 1959 in Québec and Nova Scotia (CNCI). The occurrence of P. blancardella on the West Coast is probably recent. The first con- firmed West Coast specimens were collected from cultivated apple in November 1985 from Wilsonville, Clackamas Co., Oregon (UCON), August 1986 from Concrete, Skagit Co., Washington (DESC), and Oc- tober 1987 from a nursery orchard in Chil- liwack, Vancouver District, British Colum- bia (CNCI). R. Duncan (pers. comm.) of Agriculture Canada in Victoria noted that a species of Phyllonorycter first appeared as a pest in Victoria in 1987; specimens from infested orchards, which turned out to be P. blancardella, were sent to DLW in the summer of 1988. Thus the western distri- bution of P. blancardella already may be more extensive than our records indicate. Biology.—In North America Phyllono- rycter blancardella feeds on apple, Malus spp., including apple cultivars (Malus syl- vestris (L.) Mill.) and a number of orna- mental crab apple varieties. It is not known to use native Malus (Pyrus) species here. This species is triple brooded in Connecti- cut and southern Canada (Stultz 1964, Pot- tinger and LeRoux 1971, Maier 1984, Trimble 1984). A sex attractant (E10-12: OAc) for this species was identified by Roe- lofs et al. (1977) and is now widely used by fruit growers to monitor populations of this and related Phyllonorycter species. A sec- ond, even more attractive, constituent of the female sex pheromone (E4, E10-12: OAc) was recently identified by Gries et al. (1993). Phyllonorycter crataegella (Clemens) Figs. 3, 8, 22-24, 26 Lithocolletis crataegella Clemens, 1859: 324. Busck, 1903: 190. Braun, 1908b: 301. McDunnough, 1939: 95. Phyllonorycter crataegella (Clemens): Davis 1983: 10. Lithocolletis malimalifoliella Braun, 1908a: 101 (Newsynonymy). Braun, 1908b: 300. McDunnough, 1939: 95. Phyllonorycter malimalifoliella (Braun): Davis, 1983: 10. Lithocolletes [sic] blancardella (F.): sensu Dyar, 1902: 551. Diagnosis.—In male genitalia (Fig. 3), P. crataegella is characterized by the regularly tubular costal processes with short and stout apical spines. The left costal process is about one-third the length of the right one. The apical portion of the right costal process is slightly bent. Some specimens of P. blan- cardella with an unusually long left costal process are the only other members of the group with costal processes of approxi- mately similar relative length as those of P. crataegella, but they are easily distinguished by their slender, setalike spines and narrow valvae. In female genitalia (Fig. 8), P. crataegella VOLUME 97, NUMBER 3 613 elevated portion of ovip Figs. 8-12. Female genitalia of Phyllonorycter species, ventral aspect, posterior end oriented toward top of page. Abbreviations are as follows: AA. anterior apophysis; DB, ductus bursae; OS, ostium bursae; OVIP, base of ovipositor; STG, sterigma. 8, crataegella (slide MIC 2099); 9, blandardella (slide JFL 833); 10, blancardella (slide JFL 817); 11, elmaella (slide JFL 830); 12 mespilella (slide JFL 834). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON characterized by the slight medial protru- :ion of the posterior margin of sternum VIII and the sclerotized and medially broadly elevated base of the ovipositor with a few transverse, sclerotized wrinkles just distad of the ostium bursae. The sides of the ele- vated portion are concave, giving the ap- pearance of lateral ridges. Forewing length: spring generation, 3.1- 4.3 mm (n = 30); summer generation, 2.5— 3.6 mm (n = 30), in series averaging smaller than other species, especially in summer brood and southern parts of range. Antenna uniformly dark in both generations. Me- sothoracic scutellum with 2-3 rows of in- conspicuous whitish scales. Forewing (Figs. 22-24, 26): ground color orange with de- cided metallic or coppery luster; basal area of dorsal margin with narrow line of white scales, rarely more than 2 scale rows in width; first costal strigula usually at least three times as long as broad, axis running to outer margin or apex, rarely confluent with first dorsal strigula; fourth costal strig- ula often subequal to third or absent (Fig. 26); basal streak narrow, seldom more than 4 scale rows in width, always subequal to width of orange-scaled area between basal streak and costa; outer row of lamelliform fringe scales dark for at least 7 their length, forming prominent iridescent blue-black band between outermost strigulae. Hindleg: tibia black above; all tarsomeres blackened. Individuals matching the size and fore- wing pattern of P. malimalifoliella (Fig. 26), with three rather than four costal strigulae, represent variants of the summer brood of P. crataegella. The genitalia are indistin- guishable from those of the latter. Distribution. —Phyllonorycter crataegel- la is widely distributed through apple-grow- ing regions of the East, from the Atlantic Coast westward to Arkansas, northward into southern Ontario and Nova Scotia. Western North American records (Wilson 1915, Braun 1939, Pottinger and LeRoux 1971: 16) remain unconfirmed and are probably in error. This species may be quite abundant in sprayed orchards, in those populations that have developed insecticide resistance (Maier 1983). Biology. —Phyllonorycter crataegella feeds on a variety of prunoid and pomoid Ro- saceae. Maier (1985) reared it from Ame- lanchier spp., Aronia spp., Crataegus spp., Cydonia spp., Malus spp., Prunus spp., Py- rus spp., and Sorbus spp. in Connecticut. Weires et al. (1980) noted that P. crataegella is but weakly attracted to the sex phero- mone of P. blancardella. Populations thoughout much of its range are triple brooded (Beckham et al. 1950, Maier 1981, Maier and Davis 1989). Synonymy and type material exam- ined.—Lithocolletis crataegella: Clemens gave no indication of how many specimens he had at the time of describing the species but his description of the mine as “‘usually [emphasis ours] limited by two leaf veins” suggests that he may have reared more than one specimen. Busck (1903) indicated that a single specimen was present in the Clem- ens Collection. Therefore it 1s advisable to designate that specimen as the lectotype. LECTOTYPE ¢4 in ANSP, here selected, labelled: [1] “26” [pink, handwritten]; [2] “Type 7506/ Lithocolletis/ crataegella/ B. Clemens” [red, partly printed, partly hand- written]; [3] ““Lithocolletis Type!/ cratae- gella/ AB 1902 Clemens” [handwritten by A. Busck]; [4] “‘ genitalia on/ slide 2938/ D. R. Davis” [printed with number hand- written]; [5] “Lectotype ¢ / by D. Davis” [partly printed in red, partly handwritten]; [6] “LECTOTYPE @/ Lithocolletis/ cratae- gella/ Clemens/ sel. J.-F. Landry, 1993”. The specimen is double mounted on cork. It is badly damaged, with the head broken off and stuck to the minuten pin; only the right forewing remains attached to the spec- imen; the metathorax, hindwings, and hind- legs are broken off and in a gelatin capsule pinned with the specimen. The lectotype selection by Davis is unpublished. Type lo- VOLUME 97, NUMBER 3 cality: probably Easton, Pennsylvania (see Hodges 1986: 36). Lithocolletis malimalifoliella: Braun (1908a) did not indicate the number of spec- imens in her type series of L. malimalifol- iella. In Braun’s collection at ANSP four specimens bear her red type labels but with- out indication of a species name. One male and one female are from Cincinnati, Ohio, rearing lot number B88, with emergence dates of 31 August 1907 and 1 September 1907, respectively; they are undoubtedly syntypes. Another male, also from Cincin- nati is from rearing lot B264 with an emer- gence date of 26 May 1908, and cannot be a syntype because the moth issued after the publication date of the original description (March 1908). A female (without abdomen) from Montclair, New Jersey, collected in a trap, is another syntype; Braun (1908a: 101) specifically stated that she had “‘flown spec- imens from Montclair, N.J., which are iden- tical with the bred specimens.” D. R. Davis (in litt.) has selected the B88 male as lec- totype, and the B88 and Montclair females as paralectotypes. Because his designations have not been published, a lectotype is here designated following Davis’ intent. LECTOTYPE ¢ in ANSP, here designat- ed, labelled: [1] ““Cincinnati, O./ Annette F. Braun/ VIII-31. 07”; ““B88/ apple’; [2] “TYPE/ Collection of/ Annette F. Braun” [red, printed]; [3] ““é genitalia on/ slide 3283/ D. R. Davis’; [4] “LECTOTYPE 2/ Lith- ocolletis/ malimalifoliella Braun/ By D. R. Davis’; [5] “Lithocolletis/ malimalifoliella/ Braun” [handwritten in Braun’s hand]; [6] “Figured in/ Moths of America/ North of Mexico” [blue, printed]; [7] “LECTOTYPE 6/ Lithocolletis/ malimalifoliella/ Braun/ sel. J.-F. Landry, 1993”. The indication of a “Moths of America North of Mexico” figure added by Davis as well as Davis’ lectotype selection are unpublished. The specimen is spread and double-mounted on cork. Type locality: Cincinnati, Ohio. All three specimens bear a red label 615 marked: ““TYPE/ Collection of/ Annette F. Braun.” Phyllonorycter elmaella Doganlar and Mutuura Figs. 4, 11, 27-29 Phyllonorycter elmaella Doganlar and Mu- tuura, 1980: 311. Diagnosis.—In male genitalia (Fig. 4), P. elmaella differs from all other species treat- ed here in having slender subequal costal processes with slender spines that extend to about the middle of the valvae. In female genitalia (Fig. 11), the concave, slightly crenulate posterior margin of ster- num VIII, reduction of the sterigma to very narrow rim, lack of thickening of the basal part of the ductus bursae (antrum), and lack of sclerotization in the proximal part of the Ovipositor are diagnostic. Reduction of the sterigma is also found in P. crataegella, but in this species the posterior margin of ster- num VIII is slightly protruded (Fig. 8). In P. elmaella the proximal portion of the ovi- positor is membranous, whereas it 1s scler- otized and has cuticular wrinkles in P. cra- taegella. Forewing length: spring generation, 4.0 mm (n = 1); summer generation, 3.1—4.0 mm (n = 20). Antenna uniformly fuscous in both generations. Mesothoracic scutel- lum with inconspicuous patch of white scales, 2 or 3 rows wide. Forewing (Figs. 27-29): ground color coppery to pale or- ange; basal area of dorsal margin with nar- row line of white scales, rarely more than 3 rows in width; first costal strigula often less than twice as long as broad, axis usually intersecting dorsal margin before tornus, of- ten touching or confluent with first dorsal strigula (Fig. 27); fourth costal strigula often expanded toward wing apex and broader than third; basal streak narrow, 3-4 scale rows, always subequal to width of orange- scaled area between basal streak and costa; outer row of lamelliform fringe scales dark 616 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON basal streak mesoscutal patch Fig. 13. markings. for about 2 their length, forming narrow black band without iridescent blue-black re- flections. Hindleg: coloration varying from straw colored to pale fuscous, if present, fus- cous scales cover entire dorsal surface of tibia and at least first two tarsomeres. Remarks. — The forewing pattern of P. el- maella illustrated by Doganlar and Mutu- ura (1980) is not diagnostic. Considerable variation is present in the type series, which includes individuals from two generations. The wing pattern shown by Doganlar and Mutuura is that of the dark spring form, in addition to being a variant in which the first dorsal and first costal strigulae are broadly confluent medially. The narrow slender valvae of the male genitalia described and illustrated by Do- ganlar and Mutuura (1980) is the result of a preparation artifact and is misleading the valvae are not flattened in their prepara- tions (in CNCI, examined), but merely shown on edge. The valvae do not afford a diagnostic difference from those of the Eu- ropean P. sorbi, as may be interpreted from act basal patch Dorsal aspect of Phyllonorycter species illustrating forewing pattern, with terms used to designate apical lamelliform scales their figure of the latter, which shows the valvae mounted flat. The valvae of P. el- maella are subspatulate and very similar to those of P. sorbi when both are compared in flat view. Differences between P. sorbi and P. elmaella are as follows. In the male of P. elmaella, the costal processes extend to one-third the valva length and the apical spines are nearly as long as the processes; in the male of P. sorbi, the costal processes are relatively longer and extend to about the middle of the valvae and the apical spines are proportionally shorter, at most one-third the length of the costal processes. In the female of P. e/maella, the posterior margin of sternum VIII is concave, finely serrate, and the sclerotization reduced to a very nar- row margin; in the female of P. sorbi, the posterior margin of sternum VIII has a broader sclerotization and is medially pro- duced into a short, blunt cone. There are no significant differences in the signum be- tween these two species, contrary to Do- ganlar and Mutuura’s (1980) statement. Furthermore Doganlar and Mutuura erro- VOLUME 97, NUMBER 3 617 Figs. 14-21. Forewings of apple-feeding Phyllonorycter blancardella. 14, Vancouver, B. C., ex crab apple 4.viii.1992 (CNCI); 15, Mecklenburg, Germany, ex Malus sylvestris, 6.vii.1948 (CNCI); 16, Ottawa, Ontario, ex Malus sp., 9.viii.1984 (UCON); 17, Shenandoah Nat. Pk., Virginia, ex cultivated apple, 12—22.viii.1976 (DESC); 18, Wilsonville, Oregon, ex Malus sylvestris, 23.11.1986 (UCON); 19, Hamden, Connecticut, ex apple, iv.1989 (UCON); 20, Ste. Anne de Bellevue, Québec, ex cultivated apple, 19.v.1963 (CNCI); 21, Ellington, Connecticut, ex ornamental apple, 12.vii.1993 (UCON). 618 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 28 29 Figs. 22-29. Forewings of apple-feeding Phyllonorycter crataegella and P. elmaella. 22, crataegella, South- ington, Connecticut, ex apple, 10-17.iii.1988 (UCON); 23, crataegella, Wallingford, Connecticut, ex ornamental apple, 14—-15.vi1.1993 (UCON); 24, crataegella, Wallingford, Connecticut, ex ornamental apple, 14-15.vii.1993 (UCON); 25, deceptusella, Kentucky, holotype (MCZC); 26, crataegella, Cincinnati, Ohio, ex Malus, 31.viii.1907, lectotype of Lithocolletis malimalifoliella Braun (ANSP); 27, elmaella, Burnaby, British Columbia, ex Malus, VOLUME 97, NUMBER 3 neously referred to Fig. 5 of Kumata (1963) as showing the female genitalia of P. sorbi, but the latter species is not even mentioned in Kumata’s paper. As noted in the Intro- duction, P. sorbi is not known to occur in North America. Distribution. — Phyllonorycter elmaella is presently known only from the Pacific Coast region from central Oregon to southern British Columbia. Doganlar and Beirne (1980) reported that P. elmaella occurred commonly on unsprayed apple in 1976 and 1977 in the Vancouver District. Except for 13 specimens from the type series (in CNCI), of which six examples were collected in 1978, no other specimens seem to exist to support their contention that P. e/maella was in fact the infesting species throughout the Vancouver District. No voucher mate- rial is deposited at Simon Fraser University in Burnaby (where Doganlar did his work), and no additional specimens could be found in the CNCI. DLW reared four specimens from Cra- taegus sp. near Wilsonville, Clackamas Co., Oregon, in June 1982 (UCON). In addition, we have examined photographs of genitalia of a male from Concrete, Skagit Co., Wash- ington, reared from cultivated apple in July 1986 (DESC). We examined the eight specimens from Monroe, Oregon, reared from apple foliage in July and August 1954 by Jones and Goe- den (OSUO), referred to as P. sorbi by Pot- tinger and LeRoux (1971: 35). We exam- ined a further 18 specimens from near Dal- las, Polk Co., Oregon, reared from apple foliage in August 1970 and July 1971 by S.C. Jones (OSUO). Reports of P. elmaella from western apple orchards (e.g. Weires and Forshey 1978, Orphart 1982, Hoyt 1983, Barrett and Jorgensen 1986, Barrett and Brunner 1990, Jones 1991) likely refer to misidentifications of P. mespilella. a 619 Biology.—Specimens have been reared from domestic apple cultivars, Malus spp. (CNCI, DESC, OSUO), and hawthorn, Cra- taegus sp. (UCON). The relative scarcity of this species from recent apple collections in Oregon and Washington may indicate that it is being displaced by P. blandcardella and P. mespilella. Doganlar and Mutuura (1980) noted that the species was triple brooded in the Vancouver District. The type material (CNCI) bears dates of March, late June, and September. Type material examined.—HOLOTY PE 6 in CNCI, labelled: [1] “Burnaby BC/ 3.1.1978/ Doganlar’’ [handwritten]; [2] **‘Malus/ communis”’ [handwritten]; [3] ““HOLOTYPE/ Phyllonorycter/ elmaella Do. and Mut./ CNC No.” [red, part printed, part handwritten]; [4] “genitalia on slide MIC 2094 3°. The specimen is glued on a point, unspread. Type locality: Burnaby, Vancouver District, British Columbia. PARATYPES: 3 6, 8 2 in CNCI. AL- LOTYPE g, data as holotype except dated 19.VI.1978; genitalia on slide JFL 1091). Phyllonorycter mespilella (Hiibner) Figs. 2, 12, 30-37 Tinea mespilella Hiibner, [1805]: series 8, pl, 39: fig. 272. Lithocolletis mespilella (Hiibner): Rebel, 1901: 213. Povolny, 1949. Pottinger and LeRoux, 1971: 35. Lithocolletes [sic] blancardella (Hubner): sensu Dyar, 1902: 551. Phyllonorycter mespilella (Hiibner): Brad- ley, 1972: 10. Emmet etal, (yep. om Lithocolletis pomifoliella Zeller, 1839: 218. Lithocolletis pyrivorella Bankes, 1899: 252. Diagnosis.—In male genitalia (Fig. 2) P. mespilella can be recognized by the large, dilated right costal process with a curved apical spine that is shorter than the process 3.1.1978, holotype male (CNCI); 28, e/maella, Burnaby, British Columbia, ex Malus, 3.1.1978, holotype male (CNCD); 28, e/maella, Burnaby, British Columbia, ex Malus, 19.vi.1978, allotype female (CNCD); 29, elmaella, Wilsonville, Oregon, ex Crataegus, 29.vi.1982 (UCON). 620 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON itself and the asymmetrical valvae with the ight one being broader than the left one. rhe left costal process is short and stubby, similar to that of P. blancardella. In female genitalia (Fig. 12), P. mespilella is easily distinguished from all other species of the group by the presence of an oval, ringlike sclerotization around the ostium bursae. The sclerotization of the sterigma is wider than in other species (except P. blan- cardella) and, with the apodemes, resembles an X. Forewing length: spring generation, 3.3- 4.5 mm (n = 30); summer generation, 2.7- 3.8 mm (n = 30). Antenna uniformly fus- cous, occasionally with darkened terminal articles. Mesothoracic scutellum with only small patch of white scales, 1 or 2 rows wide. Forewing (Figs. 30-37): ground fiery to red- dish coppery orange, summer brood indi- viduals often paler orange; basal area of dor- sal margin with narrow line of white scales, rarely more than 3 scale rows in width; first costal strigula usually twice as long as broad, axis usually intersecting tornus or outer margin, rarely confluent with first dorsal strigula (Fig. 37); fourth costal strigula often subequal to third, rarely confluent with third costal strigula (Fig. 32), or absent (Fig. 35); white basal streak subequal to width of or- ange-scaled area between basal streak and costa, sometimes confluent with first dorsal strigula (Figs. 32, 35-36); outer row of la- melliform fringe scales with distal %4 to 2 darkened, forming narrow iridescent blue- black band between outermost strigulae. Hindleg: tibia often smoky distad; first and second tarsomeres with dark apical bands. Distribution. — Phyllonorycter mespilella is currently known only in the West from California north to British Columbia and eastward into Utah and New Mexico. It is presumably introduced from Europe and was first reported in North America by Pot- tinger and LeRoux (1971: 35) from Cali- fornia. Our verification of vouchers has confirmed that Pottinger and LeRoux’s (1971) identification of P. mespilella was correct. The species was omitted from the most recent checklist of North American Lepidoptera (Davis 1983). A series of eight specimens in the CNCI collected at Peta- luma, Sonoma Co., California by E.C. John- ston in 1936, 1937, and 1938 also belongs to P. mespilella. Hence the species has been present in western North America for at least 50 years. Material collected from apple leaves from several locations in California, Oregon, Washington, Utah (CISC, UCON), New Mexico (DESC), and British Columbia (CNCI, UCON) reveal that P. mespilella is widespread in western North America. The species referred to as undescribed P. near elmaella by Varela and Welter (1992) is P. mespilella (vouchers examined, UCON). With the exception of the original descrip- tion, all reports of P. e/maella from western North America (e.g. Weires and Forshey 1978, Orphart 1982, Hoyt 1983, Barrett and Jorgensen 1986, Barrett and Brunner 1990, Jones 1991) likely represent P. mespilella and not P. e/maella. Biology.—Phyllonorycter mespilella has the broadest host range of any of the North American apple-feeding Phyllonorycter. In the West, it commonly mines Malus (var- ious cultivars as well as crab apples), Pru- nus, Pyrus, Cydonia, Crataegus, and Coto- neaster (Borden et al. 1953, DLW unpub- lished data). Povolny (1949) listed these same host genera for European populations of P. mespilella, as well as Amelanchier and Mespilus. In Great Britain, Prunus and Sor- bus species are the primary hosts, although a variety of other Rosaceae are occasionally used (Emmet et al. 1985). Under outbreak densities females will oviposit on apricot but few larvae reach maturity (L. Varela, in litt. 1993). Up to five generations are pro- duced in orchards of the Central Valley of California; three to four in coastal areas of California; three in the Okanagan Valley of southern British Columbia (Cossentine and Jensen 1992, Varela and Welter 1992). Var- ela and Welter (1992) used (E10)-Dodecen- yl acetate to monitor the phenology of P. VOLUME 97, NUMBER 3 mespilella in California. We examined spec- imens reared from domestic Prunus (cherry) cultivars from orchards in California and Oregon, which are true P. mespilella. How- ever, there appears to be one or more un- described species of Phyllonorycter on na- tive Prunus species in the West (Deschka, in litt., DLW, unpublished data). NOMEN DUBIUM Phyllonorycter deceptusella (Chambers) Fig: 25 Lithocolletis deceptusella Chambers, 1879: os Phyllonorycter deceptusella (Chambers): Davis, 1983: 10. This species was described from a single specimen, recorded as follows (Chambers 1879: 73): “Among my captured specimens of L. crataegella Clem. I find a specimen of this species which at the time of its capture I regarded only as a variety, but which a more attentive examination convinces me is a distinct though allied species. The ab- domen and two hinder pair of legs are want- ing, though otherwise the insect is in good condition, and though there is but a single specimen, I described it for the purpose of discriminating it from crataegella.* Lithocolletis deceptusella Chambers and L. crataegella Clemens were both synony- mized by Walsingham (1891: 328) with the European pomifoliella Zeller, now a junior synonym of mespilella (Hiibner), which at that time was confused with blancardella (F.). Busck (1903: 190), upon examining the type of crataegella, agreed with Walsing- ham. Of course at that time none of these authors examined genitalia, the study of which had not yet begun. Phyllonorycter de- ceptusella is listed as a valid species by Da- vis (1983: 10). Hagen (1884: 152) indicated that the type of this taxon was not recog- nizable. Braun (1908b: 298) indicated that based on color pattern, P. deceptusella was distinct and “not closely related to cratae- gella or the other apple feeding species.” 621 There is no indication in Chambers’ (1879) description that the species feeds on apple, rather Chambers remarked that he suspect- ed “‘that it feeds on Oak’’ (Chambers 1879: 74). Thus the statement in Pottinger and LeRoux (1971: 5) to the effect that Braun (1908b) listed deceptusella as feeding on ap- ple is in error. Although variants of P. crataegella oc- casionally have the basal and first dorsal strigula confluent, there are several other aspects of the forewing maculation that are inconsistent with those of crataegella. The first costal strigula is broader than long, and it is not obliquely angled to the outer margin as in typical crataegella.. Nor are the white stigulae margined with black scales as they should be. This later feature and the basal displacement of the first dorsal strigula would suggest affinity to eastern cherry- feeding Phyllonorycter species. Type material examined.—HOLOTY PE 2 in MCZC, labelled: [1] ““Type/1323” [red with top portion white, number handwrit- ten]; [2] ““Chambers/Kentucky”’ [partly handwritten, partly printed]; [3] “‘Lithocol- letis/deceptusella Chamb” [handwritten]. The specimen is double mounted, partly spread, with the right antenna, all but two legs, and the abdomen missing. WRONGLY RECORDED TAXA Phyllonorycter sorbi (Frey). Phyllonoryc- ter sorbi was reported from Oregon by Pot- tinger and LeRoux (1971: 35). Our verifi- cation of original vouchers (OSUO) indi- cates that their records were misidentifica- tions of P. elmaella. We have seen no specimens of P. sorbi from North America. Phyllonorycter scudderella (Frey and Boll). We have been unable to trace voucher spec- imens to check the single P. scudderella rec- ord on apple from Québec (LeRoux 1960: 107). We presume that it is based on a mis- identification or a labelling error as the spe- cies is a willow feeder (Braun 1908b: 296, Ely 1917, Needham et al. 1928). 22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 30-37. Forewings of apple-feeding Phyllonorycter mespilella. 30, San Francisco, California, ex Coto- neaster, 20.1.1986 (UCON); 31, Hood River, Oregon, ex apple, 27.ii-14.iv.1990 (UCON),; 32, Summerland, British Columbia, ex cultivated apple, 18-19.iii.1992 (CNCI); 33, Naramata, British Columbia, ex cultivated apple, 21—22.i11.1992 (CNCI); 34, Naramata, British Columbia, ex cultivated apple, 12.vi.1992 (CNCI); 35, Naramata, British Columbia, ex cultivated apple, 20-22.vi.1992 (CNCI); 36, Naramata, British Columbia, ex cultivated apple, 23.vi.1992 (CNCI); 37, Summerland, British Columbia, 5—21.viii.1992 (UCON). VOLUME 97, NUMBER 3 ACKNOWLEDGMENTS We are indebted to Gerfried Deschka (Steyr, Austria) for loaning specimens, mak- ing available his unpublished data on North American apple-feeding Phyllonorycter, and for generously sharing his knowledge of this group. Heather McBrien, Peter Shearer, Jim Troubridge, Lucia Varela, Charles Vincent, Steve Welter, and Jim Whitfield kindly sup- plied apple leaves with active mines— most of the adult photos derive from this mate- rial. Steven Passoa was instrumental in helping us obtain USDA state records, to which Gary Clement, Wheeler Foshee, Clyde Gorsuch, and Terry Harrison also contrib- uted. John Borden, Jim Corrigan, Joan Cos- sentine, Don Davis, Bob Duncan, John Heraty, Jack Lattin, E. J. LeRoux, Scott Miller, Klaus Sattler, and Felix Sperling all contributed information, records, or sug- gestions. Bernard Landry helped with spec- imen preparation and dissection of genita- lia. Julie Henry drew Fig. 13 and supplied the photograph of Vichai Malikul’s color painting of Phyllonorycter deceptusella. Dave Moorehouse inked drawings of geni- talia and prepared the plates. Bill Lukey as- sisted with photography of adult moths. This research was supported by Agriculture Can- ada Research Branch (JFL) and by NSF grant BSR-90-07671 to DLW. LITERATURE CITED Bankes, E.R. 1899. Lithocolletis concomitella, sp. n., and its nearest allies. The Entomologist’s Monthly Magazine 35: 241-255, 284-288. Barrett, B. A. and J. F. Brunner. 1990. Temporal distribution of Phyllonorycter elmaella (Lepidop- tera: Gracillariidae) and its major parasitoid, Pni- galio flavipes (Hymenoptera: Eulophidae), in Washington apple orchards. Environmental En- tomology 19: 362-369. Barrett, B. A. and C. D. Jorgensen. 1986. Parasitoids of the western tentiform leafminer, Phyllonorycter elmaella (Lepidoptera: Gracillariidae), in Utah ap- ple orchards. Environmental Entomology 15: 635- 641. Beckham, C. M., W. S. Hough, and C. H. Hill. 1950. Biology and control of the spotted tentiform leaf miner on apple trees. Virginia Agricultural Ex- 623 periment Station Technical Bulletin no. 114: 1- 19. Borden, A. D., W. H. Lange, and H. F. Madsen. 1953. A spotted tentiform leafminer outbreak in Cali- fornia orchards. Journal of Economic Entomology 46: 519. Bradley, J.D. 1972. [Microlepidoptera], pp. 1-40. Jn Kloet, G. S. and W. D. Hincks, eds., A Check List of British Insects. Second Edition (revised). Part 2: Lepidoptera. Royal Entomological Society, London. 153 pp. Braun, A. F. 1908a. New species of Lithocolletis. En- tomological News 19: 99-107. . 1908b. Revision of the North American spe- cies of the genus Lithocolletis Hiibner. Transac- tions of the American Entomological Society 34: 269-357 + 5 pls. 1939. Notes and new species: Gracillariidae (Microlepidoptera). Busck, A. 1903. Notes of Brackenridge Clemens’ types of Tineina. Proceedings of the Entomological So- ciety of Washington 5: 181-220. Chambers, V. T. 1879. Micro-Lepidoptera. The Ca- nadian Entomologist 11: 72-75. Clemens, B. 1859. Contributions to American Lep- idopterology.—No. 2. Proceedings of the Acade- my of Natural Sciences of Philadelphia 1859: 317- 328. Cossentine, J. E. and L. B. Jensen. 1992. Establish- ment of Phyllonorycter mespilella (Hiibner) (Lep- idoptera: Gracillariidae) and its parasitoid, Pni- galio flavipes (Hymenoptera: Eulophidae), in fruit orchards in the Okanagan and Similkameen Val- leys of British Columbia. Journal of the Ento- mological Society of British Columbia 89: 18-24. Davis, D.R. 1983. Gracillariidae, pp. 9-11. 7n Hodg- es, R. W. et al., eds., Check List of the Lepidoptera of American North of Mexico. E. W. Classey and The Wedge Entomological Research Foundation, London. xxiv + 284 pp. Doganlar, M. and B. P. Beirne. 1980. Parasites of Phyllonorycter elmaella (Lepidoptera: Gracilari- idae [sic]) on apple in the Vancouver District, Brit- ish Columbia. The Canadian Entomologist 1 12: 314. Doganlar, M. and A. Mutuura. 1980. A new species of Phyllonorycter Hbn. (= Lithocolletis Hbn.) (Lepidoptera: Gracilariidae [sic]) from Western North America. The Canadian Entomologist 1 12: 311-314. Dyar, H. G. 1902 [1903]. A list of North American Lepidoptera and key to the literature of this order of insects. Bulletin of the U. S. National Museum no. 52: 1-723. Ely,C.R. 1917[1918]. A revision of the North Amer- ican Gracillariidae from the standpoint of vena- tion. Proceedings of the Entomological Society of Washington 19: 29-77. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Emmet, A. M., I. A. Watkinson and M. R. Wilson. 1985. Gracillariidae, pp. 244-368. Jn Heath, J. and A. M. Emmet, eds., The Moths and Butterflies of Great Britain and Ireland. Volume 2. Cossidae- Heliodinidae. Harley Books, Colchester (Essex). 460 pp. Fabricius, J.C. 1781. Exhibentes eorum differentias specificas, synonyma auctorum, loca natalia, meta morphosin adiectis observationibus, description- ibus. Species Insectorum 2: 1-517. Gries, G., H. L. McBrien, R. Gries, J. Li, G. J. R. Judd, K.N. Slessor, J. H. Borden, R. F. Smith, M. Chris- tie, J. T. Troubridge, P. D. C. Wimalaratne, and E. W. Underhill. 1993. (E4, E10)-dodecadienyl acetate: novel sex pheromone component of ten- tiform leafminer, Phyllonorycter mespilella (Hiib- ner) (Lepidoptera: Gracillariidae). Journal of Chemical Ecology 19: 1789-1798. Hagen, H. A. 1884. The types of Tineina in the col- lection of the museum in Cambridge, Mass. Pap- ilio 4: 151-154. Hodges, R. W. 1986. Gelechioidea: Gelechiidae (in part). In Dominick, R. B., et al., eds., The Moths of America North of Mexico, fascicle 7.1. The Wedge Entomological Research Foundation, Washington. 195 pp. + xiii, 38 pls. Hoyt, S. 1983. Biology and control of the western tentiform leafminer. Proceedings of the Washing- ton State Horticultural Association 79: 115-118. Hiibner, J. [1805] (1796-[1838]). Sammlung euro- paischer Schmetterlinge. 8. Augsburg. 78 pp., 71 pls. Jones, V. P. 1991. Binomial sampling plans for ten- tiform leafminer (Lepidoptera: Gracillariidae) on apple in Utah. Journal of Economic Entomology 84: 484-488. Kumata, T. 1963. Taxonomic studies on the Litho- colletinae of Japan (Lepidoptera: Gracillariidae) Part II. Insecta Matsumurana 26: 1-48. Laing, J. E., J. M. Heraty and J. E. Corrigan. 1986. Summer diapause in Phyllonorycter blancardella (Fabr.) (Lepidoptera: Gracillariidae) in Ontario. The Canadian Entomologist 118: 17-28. Landry, J.-F. and B. Landry. 1994. A technique for setting and mounting microlepidoptera. Journal of the Lepidopterists’ Society 48: 205-227. LeRoux,E.J. 1960. Effects of “modified” and ““com- mercial’ spray programs on the fauna of apple orchards in Quebec. Annales de la Société ento- mologique du Québec 6: 87-121. Maier, C. T. 1981. Seasonal occurrence, abundance, and leaf damage of the apple blotch leafminer, Phyllonorycter crataegella, in Connecticut apple orchards. Environmental Entomology 10: 645-649. . 1983. Relative abundance of the spotted ten- tiform leafminer, Phyllonorycter blancardella (F.), and the apple blotch leafminer, Phyllonorycter cra- taegella (Clemens) (Lepidoptera: Gracillariidae), on sprayed and unsprayed apple trees in Con- necticut. Annals of the Entomological Society of America 75: 992-995. 1984. Seasonal development and flight ac- tivity of the spotted tentiform leafminer, Phyllon- orycter blancardella (Lepidoptera: Gracillariidae) in Connecticut. The Canadian Entomologist 116: 435-441. . 1985. Rosaceous hosts of Phyllonorycter spe- cies (Lepidoptera: Gracillariidae) in New England. Annals of the Entomological Society of America 78: 826-830. Maier, C. T. and D. R. Davis. 1989. Southern New England host and distributional records of litho- colletine Gracillariidae (Lepidoptera) with com- parison of host specificity in temperate regions. Miscellaneous Publications of the Entomological Society of America no. 70: 1-23. McDunnough, J. H. 1939. Check list of the Lepi- doptera of Canada and the United States of Amer- ica. Part II. Microlepidoptera. Memoirs of the Southern California Academy of Sciences 2: 1I- 7A Needham, J. G., S. W. Frost, and B. H. Tothill. 1928. Leaf-Mining Insects. The Williams and Wilkins Company, Baltimore. 351 pp. Orphart, D. R. 1982. Spotted tentiform leafminer evidences spreading in Pacific Northwest apple or- chards. The Goodfruit Grower 33 (5): 14. Pottinger, R. P. and E. J. LeRoux. 1971. The biology and dynamics of Lithocolletis blancardella (Lepi- doptera: Gracillariidae) on apple in Quebec. Mem- oirs of the Entomological Society of Canada 77: 1-437. Povolny, D. 1949. Podkopénkovi moli rodu Litho- colletis Hb. na ovocnych stromech. Acta univer- sitatis agriculturae et silviculturae, Brno, RCS C45. 57 pp. + 6 pls. Pree, D. J., D. B. Marshall, and D. E. Archibald. 1986. Resistance to pyrethroid insecticides in the spotted tentiform leafminer, Phyllonorycter blancardella (Lepidoptera: Gracillariidae), in southern Ontario. Journal of Economic Entomology 79: 318-322. Rebel, H. 1901. Famil. Pyralidae-Micropterygidae. Part 2 in Staudinger, O. and H. Rebel, eds., Cat- alog der Lepidopteren des Palaearctischen Fau- nengebietes. R. Friedlander & Sohn, Berlin. 268 pp. Robinson, G. S. 1976. The preparation of slides of Lepidoptera genitalia with special reference to the Microlepidoptera. Entomologist’s Gazette 27: 127- 132: Roelofs, W. L., W. H. Reissig and R. W. Weires. 1977. Sex attractant for the spotted tentiform leafminer moth Lithocolletis blancardella. Environmental Entomology 6: 373-374. VOLUME 97, NUMBER 3 Stultz, H. T. 1964. Lithocolletis blancardella Fabri- cius (Lepidoptera: Gracilariidae [sic]) on apple in eastern North America with notes on other species occurring on Rosaceae. The Canadian Entomol- ogist 96: 1442-1449. Trimble, R. M. 1984. Effect of carbohydrate on the fecundity of the spotted tentiform leafminer, Phy/- lonorycter blancardella (Lepidoptera: Gracillari- idae), in sprayed and unsprayed apple orchards in Wisconsin. Environmental Entomology 14: 459- 463. Varela, L. G. and S. C. Welter. 1992. Parasitoids of the leafminer, Phyllonorycter nr. elmaella (Lepi- doptera: Gracillariidae), on apple in California: Abundance, impact on leafminer, and insecticide- induced mortality. Biological Control 2: 124-130. Walsingham, T.G. 1882. Notes on Tineidae of North America. Transactions of the American Ento- mological Society 10: 165-204. 625 . 1891. Steps towards a revision of Chambers’ index, with notes and descriptions of new species. Insect Life 3: 325-329. Weires, R. W. and C. G. Forshey. 1978. Leafminer stages a comeback. American Fruit Grower 98(3): 26, 50. Weires, R. W., D. R. Davis, J. R. Leeper, and W. H. Reissig. 1980. Distribution and parasitism of gracillariid leafminers on apple in the Northeast. Annals of the Entomological Society of America 73: 541-546. Wilson, H. F. 1915. An apple leaf miner, Phyllono- rycter (Lithocolletis) crataegella, Clems. (Second Biennial Crop Pest and Horticultural Report, 1912- 1914). Annual Report of Oregon Agricultural Ex- periment Station 1915: 119-120. Zeller, P. C. 1839. Versuch einer naturgemassen Eintheilung der Schaben. Isis von Oken 3: 167- 220. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 626-633 NOTES ABOUT VESCINAE, A KEY TO THE WORLD GENERA, AND DESCRIPTION OF TWO NEW CHOPARDITA (HETEROPTERA: REDUVIIDAE) J. MALDONADO CAPRILES Department of Crop Protection, University of Puerto Rico at Mayag ez, Mayaguez, Puerto Rico 00681; mailing address: Urb. Aponte 6 I 1, Cayey, Puerto Rico 00736. Abstract. —New characters to separate Vescinae from all other Reduviidae subfamilies are discussed, a key to the World genera is presented, and the new African species Cho- pardita granulosa and C. mimetica are described. Key Words: species, Reduviidae China and Usinger (1948) remarked that the validity and coherance of Vescinae could be questioned. Their arguments were: a) the genitalia of the American Vescia Stal and the African Chopardita Villiers show close relationship whereas those of Pessoaia Cos- ta Lima are quite different; b) Pessoaia, Mi- crovescia Wygodzinsky and Mirambulus Breddin run to Peiratinae, c) all genera lack ocelli except Mirambulus, and d) Vescia and Chopardita lack claspers. They placed Mir- ambulus and Megavescia in two different subfamilies, unaware that these genera are synonyms. Wygodzinsky (1950) refuted the above points and stated that the “apical prolongation of the fore tibia beyond the insertion of the tarsus serves to distinguish all Vescinae.” Chopardita, an African genus, is the only vescine outside the Neotropics. It is closely related to Vescia, an affinity that could be a result of fragmentation of Gondwanaland. Measurements are in mm. The holotype of Chopardita mimetica is in the Paris Mu- seum, France, and of C. granulosa in the Natural History Museum, London, En- gland. Vescinae, diagnosis, key to genera, Chopardita granulosa, C. mimetica new SUBFAMILY CHARACTERS IN VESCINAE All genera have a flattened, glabrous area, demarcated or not by a carina, on the apical frontal face of the fore tibia. The carina may be triangular as in Vescia or oblong-ovate as in Chopardita. This area may include the insertion of the tarsi, as in Vescia, or the insertion may be apicad to it as in Chopar- dita. This area may be related to the flexed position of the tarsi, but not for their re- ception, as the tarsi are longer (Fig. 8). It is in the same position as the furrow for the reception of the tarsi in some apiomerine genera. Some apiomerine genera lack this furrow and instead have a demarcation quite similar to that of Chopardita. In the apiom- erines this sulcus or flat area occurs also in the middle tibia but in the vescines the flat- tened area is limited to the protibia. In all vescine genera the apex of the fore tibia is extended and curved backward past the insertion of the tarsi (Figs. 1, 8). Their tarsi are filiform, between '2 (fore) and 13 (middle and hind) the length of the corre- sponding tibia (Figs. 18, 19). The first seg- ment is the shortest, the last two subequal VOLUME 97, NUMBER 3 627 Figs. 1-9. 1. Mirambulus sp., fore leg. 2 Vescia sp., pronotum and mesopleura (mpl), lateral view. Chopardita mimetica n. sp., female, 3. pronotum, dorsal view, 4. profemur, dorsal view, 5. head, dorsal view, 6. posterior margin of pronotum and abdomen, dorsal view. Chopardita mira, female, 7. head, dorsal view, 8. apex of fore tibia, frontolateral view, 9. posterior margin and abdomen, dorsal view. or the third slightly longer than the second and twice as long as the first. The claws are long, tapering, unnotched basally. A broad triangular gena is common to all genera (Fig. 13). Its blunt apex barely reach- es the margin of the clypeus. Its upper mar- gin ends below the antenniferous tubercle and the lower above the upper margin of the buccula. Wygodzinsky gives a partial drawing of the gena of Microvescia (1943: 221, Fig. 87). The pterostigma of the Heteroptera is a slightly chitinized, more or less expanded area, at the external end of the corium. It is 628 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON limited externally by C or Sc and internally by R. Both converge to form its apex. In Vescinae it is broader and longer than in all other subfamilies. In this subfamily it starts just past the level of the apex of the scutel- lum, notably widens, and ends about level with the apex of the inner membranal cell. Along its apical half R runs very close to the basal veins of the membranal cells (Figs. 6, 9). In Vescinae, veins C and Sc are not visible from above on the basal third or fourth of the fore wings. The area they occupy 1s bent downward, vertical to the wing blade. Vein R consequently is the first lateral vein in this area. Vein R is much thicker than the other veins. Some species of Evagoras Bur- meister, a typical harpactorine, have a thick R parallel to the normally located C and Sc, and the latter form a narrow and short pter- ostigma. The fore wings of the vescines nar- row to about level of the apex of the scu- tellum, then broaden in an oval fashion. The connexivum is exposed starting from the level of the apex of the scutellum. The constriction is less marked in Pessoaia. On- cerotrachelus Stal and other saicines and the reduviine Microlestria Stal have a thick R that runs parallel to the costal margin and a relatively wide, but shorter pterostigma. The partially marginal, thick R vein and the long and wide pterostigma are characteristic of the vescine. The subcircular, larger, anterior lobe and a shorter, apron-shaped, posterior lobe of the pronotum is similar in all vescine genera (Fig. 3). The anterior lobe is inflated in all genera except, Chopardita, but its pronotum has the typical vescine outline. The upper surface of the anterior lobe of the other gen- era is above the surface of the posterior lobe (Fig. 2). The conical spinules of the anterior lobe, usually two anterior and four poste- rior, and four anteapicals on the posterior lobe are in the same relative position in all genera (Fig. 3). In many reduviines a some- what similarly shaped pronotum occurs, but the anterior lobe is narrower, never inflated and the posterior lobe is spined differently. The inflated anterior lobe of the harpacto- rine Notocyrtus Burmeister is quite different and has no spinules. Longitudinal, short crenulations occur between both lobes of the pronotum in all vescines. The basal half of fore femora 1s incrassate in all genera except Pessoaia. In the latter the tibiae tapers from a slightly broad base to apex. The basal half of the internal face of the fore tibia is flattened in Vescia, Mir- ambulus, and Chopardita only (Fig. 4), a character not present in other reduviids. Wygodzinsky noticed this character in Mir- ambulus. The middle and hind femora are straight, cylindrical in all vescine genera. The armature of the anterior femora and tibiae of all vescines consists of teeth and round or spiniform tubercles, never spines (Figs. 1, 8). Middle and posterior femora are unarmed. The scutellar spine is preap- ical in all genera (Figs. 11, 16). Wygodzinsky (1943) noticed that the microspines of the coxae and trochanters of the fore legs are good specific characters. The barrel-shaped fore coxae are about a third as long as the fore femora and not flattened laterally as they are in Peiratine. Elongate coxae is a relatively uncommon reduviid character. Emesines, bactrodines, and saicines also have long coxae but their overall shape is quite different from that of the vescines. Vescine species are black, dark ferrugin- eous or dark or pale brown. The coria may have an extensive pale area or be slightly ornamented with yellowish, ivory or whit- ish. The membrane is grayish or ferrugin- eous, may have more or less extensive paler areas, and veins are brown usually. The con- nexivum can be either unicolorous, usually a shade of brown, or the segments may be brownish at middle and paler at sutures. The sutures separating the connexival seg- ments are usually very tenuous (Figs. 12, LZ): VOLUME 97, NUMBER 3 10 629 Figs. 10-19. Chopardita villiersi, 10. pronotum, dorsal view, 11. scutellum, lateral view, 12. posterior margin of pronotum and abdomen, dorsal view. Chopardita granulosa n. sp., female holotype, 13. head, lateral view, 14. genital segments, lateral view, 15. genital segments, caudal view, 16. scutellum, lateral view, 17. abdomen, dorsal view. Vescia sp., 18. tarsi, hind leg, 19. tarsi, middle leg. KEY TO THE GENERA IN VESCINAE 1. Anteocular space shorter than postocular; first antennal segment longer than anteocular space (Rigs) ere nice erate eat oe rca: — Anteocular space longer than postocular; first antennal segment shorter than anteocular space tO Spine between antenniferous tubercles present; WiIthOUtOCEICN ici hace AA Heer cere 3 Spine between antenniferous tubercles absent; ocelli present ......../ Mirambulus Breddin 1901 Anterior prontal lobe not inflated, its surface level with that of posterior lobe, with 2 short, vertical anterior and four preapical spinules; longitudinal sulcus of anterior lobe begins at 63( PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON midlength with an oval pit, reaching transverse constriction between lobes — Anterior pronotal lobe inflated subglobularly, surface above level of posterior lobe (Fig. 2), with or without anterior short spinules; lon- gitudinal sulcus from behind collar to level of posterior spinules of same lobe, not reaching transverse constriction between lobes ....... PER ees SSE Eos acidic aoe Vescia Stal 1865 4. Head almost and pronotum glabrous, some species with few scattered setae, anterior lobe of pronotum with short spinules and fine cor- rugations; humeral angles spined or not; apex of prosternum reaching mesosternum PE NE eet, 2 io 3 A ated Pessoaia Costa Lima 1940 — Head and pronotum densely pilose; anterior lobe of pronotum without spinules and cor- rugations; humeral angles spined; apex of pro- sternum not reaching mesosternum ec Re oh a Oe Microvescia Wygodzinsky 1943 THE GENERA AND SPECIES OF VESCINAE For synonyms of species see Wygodzin- sky (1949) or Maldonado (1990). Miller (1951: 465) described Eremovescia and included it in Vescinae. Villiers (1954: 221) synonymized it with Pasira Stal, a Re- duviinae. This genus occurs from Maure- tania to the Canary Islands. Chopardita Villiers Chopardita Villiers 1944: 79. Type: Cho- pardita mira Villiers 1944: 80. Ivory Coast. Chopardita mira Villiers Villiers 1944: 80. Occidental Africa, Zaire, Oubanghi-Chari, Sudan. Figs. 7-9. Chopardita villiersi China and Usinger China and Usinger 1943: 509. Sudan. Figs 10-12. Chopardita mimetica Maldonado, NEw SPECIES Figs. 3-6 Female.— Brown: head, first segment of rostrum, anterior femora and tibiae, pos- terior lobe of pronotum, abdomen ventral- ly, connexivum above and below. Dark brown: anterior lobe of pronotum, scutel- lum, tubercles of fore femur and tibia. Pale brown: antenna, femora ventrally, middle and hind legs, second and third segments of rostrum. Hemelytra: basal angle and pter- ostigma blackish brown; inverted trapezoi- dal, dirty-yellow area from level of apex of scutellum, margined laterally and apically to base of inner cell of membrane by black R (Fig. 6); membrane gray, cells same color as pale area of corium, veins dark brown. Head: \ength ofanterior lobe 0.40, length of posterior lobe 0.62, greatest width of pos- terior lobe 0.59, depth of posterior lobe 0.59, interocular space 0.28, interantennal spine reaching one third of first antennal segment. Antennal segments: I. 0.43; II. 1.25; II] and IV missing; glabrous. Thorax: anterior lobe—length 0.75, width 1.00, two anterior vertical spinules and 2+2 subapical trian- gular spinules irregularly spaced; lateral margin not carinate, with six small granules of equal size; posterior lobe: length 0.53, width 1.31, with 2+2 subapical triangular spinules. Legs—anterior femur (Fig. 4): in- crassate basally, narrowing to apex, without thick laterosubapical tubercle, ventral tu- bercles small, in two irregular rows, internal face flattened; length 1.62, greatest thick- ness 0.50; tibia: curved and bent apically, length 1.53, with typical dorsal granules, tarsi missing; middle leg: femur length 1.56; tibia straight, length 1.72; hind leg: femur length 2.06, tibia straight, length 2.00. Scu- tellar spine broken, preapical. Hemelytra reaching apex of abdomen; pterostigma: length 1.28, width 0.37. Connexival sutures obsolete. Abdomen length 2.60, greatest width 1.53. Overall length 4.90. Holotype-female, COTE D’IVOIRE, bords du Volta-rouge, december 1938, L. Chopard collector, in Paris Museum. The holotype is the specimen misidenti- fied as Chopardita mira by Villiers (1944: 80). It was compared by me with a speci- men, correctly identified, from Republic of Tchad, Bas Chari, 13.viii.1963, environs du VOLUME 97, NUMBER 3 Douggia, labeled by Villiers. My drawings of C. mira are from this latter specimen. Villiers did not declare paratypes any of these specimens. This species has the basal third of middle and hind femora, the ros- trum, tibia, and apex of fore femora yellow; the abdomen 1s slightly longer and narrower (2.60:1.53::2.40:1.62), and the width across the eyes greater than across posterior lobe of head than in C. mimetica sp. nov. (0.66: 0.53::0.62:0.59). In the latter species the an- terior pale area of the hemelytra is limited laterally by R, whereas in C. mira the pale area reaches the lateral margins of the wing (Fig. 9). C. mira has a pale area on the mem- brane, caudad of the pterostigma, that is absent in C. mimetica. Choparaita villiersi is pale brown. Differs from C. mira and C. mimetica by having three small spinules (Fig. 10) on the thinly carinate lateral margin of the anterior lobe of the pronotum, and a smooth vertex. The last two species have five or six lateral gran- ules and their vertex is shagreen or smooth. These species are about 5.3 mm long. Chopardita granulosa Maldonado, NEW SPECIES Figs. 13-17 Female.— Black: anterior lobe of prono- tum, scutellum, humeral angles of hemely- tra, pterostygma, mesopleura, and meta- pleura. Brown: antenna, anterior lobe of head. Stramineous: second and third seg- ments of rostrum, connexivum above, fem- ora, scutellar spine. Sort of greenish brown: posterior lobe of pronotum, abdominal ster- na. White: most of discal area of clavus. Gray: membrane, lateroposterior angles of white discal area. Head.—minutely granulose, length 2.19, anteocular space 0.88 (to apex of head); postocular space 1.25, length of interanten- nal spine 0.38, length of eye 0.50; collum poorly defined, length 0.05; width across eyes 1.33, interocular space 0.56, width across widest part of posterior lobe 1.13. Antennae missing, only part of one first seg- 631 ment present. Rostral segments: I. 0.88; II. 0.94; III. 0.50. Pronotum-anterior lobe: globular, longitudinal sulcus beginning at midlength with a small pit, then deep to transverse constriction, margin with 6 or 7 minute granules, collar with angles project- ed laterally; with two vertical anterior and 2+2 posterior spinules; greatest width 1.94, length 1.50; posterior lobe: length 1.06, hu- meral width 2.75 (to base of spinules), finely corrugate and minutely, sparsely granulose; longitudinally crenulate between lobes. Meso- and metapleurae vertically rugose, sparsely granulose. Scutellum long trian- gular, basal width 0.44, length to base of spine 0.38, length to apex of spine 1.31. Abdomen: length to apex of genital segment 1.70, greatest width 2.25. Genital segments as in Figs. 14, 15. Total body length 5.39. Holotype.—female, N. NIGERIA, Azar, 1928-1929, Dr. Ll. Lloyd collector; para- type same collection data, both in NMNH, London. KEY TO THE SPECIES IN CHOPARDITA 1. Lateral margin of anterior lobe of pronotum with three spinules; fore wings as in Fig. 12 ... Ferenc tae ee C. villiersi China and Usinger — Lateral margin of anterior lobe of pronotum with 5-7 aligned small granules; fore wings not ASHI I Se Deh Canin. hs cei eee ee Z . Posterior lobe of head and posterior lobe of pronotum very finely granulose; fore wings as OO) aes II aie nee C. granulosa Maldonado n. sp. — Posterior lobe of head and posterior lobe of pronotum smooth or shagreen; fore wings as in Figs. 6 or 9 3. Femora bicolored; basal third and apex of fem- ora, rostrum yellow; head pale brown; fore wings as in Fig. 9 C. mira Villiers — Femora unicolored, pale brown; first rostral segment pale brown, last two brown; head brown; fore wings as in Fig. 6 Be PRES Sse cuca C. mimetica Maldonado n. sp. tO Microvescia Wygodzinsky Microvescia Wygodzinsky 1943: 206, 220. Type: Microvescia costalimai Wygodzin- sky 1943: 220. Brazil. Monotypic. Mirambulus Breddin rambulus Breddin 1901: 74. Ecuador. Type: Mirambulus niger Breddin 1901: 75. Ecuador. Megavescia Wygodzinsky 1947: 414. Type: Megavescia cazieri Wygodzinsky 1947: 412, 414. Guyana. Synonymized by Wy- godzinsky 1950: 266. The genus includes: M. morio Breddin 1903. Bolivia. M. niger Breddin 1901. Ecuador. Pessoaia Costa Lima Pessoaia Costa Lima 1940: 487. Type: Pes- soaia piratoides Costa Lima 1940: 487. Brazil. Abalos (1945) keyed the species. P. argentina Wygodzinsky 1943. Argen- tina. P. limai Usinger 1942. Costa Rica, Co- lombia. P. maculata Wygodzinsky 1943. Brazil. P. parkoi Wygodzinsky 1943. Brazil. P. piratoides Costa Lima 1940. Brazil. Vescia Stal Vescia Stal 1866: 123. Type: Vescia spicula Stal 1866: 166. Brazil. In Acanthaspidi- nae. Geaya Villiers 1944: 81. Type: Geaya di- latata Villiers 1944: 81. Venezuela. Syn- onymized by China and Usinger (1948: 603). Wygodzinsky (1943: 207) keyed five of the eight species. V. adamanta Brindley 1931. Guyana V. angrensis Seabra & Hathaway 1942. Brazil. V. brachyptera Usinger 1942. Paraguay. V. dilatata (Villiers) 1944. Venezuela. V. minima Fracker & Bruner 1924. Bra- 7Al. V. nostratis Drake and Harris 1945. Bra- zil. V. penningtoni Drake 1943. Paraguay, Brazil. V. spiculata Stal 1866. Brazil. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ACKNOWLEDGMENTS For the loan of specimens for study I am grateful to D. Plout-Sigwalt, National Mu- seum of Natural History, Paris, France, and Mrs. J. Margerison-Knight, Department of Entomology, Natural History Museum, London, Great Britain. LITERATURE CITED Abalos, J. W. 1945. Pessoaia alvaradoi n. sp. Anales del Instituto de Medicina Regional, Tucuman |: 229-237. Breddin, G. 1901. Neue tropische Wanzen und Zir- pen. Societas Entomologica 16: 74-76. . 1903. Uber neue Palaéotropische Reduviinen. Gesellschaft Natur forschender Freunde 3: 111- 129. China, W. E. & R. L. Usinger. 1948. A new species of Chopardita from the Anglo-Egyptian Sudan with notes on the subfamily Vesciinae. Hemiptera Re- duviidae. Annals and Magazine of Natural History (12)1: 598-604. Costa Lima, A. da. 1940. Novo Hemiptero da sub- familia Vesciinae. Revista do Museu Paulista 1: 485-490. Drake, C. J. 1943. A new neotropical vesciid (He- miptera: Reduviidae). Boletin de entomologia ve- nezolana 2: 207-209. Drake, C. J. & H. M. Harris. 1945. Concerning the subfamily “‘Vescinae.”’ Revista Brasileira de Bio- logia 5: 155-156. Fracker, S. B. & S.C. Bruner. 1924. Notes on some Neotropical Reduviidae. Annals of the Entomo- logical Society of America 17: 163-174. Haviland, Maud D. (Mrs. H. H. Brindley). 1931. The Reduviidae of Kartabo, Bartica District, British Guiana. Zoologica 7: 129-154. Maldonado Capriles, J. 1990. Systematic Catalogue of the Reduviidae of the World. Special Publica- tion. Caribbean Journal of Science, University of Puerto Rico at Mayagiiez. i-x, 1-694 pp. Miller, N. C. E. 1951. New Reduviidae in the col- lection of the British Museum (Natural History)— V. Annals and Magazine of Natural History (12)4: 465-480. Seabra, C. A. C. & C. R. Hathaway. 1942. Especies de Vescia del Brasil (Hemiptera: Reduviidae). Me- morias del Instituto Oswaldo Cruz 37: 539-541. Stal, C. 1866. Hemiptera Africana, Vol. 3. Ofversigt Kongliga Akadiemens Foerhandlingar 3: 1-200. Usinger, R. L. 1942. Key to genera and species of Vesciinae with description of two species (Heter- optera, Reduviidae). Revista de Entomologia 13: 290-296. VOLUME 97, NUMBER 3 633 Villiers, A. 1944. Note sur deux Réduviides africano- (Reduviidae, Hemiptera). Revista de Entomologia bresiliens constituant une nouvelle sousfamille. 18: 411-416. Bulletin de la Société Entomologique de France 1949. Elenco sistematico de los reduvi- 49: 79-83. iformes americanos Instituto de Medicina Re- Wygodzinsky, P. 1943. Contribuicao ao conheci- gional, Tucuman. Monografia. 1: 1-102. mento da subfamilia Vesciinae (Hemiptera, Re- 1950. On the genus Mirambulus Breddin, duviidae). Revista Brasileira de Biologia, 3: 203- with general notes on the subfamily Vescinae (Re- 223 duviidae, Hemiptera). Annals and Magazine of 1947. Sobre um novo genero neotropico de Natural History (12)3: 265-268. Vesciinae, com consideracoes sobre a subfamilia PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 634-638 INTRAGUILD PREDATION BETWEEN SYMPATRIC SPECIES OF MANTIDS (MANTODEA: MANTIDAE) MATTHEW D. MORAN Ecology Program, Department of Biology, University of Delaware, Newark, Delaware 19716. Abstract. —I tested the importance of predation versus competition in two congeneric sympatric species of mantids, Tenodera aridifolia sinensis (Saussure) and T. angustipennis (Saussure), during the late portion of the juvenile stage of their life cycle. Tenodera angustipennis abundance was reduced through predation by the larger T. a. sinensis, but no evidence of competition for resources was demonstrated for either species. Tenodera a. sinensis gained more body mass in the presence of 7. angustipennis indicating that the benefit of consuming smaller predators may outweigh the cost of competing for resources. Key Words: Although the main factor limiting the abundance of predators is predicted to be food (Hairston et al. 1960), regulation of population size by food limitation has been difficult to demonstrate among many pred- atory arthropods (Riechert and Cady 1983, Polis and McCormick 1986, Wise 1981, 1993). Among generalist predators, intra- guild predation seems to be a prevalent in- teraction and may be important in regula- tion of abundance (Polis et al. 1989). In the northeastern United States three species of mantids co-occur: Tenodera ar- idifolia sinensis, T. angustipennis, and Man- tis religiosa L. (Rathet and Hurd 1983). As generalist predators, they have the potential to compete for prey, but differences in their life history apparently alleviate this. Ten- odera angustipennis and M. religiosa hatch later than 7. a. sinensis, establishing a size difference that allows them to consume dif- ferent prey items (Hurd 1988) Mantis reli- giosa 1s spatially separated from the two Tenodera congeners by occupying a lower height in the vegetation (Rathet and Hurd 1983). Although these characteristics can Intraguild predation, competition, Tenodera, Mantodea, Mantidae reduce competition for prey, size differences between the species are great enough to pro- mote intraguild predation (Hurd 1988, Hurd and Eisenberg 1990a). Since 7. angustipen- nis and T. a. sinensis occupy the same veg- etational strata, this interaction may be 1m- portant for these two species. This experiment was designed to test whether competition for prey or intraguild predation is a more important factor during the late portion of the juvenile stage (6-7 stadia) for these species and how these in- teractions affect the potential growth of in- dividuals. MATERIALS AND METHODS The study site was an old field on the Experimental Farm of the University of Delaware. It consisted of mixed grasses and forbs with Poa spp. and Solidago spp. being the most common plants. On 2 August, 12 enclosures each measuring | m? were placed in the field in a 6 x 2 array. Each enclosure consisted of a PVC frame which was cov- ered by fine nylon mesh (Bioquip Products, Gardena, CA). The enclosures were quickly VOLUME 97, NUMBER 3 placed on the ground to prevent the escape of resident arthropods. From 3 to 5 August, both species of man- tids (all females) were collected from a near- by field. This field has had a large popula- tion of both species for several years (per- sonal observation). Each captured mantid was weighed (nearest 0.01 g), individually marked with nail polish and randomly as- signed to a treatment group. The treatment groups were as follows: 1) three 7. a. sinen- sis, 2) three T. angustipennis, and 3) three T. sinensis and three T. angustipennis. Each treatment consisting of four replicates was intentionally interspersed within the enclo- sure array. The mantids were introduced into the enclosures on 5 August. Each day following, the enclosures were inspected, and any mantid that had molted was marked again. On 16 August, all cages were sampled by a combination D-vac and hand search. All surviving mantids were weighed to the near- est 0.01 g, and all other arthropods were separated into their respective orders. RESULTS At no time did more than one mantid molt during one 24 hour period. Therefore, I was able to track every surviving individ- ual and recorded both an initial and final weight for each. The weight of the mantids at the beginning of the experiment did not differ between treatments (7. a. sinensis t5> = 0.41; P = 0.68: T. angustipennis t,. = 1.41; P=0.17) for either species. I therefore assume that any response seen at the end of the experiment was not the result of initial bias. Survivorship of 7. a. sinensis was not sig- nificantly different (t, = 0.52; P = 0.62) be- tween the two treatments (Table 1) while 7. angustipennis showed decreased survivor- ship in treatment 3. Since there was only one surviving 7. angustipennis in one rep- licate for treatment 3, it was not possible to do a statistical comparison. However, all replicates in treatment 2 had either two or 635 Table 1. Mean survivorship (+ | SE) of both spe- cies of mantids in their respective treatments. Treat- ment | = T. a. sinensis, Treatment 2 = 7. angustipen- nis, Treatment 3 = TJ. a. sinensis + T. angustipennis. Species Treatment | Treatment 2 Treatment 3 T. sinensis De2> —_ 2.00 (0.25) (0.41) T. angustipennis _ DDS O25 (0.25) (0.25) three surviving 7. angustipennis, while three of the four replicates in treatment 3 had no survIVOTS. The mean weight of individual 7. a. si- nensis was significantly greater in treatment 3 (1.70 ¢ = 0.12) than in. treatment 1 (1.35 g + 0.06) (t,; = 2.61; p = 0.02) (Fig. 1). An analysis of variance on arthropod abun- dance for the three treatments showed that there was no statistical significance between treatments for any order or for total abun- dance of arthropods (Table 2). DISCUSSION At the conclusion of the experiment, there were fewer surviving 7. angustipennis in treatment 3, where 7. a. sinensis was also present, compared to treatment 2, where 7. a. sinensis was absent. The high mortality was most likely due to predation by 7. a. sinensis, as there were no other arthropods large enough to capture 7. angustipennis, and I observed several such predation events during my daily monitoring. However, the removal of 7. angustipennis by T. a. sinen- sis did not occur immediately, as 7. angus- tipennis were observed in all the treatment 3 replicates through the sixth day of the experiment. Therefore, mantid densities were elevated in treatment 3 for a large por- tion of the experiment. The differences in final weight between treatment | and treatment 3 show 7. a. si- nensis accumulated more biomass in the presence of 7. angustipennis. This indicates that the value of consuming 7. angustipen- nis outweighed the potential cost of in- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 25 — oO Mean Weight (+ 1SE) (g) © oO 6) Treatment 1 Treatment 3 Fig. 1. Final mean weight (+1 SE) for 7. a. sinensis in Treatment | (7. a. sinensis only) and Treatment 3 (T. a. sinensis and T. angustipennis). * P < 0.05. creased competition for prey. The amount of biomass acquired during the juvenile stages of the life cycle affects the size a man- tid will reach at adulthood. This determines the maximum weight gain possible which in turn affects the number of eggs oviposited (Eisenberg et al. 1981). Therefore, the 7. a. sinensis individuals in the presence of 7. angustipennis may have been able to pro- duce more eggs. VOLUME 97, NUMBER 3 Table 2. Arthropod abundance (+ SE) and corre- sponding ANOVA analysis. Orders are arranged in de- scending abundance for Treatment | (T1). Tl = 7. a. sinensis, T2 = T. angustipennis, T3 = T. a. sinensis + T. angustipennis. Order Tl m2 T3 F,5 P Araneae 338}(0) NES} 21.0 0.98 0.41 (8.2) (6.5) (5.4) Homoptera D3ES e29eS 48.8 3.47 0.07 (4.3) (4.9) (10.5) Hymenoptera 11.8 6.3 LES es 4e0 si (3.8) (1.3) (2:3) Diptera 7.0 4.8 5), SORT Ol 7 (@55) ees G@ES) (233) Hemiptera 6.0 6.3 PAO SS Messi (077 (3:3) 163) (0:4) Thysanoptera 35 1.0 OR ESS 0227 (2.9) (0.0) (1.0) Coleoptera 0.5 le3 DPS OGOs39 (0.3) (0.6) (1.8) Total S50 One lel Oales 762030 (10.9) (8.0) (27.8) The analysis of the arthropod assemblage showed there was no difference between treatment groups in either total abundance or within any individual order. It would be expected that treatment 3 would have had lower abundance of arthropods, since the density of mantids was double the other treatments. Actually, the trend was for el- evated arthropod density in treatment 3 al- though this was not significant. It has been shown in recent experiments that increasing the density of predators may have little im- pact on the prey density (Hurd and Eisen- berg 1990b, Fagan and Hurd 1991, Wise 1993). This was a short-term experiment and significant depressions of prey in the presence of elevated predator densities may not have had time to occur. A previous ex- periment (Moran and Hurd 1994) indicated that important short term interactions dur- ing elevated predator densities were intra- guild predation and emigration by predators small enough to be potential prey. Increased food limitation became a factor only later in the study. This experiment showed that the major 637 short term interaction between these two species was predation while competition was apparently absent. That 7. a. sinensis dem- onstrated greater weight gain in the presence of 7. angustipennis indicates that top level predators may benefit from other predators being present. The benefit of using other predators as prey may therefore outweigh the cost of interspecific competition within this guild, at least in the short-term. ACKNOWLEDGMENTS I am grateful to T. P. Rooney for assis- tance with the D-vac and L. E. Hurd for reading an earlier draft of the manuscript. LITERATURE CITED Eisenberg, R. M., L. E. Hurd, and J. A. Bartley. 1981. Ecological consequences of food limitation for adult mantids (Tenodera sinensis Saussure). American Midland Naturalist 106: 209-218. Fagan, W. F. and L. E. Hurd. 1991. Direct and in- direct effects of generalist predators on a terrestrial arthropod community. American Midland Nat- uralist 126: 380-384. Hairston, N. G., F. E. Smith, and L. B. Slobodkin. 1960. Community structure, population control, and competition. American Naturalist 94: 421- 425. Hurd, L. E. 1988. Consequences of divergent egg phenology to predation and coexistence in two sympatric, congeneric mantids (Orthoptera: Man- tidae). Oecologia 76: 549-552. Hurd, L. E. and R. H. Eisenberg. 1990a. Arthropod community responses to manipulation of a bi- trophic predator guild. Ecology 76: 2107-2114. 1990b. Experimentally synchronized phe- nology and interspecific competition in mantids. American Midland Naturalist 124: 390-394. Moran, M. D. and L. E. Hurd. 1994. Short-term responses to elevated predator densities: Noncom- petitive intraguild interactions and behavior. Oe- cologia (In press.) Polis, G. A. and S. J. McCormick. 1986. Scorpions, spiders, and solpugids: predation and competition among distantly related taxa. Oecologia 71: 111- 116. Polis, G. A., C. A. Myers, and R. D. Holt. 1989. The ecology and evolution of intraguild predation. An- nual Review of Ecology and Systematics 20: 297— 330. Rathet, I. H. and L. E. Hurd. 1983. Ecological rela- tionships of three co-occurring mantids, Tenodera 638 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sinensis (Saussure), T. angustipennis (Saussure), density manipulations upon females of two orb- and Mantis religiosa (Linnaeus). American Mid- weaving spiders (Araneae: Araneidae). Oecologia land Naturalist 110(2): 240-248. 48: 252-256. Riechert, S. E. and A. B. Cady. 1983. Patterns of Wise,D.H. 1993. Spiders in Ecological Webs. Cam- resource use and tests for competitive release in a bridge University Press, Cambridge, Great Brit- spider community. Ecology 71: 1441-1450. ain. Wise, D. H. 1981. Inter- and intraspecific effects of PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 639-648 NEW GENERA AND SPECIES OF PERUVIAN PHYCITINAE (LEPIDOPTERA: PYRALIDAE) H. H. NEUNZIG Department of Entomology, North Carolina State University, Raleigh, North Carolina 27695-7613. Abstract.—Three new genera of phycitine moths from Peru are described. They are Depeadus n. gen. with D. deiulus n. sp., Montestra n. gen. with M. dentata n. sp., and Eurythmioides n. gen. with E. carasensis n. sp. Also described from Peru is Caviana peruviensis n. sp. Key Words: Moths, phycitines, Peru In 1987, O. Karsholt, of the Zoological Museum of the University of Copenhagen (ZMUC) collected a series of phycitine moths in Peru. He generously offered this material, along with a smaller amount of other Peruvian Phycitinae in the ZMUC, to me for study. Included in the loan were 4 new species, descriptions of which are given here. Most previously described Peruvian phycitines (Heinrich 1956) have been col- lected at low to moderate elevations, mainly along the coast. Two of the species de- scribed herein were obtained at mountain- ous sites of 3000, to over 4000, meters. Depeadus Neunzig, NEw GENUS Gender. — Masculine. Type species. — Depeadus deiulus Neunzig. Antenna of male (Fig. 9) with distinct si- nus at base of shaft; elongate scales clothing distoanterior part of sinus; sensilla tricho- dea (cilia) of shaft of male moderately abun- dant, and near base of shaft about '2 as long as basal diameter of shaft. Front convex, with scales produced anteriorly. Labial pal- pus robust, upturned. Haustellum well de- veloped. Ocellus present. Basal half of costa of forewing of male (Fig. 6) slightly convex; underside of male wing without costal fold. Forewing without raised scales; with eleven veins, R, distant at base from R;,, and R, and upper outer angle of cell; R;,, and R, stalked for slightly over ' their lengths; M, straight; M, and M; stalked for about '4 their lengths; CuA, from lower outer angle of cell; CuA, from well before outer angle of cell. Hindwing (Fig. 6) with eight veins (LA, 2A and 3A considered to be one vein); Sc + R, and Rs fused for about ' their lengths be- yond cell; M, from upper outer angle of cell and its basal '4 fused with Sc + R, and Rs; M, and M, fused for about '2 their lengths; CuA, fused at base with stalk of M, and M,; CuA, from well before angle of cell; cell about '3 length of wing. Male abdominal segment 8 with ventral scale tufts; sclero- tized element associated with tufts U-shaped and with distally swollen medial projection. Male genitalia (Figs. 19, 20) with uncus nar- row, triangular, and weakly sclerotized; gna- thos absent; transtilla with long, setiferous, bladelike, lateroposteriorly directed arms; juxta broadly V-shaped with lateral, seti- ferous protuberances; valva short, strongly knobbed, and with long, slender, costal pro- jection at base, and large, posteriorly pro- jecting, hooked, apically swollen and spined element; sacculus delineated from valva by 40 ( / j a PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Male adults. 1, Depeadus deiulus n. sp., holotype (11.5 mm). 2, Caviana peruviensis n. sp., holotype (6.0 mm). 3, Eurythmioides carasensis n. sp., holotype (5.5 mm). 4, Montestra dentata n. sp., holotype (20.0 mm). (Length of forewing in parentheses.) strongly sclerotized ridge; dorsodistal sur- face of valva with many long, slender scales; linear aggregation of long slender scales also originating at base of valva; aedoeagus ro- bust with apical '4 strongly spined; cornutus a strongly sclerotized blade; vinculum lon- ger than greatest width, anterior margin with concavity. Female genitalia (Fig. 22) with ductus bursae short, sclerotized; dorsal plate behind genital opening with granulate, pocketlike, lateral lobes; corpus bursae elongate, about 22x longer than ductus bursae, membranous, and with posterior '4 to '2 with numerous spines and scobina- tions; signum absent; ductus seminalis at- tached to corpus bursae near junction with ductus bursae. Depeadus shares the following features with Peadus Heinrich: (1) forewing with eleven veins, and hindwing with eight veins; (2) male antenna with sinus at base of shaft; (3) male genitalia with uncus greatly re- duced, gnathos indistinguishable, valva short, sacculus bearing large scale tufts, ae- doeagus spined at apex; (4) female genitalia with ductus bursae sclerotized and much shorter than corpus bursae, and with scler- otized dorsal plate behind genital opening. Differences between the two genera include: (1) M, and M,; of forewing separate in Pead- us, stalked for basal 3 in Depeadus; (2) Sc + R, and Rs of hindwing briefly fused at base in Peadus, fused for 2 their lengths beyond cell in Depeadus; (3) male genitalia without transtilla, and with divided valva in Peadus, with well developed transtilla, and entire valva in Depeadus; (4) female genitalia with corpus bursae with single VOLUME 97, NUMBER 3 Figs. 5-7. thornlike spine (signum) in anterior '2 in Peadus, without signum, but with numer- ous spines in posterior '3—'2 in Depeadus. Depeadus deiulus Neunzig, NEw SPECIES Bigs.al6..9 9.20.22 Type locality.—5 km E. Limbani, 3000 my Dept, Puno; Peru: Male wing venation. 5, Eurythmioides carasensis n. sp. (5.5 mm). 6, Depeadus deiulus n. sp. (11.5 mm). 7, Montestra dentata n. sp. (20.0 mm). (Length of forewing in parentheses.) Diagnosis.—The slender, bladelike, lat- eral arms of the transtilla, and the long cos- tal projection, and large, hooked and spined element, on the valva of the male genitalia are diagnostic. Description.—Length of forewing 11.5 mm. Head with frons and vertex pale brown and white with a few reddish brown and 642 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LA 4 Z Ss >5 =) a= Z CA es: Yeh ass: Ny 2 ess a> gig KEE: : SEEPS Figs. 8-13. Male antennae and labial palpi. 8, urythmioides carasensis n. sp., frontal view of basal segments. 9, Depeadus deiulus n. sp., frontal view of basal segments. 10, Caviana peruviensis n. sp., frontal view of basal segments. 11, Eurythmiodes carasensis Nn. sp., lateral view. 12, Caviana peruviensis n. sp., lateral view. 13, Montestra dentata n. sp., lateral view. (All scale lengths 0.5 mm.) 13 VOLUME 97, NUMBER 3 dark brown scales in male, pale brown and white in female; labial palpus mostly pale brown with some reddish brown and dark brown to black scales in both sexes; max- illary palpus mostly pale brown with dark brown at base. Thorax dorsum and collar mostly pale brown to pale reddish brown (most scales paler at tip) with scattered dark brown, black, red and reddish brown scales. Forewing with ground color pale brown dusted with white, with scattered ochre and dark brown to black scales; antemedial line white to brownish white, moderately dis- tinct in posterior half of wing; patches of black at costa basad of antemedial line and basad and distad of antemedial line in pos- terior half; postmedial line weakly devel- oped, best defined in costal half where patches of black scales establish its proximal and distal borders; patches of, and a few isolated, red or reddish brown, scales, most- ly in distal half; discal spots dark brown to black, moderately distinct. Underside of forewing of male without sex-scaling. Hind- wing pale smoky brown, darker on veins and near costal and outer margins. Male and female genitalia as given under generic de- scription. Distribution.— Known only from south- eastern Peru. Holotype.—¢—5 km E. Limbani, 3000 m, Dept. Puno, Peru, 28.111.1987, O. Karsholt leg., genitalia slide 3450 HHN (ZMUC). Paratype.—2—Same collection data as holotype, genitalia slide 3451 HHN (ZMUC). Caviana Neunzig and Dow, 1993 Caviana peruviensis Neunzig, NEW SPECIES Figs, 2, 012.1415 Type locality.— 15 km N. Caras, Rio Sal- ta Valley, ca. 2000 m, Ancash, Peru. Diagnosis.—C. peruviensis has the fol- lowing combination of features: a shallow, mostly exposed depression on the frons just anterior to the antennae; male genitalia with a distally attenuated uncus and a dorsally directed spur on the valva. 643 Description.— Length of forewing 6.0 mm. Head with frons with shallow concav- ity partially covered anteriorly by dome of posteriorly and mesially projecting black and white scales; surface of concavity with many, white microscales; vertex white, fuscous and black; male antenna (Fig. 10) simple; labial palpus (Fig. 12) upcurved, extending above vertex, mostly fuscous and black, white at base of first and second segments and at tip of second and third segments; maxillary pal- pus simple, fuscous, black and white. Tho- rax dorsum and collar fuscous and black, dusted with white. Forewing mostly fuscous and black, dusted with white; antemedial line distinct, white, extending diagonally from costa distally to posterior margin; postmedial line indistinct, white; small patches of pale reddish brown scales sub- basally, near posterior margin and in medial area adjacent to antemedial line; discal spots not apparent, blending with black and white pattern of wing. Underside of wing of male with elongate costal fold. Hindwing mostly white, fuscous near costal and outer mar- gins. Male abdominal segment 8 simple. Male genitalia (Figs. 14, 15) with uncus strongly tapered and slender distally; gna- thos with distal part a single, short, straplike element; transtilla incomplete; juxta a plate with slender, lateral lobes; valva with sub- apical, dorsoposteriorly directed spur on costa; vinculum slightly longer than greatest width; aedoeagus robust with platelike, folded cornutus. Female unknown. C. peruviensis differs most noticeably from Caviana fuscella Neunzig and Dow, the only other known species in the genus, 1n having: (1) a strongly developed, straight anteme- dial line (rather than an indistinct, dentate antemedial line), and (2) a projection at the apex of the costa of the valva that 1s robust, blunt and directed dorsoposteriorly (rather than an apical, costal projection that is spinelike and curved ventrally). Distribution.— Known only from north- western Peru. Holotype.—é—15km N. Caras, Rio Salta PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Valley, ca. 2000 m, Dept. Ancash, Peru, 19- 1 .ii.1987, O. Karsholt, genitalia slide 3469 HHN (ZMUC). Montestra Neunzig, NEw GENUS Gender. — Feminine. Type species.—Montestra dentata Neun- zig. Antenna simple in both sexes; sensilla tri- chodea (cilia) of male moderately abundant and at base of shaft about *4 as long as basal diameter of shaft. Frons convex. Labial pal- pus (Fig. 13) porrect, long, extending, in both sexes, beyond head about 4 x length of head. Maxillary palpus simple. Haustellum well developed. Ocellus absent. Forewing, of male, (Fig. 7) with basal half of costa slightly convex; underside of wing, of male, with costal fold; with ten veins; R,,,and R; com- pletely united; M, from well below upper angle of cell; M, and M; stalked for slightly less than 2 their lengths; CuA, from lower angle of cell; CuA, from well before lower angle of cell. Hindwing (Fig. 7) with seven veins (LA, 2A and 3A together considered as one vein); Sc + R, and Rs fused for most of their lengths; M, originating slightly be- low upper angle of cell; M, and M, com- pletely fused; CuA, from slightly before lower angle of cell; CuA, from well before lower angle of cell; cell less than '2 length of wing. Male abdominal segment 8 without scale tufts. Male genitalia (Figs. 18, 21) with uncus subtriangular, broadly rounded api- cally; distal process of gnathos a short, bluntly pointed hook; transtilla complete, triangularly shaped; juxta a narrow U-shaped band with very short, setiferous lobes; valva with short, costal tooth; ae- doeagus with elongate, thin cornutus; vin- culum about as long as greatest width. Fe- male genitalia (Fig. 23) with ductus bursae sclerotized for slightly over 1 1ts length; cor- pus bursae membranous, with moderately distinct signum consisting of a loose aggre- gation of 5-6 thornlike spines; ductus sem- inalis joined to corpus bursae just anterior to signum. The male and female genitalia of Mon- testra resemble those of Vitu/a Ragonot and Volatica Heinrich—in all three genera the males have a short, simple, distal hook on the gnathos, a complete, triangularly shaped transtilla, and a juxta with short setiferous arms, and the females have the ductus bur- sae sclerotized for about '2 its length and a membranous corpus bursae. Nevertheless, the transverse markings on the forewing characteristic of Vitula and Volatica are completely lacking in Montestra; instead Montestra has longitudinal bands on a nar- rower forewing. Also, Montestra differs from both Vitula and Volatica in that the costa of the valva has a distinct, dorsally directed toothlike element, a feature lacking in the other genera. Furthermore, the elongate la- bial palpi of Montestra are similar to those of Volatica, but the number of veins in the forewing are fewer in Montestra than in Vol- atica;, the number agrees with the venation found in Vitula, but Vitula has rather short labial palpi. Montestra dentata Neunzig, NEw SPECIES Figs..4, 75, toe Sie Type locality. —35 km SE. Huaraz, Dept. Ancash, Peru. Diagnosis.—M. dentata is a rather large phycitine (average forewing length of eight specimens 18.0 mm) with longitudinally streaked forewings, long, porrect labial palpi, and a costal tooth on the valva of the male genitalia. Description.—Length of forewing 14.0- 20.0 mm. Head with frons and vertex pale brown to pale reddish brown; labial palpus (Fig. 13) brown to pale brown dusted with white (venter of segments, particularly basal segments, sometimes completely white); maxillary palpus brownish white. Thorax with dorsum and collar pale brown to pale reddish brown. Forewing usually mostly pale brown to pale reddish brown with broad VOLUME 97, NUMBER 3 645 Figs. 14-19. Male genitalia. 14, Caviana peruviensis n. sp., aedoeagus omitted. 15, aedoeagus. 16, Euryth- mioides carasensis n. sp., aedoeagus omitted. 17, aedoeagus. 18, Montestra dentata n. sp., aedoeagus omitted. 19, Depeadus deiulus n. sp., aedoeagus and some scales omitted. (All scale lengths 0.5 mm.) brown to dark brown subcostal and median longitudinal streaks and yellowish white pale scales between streaks; forewing of some (like the holotype) with darker scales over almost entire surface, obscuring most lon- gitudinal streaks; antemedial and postme- dial lines not present; discal spots dark brown to black, moderately distinct to ob- 646 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 20-25. Male and female genitalia. 20, Depeadus deiulus n. sp., aedoeagus. 21, Montestra dentata n. sp., aedoeagus. 22, Depeadus deiulus n. sp., ventral view of ductus bursae, corpus bursae and ductus seminalis. 23, Montestra dentatan. sp., ventral view of ductus bursae, corpus bursae and ductus seminalis. 24, Eurythmioides carasensis n. sp., ventral view of posterior part. 25, ventral view of anterior part. (All scale lengths 0.5 mm.) VOLUME 97, NUMBER 3 scure. Underside of forewing, of male, with- out sex-scales. Hindwing pale smoky fus- cous. Male and female genitalia as given under description of genus. Distribution.—collected at 3240 to 4100 m in the Peruvian Cordillera Negra and Cordillera Vilcabamba. Holotype.—é—Quabrada_ Pucavado, Cerro Cahuish, 35 km SE. Huaraz, 4100 m, Dept. Ancash, Peru, 15—18.11.1987, O. Kar- sholt leg., genitalia slide 3463 HHN (ZMUC). Paratypes.—2 2, same collection data as holotype, genitalia slides 3464, 3465 HHN (ZMUC); 2 2, 5 km E. Laguna Pomacanchi, 40 km NW. Sicuani, 3240 m, Dept. Cuzco, Peru, 24.111.1987, O. Karsholt leg. (ZMUC); 3 2, Pueblo Quichas, Quabrada Quichas, 10 km N. Oyon, 4000 m, Dept. Lima, Peru, 24-26.11.1987, O. Karsholt leg (ZMUC). Eurythmioides Neunzig, NEw GENUS Gender. — Masculine. Type species. —Eurythmioides carasensis Neunzig Antenna of both sexes simple (Fig. 8). Front convex with most scales directed an- teriorly to form a conelike projection. La- bial palpus oblique (Fig. 11). Maxillary pal- pus simple. Haustellum well developed. Ocellus ?present. Basal half of costa of fore- wing of male (Fig. 5) slightly convex; un- derside of wing of male with costal fold. Forewing without raised scales; with nine veins; R;,, and R; completely fused; M, from below upper angle of cell; fused M,,, stalked with CuA, for slightly more than '2 their lengths; CuA, from close to lower angle of cell. Hindwing (Fig. 5) with six veins (1A, 2A, and 3A considered together to be one vein); Sc + R, and Rs completely fused; M, from upper angle of cell; M,,; and CuA, stalked for about 2 their lengths; CuA, from close to lower angle of cell; cell about '2 length of wing. Male abdominal segment 8 with scale tufts; some scales forming tufts straight, others sinuous. Male genitalia (Figs. 16, 17) with uncus broadly triangular; gna- 647 thos distally a broad, tonguelike plate; tran- stilla complete, with slender, distal hook; juxta a U-shaped plate with short, setiferous lateral lobes; valva with costa strongly scler- otized for about %4 its length and with scler- otized part terminating in short, slightly curved, dorsal projection; inner base of val- va with setiferous, triangular lobe; aedoea- gus enlarged at base and with a well-scler- otized, saw-toothed, spiral element; vin- culum longer than greatest width. Female genitalia (Figs. 24, 25) with very short apophyses posteriores and apophyses an- teriores; ductus bursae longer than corpus bursae, sclerotized and granulate for about 7s its length with a strongly sclerotized shield behind genital opening, and with a sclero- tized band posterior to opening; ductus bur- sae membranous, oval, with signum a fused cluster of a few thornlike spines near ante- rior end of ductus bursae; ductus seminalis attached to corpus bursae close to signum. The female genitalia of Eurythmioides, especially the very short apophyses, are like those of Ephestiodes Ragonot, but, the greatly reduced wing venation, and the complete transtilla and toothed and spiraled cornutus of the aedoeagus of the male gen- italia of the former genus differ from those of the latter group and, therefore, suggest that Eurythmioides is more closely allied to Eurythmia Ragonot. Eurythmioides carasensis Neunzig, NEW SPECIES Figs.93; 294.05 te Gala a Type locality.— 15 km N. Caras, Rio Sal- ta Valley, ca. 2000 m., Ancash, Peru. Diagnosis.— The platelike distal element of the gnathos, and the slender, distal hook of the transtilla, of the male genitalia are diagnostic. Description. — Length of forewing: 5.5—6.5 mm. Head with front and vertex brown, some scales, particularly near eyes, brown- ish white or tipped with brownish white; labial palpus brown and brownish white, palest on basal segment; maxillary palpus 548 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON srownish white. Thorax with dorsum pale brown to brown, some brown scales faintly tipped with brownish white, collar generally slightly paler than dorsum. Forewing with ground color brown to dark brown dusted with white and brownish white; weakly de- veloped reddish brown streaks or patches mostly in posterior half; costal half of basal area with obscure pale longitudinal streak; antemedial and postmedial lines indistinct, white and brownish white; discal spots dark brown to black, obscure. Hindwing smoky fuscous, darker along veins and near costal and outer and posterior margins. Male and female genitalia (Figs. 16, 17, 24, 25) as in description of genus. Distribution.— Known only from north- western Peru. The few specimens collected have been taken near a river or in a river valley at low to moderate elevations. Holotype.—é— Rio Salta Valley, ca. 2000 m., 15 km N. Caras, Dept. Ancash, Peru, 19-21.i1.1987, O. Karsholt leg., genitalia slide 3470 HHN. (ZMUC) Paratypes.— 2 2—Same collection data as holotype, genitalia slides 3471, 3473 HHN. (ZMUC). 1 6—7 km E. Ogatun, 100 m. E. side of Rio Zana, 35 km SE. Chiclayo, Dept. Lambayeque, Peru, 18.v.1987, N. Krabbe leg., genitalia slide 3481 HHN (ZMUC). ACKNOWLEDGMENTS I thank O. Karsholt (ZMUC, Copenha- gen, Denmark) for giving me the opportu- nity to study the specimens, and R. L. Blinn, L. L. Deitz and D. L. Stephan (NCSU, Ra- leigh, North Carolina) for critically review- ing a draft of this paper. LITERATURE CITED Heinrich, C. 1956. American moths of the subfamily Phycitinae. United States National Museum Bul- letin 207: 1-581. Neunzig, H. H. and L.C. Dow. 1993. The Phycitinae of Belize (Lepidoptera: Pyralidae). North Carolina Agricultural Research Service Technical Bulletin 304: 1-131. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 649-653 NOMENCLATURAL AND SYNONYMICAL NOTES ON THE GENERA DIPLONYCHUS LAPORTE AND APPASUS AMYOT AND SERVILLE (HETEROPTERA: BELOSTOMATIDAE) JOHN T. POLHEMUS University of Colorado Museum, 3115 South York St., Englewood, Colorado 80110. Abstract.—The genus Appasus Amyot and Serville is resurrected from synonymy, and Muljarus Lee placed as a junior synonym (new synonymy). Revised synonymies are given for the genera Appasus and Diplonychus Laporte, and for Diplonychus annulatus Fabricius. Diplonychus heeri Polhemus, New Name, is proposed as a replacement name for Diplon- ychus rotundatus Heer, 1853, a junior secondary homonym of Diplonychus rotundatus (Laporte, 1833). Diplonychus indicus Venkatesan and Rao, 1980 is placed as a junior synonym of Diplonychus rusticus (Fabricius, 1781). Key Words: Genus Appasus Amyot and Serville 1843, REVISED STATUS Naucoris Fabricius, 1803: 111 (in part) Appasus Amyot and Serville, 1843: 430. Type species: Appasus natator Amyot and Serville, 1843: 431 (= Naucoris nepoides Fabricius 1803: 111), Monobasic. Amyotella Spinola, 1850: 48 (no included species). Type species: [Appasus] natator Amyot and Serville, 1843: 431 (= Nau- coris nepoides Fabricius, 1803: 111); fixed by Kirkaldy, 1906. Muljaris Lee, 1991: 10. Type species: Ap- pasus japonicus Vuillefroy, 1864: 141 (as Diplonychus japonicus). Original desig- nation. New Synonymy. Fabricius (1803) described the new spe- cies nepoides under Naucoris. However he retained both annulata Fabricius, 1781, and rustica Fabricius, 1781, under Nepa. Lee (1991) proposed the genus Muljaris to hold Diplonychus japonicus Vuillefroy and Diplonychus major Esaki, which he sep- arated from Diplonychus on the basis of a number of characters. A new generic entity was unnecessary because the type species of Heteroptera, Belostomatidae, Diplonychus, Appasus, taxonomy, synonymy Appasus, Naucoris nepoides Fabricius, has all of the characters used to separate the group containing japonicus and major, thus Muljaris must fall as a junior synonym of Appasus. Lee failed to analyze the described species previously held under Diplonychus and place them into their respective genera, or undoubtedly he would have realized that a genus group name was already available for his newly conceived group. A review of the genera germane to this work was accom- plished by examination of specimens in the Polhemus Collection, which currently holds all described species except stali Mayr, placed by Lee with the species retained in Diplonychus, a placement that seems cor- fect: Lee’s separation of Diplonychus into two genera seems to be well founded on both somatic and male genitalic characters. However several of the characters need modification or clarification, e.g. the mor- phological differences of the hemelytral membrane, and the “lips marks’ (sic) on the scutellum. The purported generic differences of the hemelytral membrane (Lee, 1991, figs. 3A, 650 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ,B) actually represent wing morphs found . a single population of D. rusticus (F.) from /iet Nam that also contains an intermediate morph. The flightless morph with the mem- brane reduced or absent has in addition re- duced straplike flight wings, narrowed and angulate hemelytra distally, and sharp pos- terolateral pronotal angles. The fully alate morph has a well-developed membrane with veins in addition to the transparent margin present in all morphs, normally rounded he- melytra distally, fully developed flight wings, and the posterolateral pronotal angles slop- ing anteriorly. This phenomenon of linked characters in different flight morphs is also common in Naucoridae. The “‘lips marks”’ of the posteroapical an- gle of the scutellum, said to be present in the “rusticus group,” but absent in the “‘ja- ponicus group” is an inconsistent color character that, in my view, should not be used to separate genera. Other differences given by Lee are useful, and found in the key couplet that follows. For details of the male genitalic structures, see figures in Lee (1991). In addition to the differences given by Lee, Appasus species have prominent eyes, very different from those of Diplonychus species, in which the eyes are “streamlined” and flush with the lateral margins of the head. KEY TO APPASUS AND DIPLONYCHUS 1. Lateral eye margins flush with lateral margin of head, not protruding laterally. First antennal segment equal to or longer than the lateral pro- longations of segments II and III, and segment IV. Male pygophore tapering more or less evenly between basal portion and apical semitubular portion. Lateral arms of phallic basal plate each with a low angular medial projection; phallus laterally flat, exophallotheca with sclerotized lateral rhomboid plates; endosoma laterally flat, hatchet-shaped 2.2 3..c1.-.5- Diplonychus Laporte — Lateral eye margins not flush with lateral mar- gin of head, prominent, protruding laterally. First antennal segment shorter than the lateral prolongations of segments II and III, and seg- ment IV. Male pygophore with an abrupt sculptured shoulder between basal portion and apical semitubular portion. Lateral arms of phallic basal plate without projections, smooth; phallus tubular, exophallotheca with narrow sclerotized lateral plates; endosoma tubular, not hatchet-shaped. .... Appasus Amyot and Serville The following species are now assigned to Appasus, with original genus given in brack- ets if applicable: ampliatus alluaudi (Montandon, 1914). [Diplonychus]. New Combination ampliatus ampliatus (Montandon, 1914). [Diplonychus]. New Combination capensis (Mayr, 1871). [Diplonychus]. New Combination grassei ghesquieri (Poisson, 1940). [Diplo- nychus]. New Combination grassei grassei (Poisson, 1937). [Diplony- chus]. New Combination grassei luitikilae (Poisson, 1968). [Diplony- chus]. New Combination kjellanderi (Menke, 1962). [Diplonychus]. New Combination japonicus Vuillefroy, 1864. Restored Com- bination major (Esaki, 1934). [Diplonychus]. New Combination nepoides (Fabricius, 1803). [Naucoris]. Re- stored Combination procerus divoi (Poisson, 1968). [Diplony- chus]. New Combination procerus procerus (Gerstsaecker, 1873). [Diplonychus]. New Combination quadrivittatus Bergroth, 1893. Restored Combination stappersi (Montandon, 1916). [Sphaerode- ma]. New Combination urinator sudanensis (Linnavuorl, 1971). [Diplonychus]. New Combination urinator urinator Dufour, 1863. Restored Combination wittei (Poisson, 1949). [Diplonychus]. New Combination Genus Diplonychus Laporte, 1833 Nepa Fabricius, 1781, vol. 2: 333 (in part) Diplonychus Laporte, 1833: 18. Type spe- cies: Nepa rustica Fabricius, 1781, vol. 2: VOLUME 97, NUMBER 3 333 (= Nepa plana Sulzer, 1776: 92). Monobasic. Sphaerodema Laporte, 1833: 18 (as sub- genus of Diplonychus). Type species: Sphaerodema rotundata Laporte, 1833: 18 (= Nepa annulata Fabricius, 1781: 333). Monobasic. Synonymized by Kir- kaldy, 1906: 151. Diplonycha Spinola, 1837: 53 (variant spell- ing). Atomya Spinola, 1850: 48 (no included spe- cies). Synonymized by Kirkaldy, 1906: ie Nervinops Dufour, 1863: 348. Type species: Nepa rustica Fabricius, 1781, vol. 2: 333 (= Nepa plana Sulzer, 1776: 92). Mono- basic. Synonymized by Kirkaldy, 1906: Hesilie Cyclodema Dufour, 1863: 397 (suggested new name for Sphaerodema Laporte). Synonymized by Kirkaldy, 1906: 151. Nectocoris Mayr, 1871: 432. Type species: Nectocoris stali Mayr, 1871: 432. Mono- basic. Synonymized by Lauck and Menke, 1961: 649. Polhemus (1994) has shown that the type species of Diplonychus Laporte, Nepa rus- tica Fabricius, 1781, is ajunior synonym of Nepa plana Sulzer, 1776, and a primary homonym of Nepa rustica Fabricius, 1775. A petition has been made to the ICZN to preserve the name Nepa rustica Fabricius (Polhemus and Kerzhner, 1995). Therefore, I continue here to use Nepa rustica Fabricius as the type species of the genus Diplonychus. I have studied specimens of Diplonychus indicus Venkatesan and Rao, 1980, kindly furnished by Dr. Venkatesan, and find that this species is inseparable from Diplonychus rusticus (Fabricius) and, therefore, must fall as a junior synonym (New synonymy). Lauck and Menke (1961) gave the syn- onymy of Diplonychus along with an exten- sive discussion. However, at that time Ap- pasus was a synonym. The above synonymy gives the current status of the synonyms still 651 remaining under Diplonychus with the re- moval of Appasus. The following species are now assigned to Diplonychus, with original genus given in brackets if applicable: annulatus (Fabricius, 1781) [Nepa] eques (Dufour, 1863) [Appasus] esakii Miyamoto and Lee, 1966 rusticus (Fabricius, 1781) [Nepa] = planus (Sulzer, 1776) [Nepa] = indicus Venka- tesan and Rao, 1980. stali (Mayr, 1871) [Nectocoris] Diplonychus annulatus (Fabricius, 1781) Nepa annulata Fabricius, 1781: 333; type locality, “in Coromandel Mus. Dom. Banks” (= Coromandel Coast, Madras area, South India); types in London (1) and Copenhagen (2). Belostoma marginata Gray, 1832: 248, pl. 93, fig. 4; type-locality ‘““China.” Synon- ymized by Mayr, 1871: 434. Sphaerodema rotundata Laporte, 1833: 18; type-locality not given. Synonymized by Mayr, 1871: 434. Diplonychus annulatus: Lauck and Menke, 1961: 649. Abedus sangameswari Rupavathi, 1985: 169; type-locality Guntur District, An- dhra Pradesh, India (manuscript species; nomen nudum). See also: Sphaerodema annulatum: Distant, 1906: 35 (brief redescription). Sphaerodema annulatum: Hoffmann, 1941: 8 (catalog; synonymy; distribution). This species has a much more restricted distribution than D. rusticus (Fabricius); however, the two species are geographically sympatric in southern India. The two spe- cies are easily distinguished, as the latter is much smaller and less rotund. The synonymy above places the species Sphaerodema rotundata Laporte in the ge- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Diplonychus, and establishes the sec- ndary homonym discussed below. Diplonychus heeri Polhemus, NEw NAME Diplonychus rotundatus Heer, 1853: 85 (Tertiary fossil). Junior secondary hom- onym of Diplonychus rotundatus (La- porte, 1833: 18) = Nepa annulata Fabri- cius, 178123333 Diplonychus rotundatus Heer, 1853 is a junior secondary homonym of Sphaerode- ma rotundata Laporte, 1833. The latter and its senior synonym Nepa annulata Fabri- cius, 1781 were placed in the genus Diplon- ychus by Lauck and Menke (1961) when they synonymized Sphaerodema (see syn- onymy of Diplonychus above). This leaves Diplonychus rotundatus Heer without a name; thus, for this taxon I propose the name Diplonychus heeri Polhemus. Another fossil, Sphaerodema jurassicum Oppenheimer, 1888, was transferred by Handlirsch (1906: 543) to his new genus Sphaerodemopsis, and is presently placed in Naucoridae. ACKNOWLEDGMENTS This research was sponsored by a series of grants from the National Geographic So- ciety, Washington, D.C. (1806-77, 2698-83, 3053-85, 3398-86, 4537-91), and by grant BSR-9020442 from the National Science Foundation, Washington, D.C. We thank these organizations for their continued sup- port on research into the systematics and zoogeography of aquatic Heteroptera. LITERATURE CITED Amyot, C. J. B. and J. G. A. Serville. 1843. Histoire naturelle des insectes, Hémiptéres. Libraire En- cyclopedique de Roret, Paris. Ixxvi + 675 + 6 pp., 12 pls. Distant, W. L. 1906. The Fauna of British India In- cluding Ceylon and Burma. Rhynchota. Vol. III (Heteroptera-Homoptera). Taylor & Francis, Lon- don. xiv + 503 pp. Dufour, L. 1863. Essai monographique sur les Bé- lostomides. Annales de la Société Entomologique de France (4)3: 373-400. Fabricius, J.C. 1775. Systema Entomologiae, Sistens Insectorum Classes, Ordines, Genera, Species, Adjectis Synonymis, Locis, Descriptionibus, Ob- servationibus. Kortii, Flensburgi & Lipsiae. xxx + 832 pp. 1781. Species insectorum exhibentes eorum differentias specificas, synonyma auctorum, etc. . . Bohn, Hamburg & Kilonil. Vol. 2, 494 pp. + Ap- pendix (pp. 495-514) + Index (pp. 515-517). . 1803. Systema Rhyngotorum Secundum Or- dines, Genera, Species Adjectis Synonymis, Locis, Observationibus, Descriptionibus. Carolum Rei- chard, Brunsvigae. vi + 314 pp. Gray, G. 1832. Notices of new genera and species in Griffith, E. & E. Pidgeon, eds. The class Insecta arranged by the Baron Cuvier, with supplementary additions to each order, volume 2, 796 pp.; vol- ume 15 of Griffith, E., The animal kingdom ar- ranged in conformity with its organization by the Baron Cuvier, with additional descriptions of all the species hitherto named, and of many not be- fore noticed. Geo. B. Whittaker, London. 16 vols. Handlirsch, A. 1906-1908. Die Fossilen Insekten und die Phylogenie der Rezenten Formen. Part IV, pp. 481-672, 1906. Engelmann, Leipzig. xi + 1430 pp., 51 pls. Heer, O. 1853. Die Insectenfauna der Tertiargebilde von Oeningen und Radoboj in Croatien. William Engelmann, Leipzig. Vol. 3, iv + 138 pp. Hoffmann, W.E. 1941. Catalogue of the aquatic He- miptera of China, Indo-China, Formosa, and Ko- rea. Lingnan Science Journal 20: 1-78 + 5 pp. Kirkaldy, G. W. 1906. List of the genera of the pa- giopodous Hemiptera-Heteroptera, with their type species, from 1758 to 1904 and also of the aquatic and semi-aquatic trochalopoda. Transactions of the American Entomological Society 32: 117-156 + 156a, 156b. Laporte, F. L. de. 1832-1833. Essai d’une classifi- cation systematique de l’ordre des Hémiptéres. Magasin de Zoologie, Paris 2 (Suppl.): 1-88 (pp. 16-88 published in 1833). Lauck, D. R. and A. S. Menke. 1961. The higher classification of the Belostomatidae. Annals of the Entomological Society of America 54: 644-657. Lee, C. E. 1991. Morphological and phylogenetic studies on the true water bugs (Hemiptera: Het- eroptera). Nature and Life 21: 1-183. Mayr, G. 1871. Die Belostomiden. Monographisch bearbeitet. Verhhandlungen der Zoologisch-Bo- tanischen Gesellschaft in Wien 21: 379-440. Polhemus, J. T. 1994. The identity and synonymy of the Belostomatidae (Heteroptera) of Johann Christian Fabricius 1775-1803. Proceedings of the Entomological Society of Washington 96: 687-695. Polhemus, J. T. and I. M. Kerzhner 1995. Nepa rus- tica Fabricius, 1781 and Zaitha stollii Amyot & Serville, 1843 (currently Nepa rustica and Zaitha VOLUME 97, NUMBER 3 stollii: Insecta, Heteroptera): proposed conserva- tion of the specific names. Bulletin of Zoological Nomenclature 51: 40-43. Rupavathi, A. 1985. Systematics and ecology of aquatic Hemiptera of Guntur district, Andhra Pra- desh, India. Unpubl. Ph.D. Dissertation, Nagar- juna Univ., Nagarjunanagar, A. P., India. vii + 298 pp. Spinola, M. M. 1837. Essai sur les insectes Hémip- téres L. ou Rhyngotes F. et a la section des Hét- eroptéres Duf. Yves Graviers, Geneva. 383 pp., 5 folding tables. (Reprinted in 1840, Bailliere, Paris, with slightly different title.) 1850. Tavola sinottica dei generi spettanti 653 alla classe degli insetti artroidignati, Hemiptera, Linn. Latr.-Rhyngota, Fab.-Rhynchota, Burm. R. D. Camera, Modena. 138 pp. Sulzer, J. H. 1776. Abgekiirzte Geschichte der In- secten nach dem Linnéischen System. H. Steiner u. Co., Winterthur. 2 Teile; Teile 1, 27 + 274 pp.; Teile 2, 71 pp., 32 col. pls. Venkatesan, P. & T. K. R. Rao. 1980. Description of a new species and a key to Indian species of Belostomatidae. Journal of the Bombay Natural History Society 77: 299-303. Vuillefroy, F. de. 1864. Hémiptéres nouveaux. An- nales de la Société Entomologique de France 4: 141-142. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 654-658 THE IDENTITY OF PELOCORIS BIIMPRESSUS MONTANDON AND SYNONYMY OF PELOCORIS SPECIES IN THE SOUTHWESTERN UNITED STATES (HETEROPTERA: NAUCORIDAE) JoHN T. POLHEMUS AND ROBERT W. SITES (JTP) University of Colorado Museum, 3115 S. York St., Englewood, CO 80110; (RWS) Wilbur R. Enns Entomology Museum, Department of Entomology, University of Mis- souri, Columbia, Missouri 65211. Abstract. — Pelocoris shoshone amargosus La Rivers 1956 is synonymized with Pelocoris biimpressus Montandon 1898, NEW SYNONYMYyY; a lectotype is designated for the latter. Pelocoris femoratus (Palisot de Beauvois) 1820 is compared to P. biimpressus and dis- cussed, and a neotype is designated. Key Words: A recent review (JTP in prep.) of the ge- nus Pelocoris Stal revealed that P. biim- pressus Montandon is the most common species in northern Mesoamerica with a range extending from Guatemala north- ward into Texas, Louisiana, Arizona, Ne- vada and California, and that Pelocoris shoshone amargosus La Rivers 1956 is a synonym. To characterize species of this difficult ge- nus, several morphological characters have been found to be useful in addition to those used by La Rivers (1948, 1956) and Nieser (1975) (e.g. shape of male ventral lateroter- gites VI and VII, shape of female ventral laterotergite VI, sculpturing of male tergite V, dentition of female ovipositor lobes, shape and setiferation of male parameres). The characters used by La Rivers (loc. cit.) to separate species include the morphology of the female sternum VII (subgenital plate), male aedeagus, male dorsal aedeagal plate, coloration of dorsum, and body size; Nieser added the coloration of the fore femora and hemelytra, and development of spines on the connexiva. Some, but not all, of these Insecta, Heteroptera, Naucoridae, synonymy characters are used below in distinguishing North American species. This contribution is intended to clarify the status of the Pelocoris species of the cen- tral and southwestern United States, from Louisiana and Texas westward; therefore, the voluminous Mexican and Mesoameri- can material in the Polhemus Collection is not treated here, except for a few examples. The resolution of Pelocoris species-group problems in the southeastern United States (particularly Florida), and Mesoamerica plus northern South America are beyond the scope of this paper, and will be treated in future publications. Pelocoris biimpressus biimpressus Montandon Pelocoris biimpressus Stal in litt.; see Mon- tandon 1898: 285, and Champion 1901: 360. Manuscript name, unavailable. Pelocoris biimpressus Montandon 1898. Bull. Soc. Sci. Buc.-Roum. 7: 285. Lec- totype, male, here designated, Mexico, in Stockholm Museum. VOLUME 97, NUMBER 3 Figs. 1-4. femoratus (arrow, posterior margin of laterotergite VI). 3, Male genitalia of Pelocoris biimpressus. 4, Male genitalia of Pelocoris femoratus. Pelocoris biimpressus, Van Duzee 1916. Check List Hem., p. 52. Pelocoris shoshone amargosus La Rivers 1956. Wasmann J. Biol. 14: 155. NEW SYNONYM Y. Diagnosis. — Females of P. biimpressus can easily be distinguished from those of P. fe- moratus by the shape of the posterior mar- gin of ventral laterotergite VI: straight in the former (Fig. 1), distinctly curved in the latter (Fig. 2, arrow). Males can be distin- guished by the shape of the aedeagus (Figs. 3, 4) (see also La Rivers 1948, Figs. 1 M, N). Pelocoris femoratus ordinarily is some- what smaller in stature and lighter in color than is P. biimpressus, which is particularly noticeable in the region where the species co-occur. Both color and size are extremely variable over the larger geographic range of each species and indeed even within a single series and are thus unreliable as key char- acters over most of the range. Possibly, with better methods to discriminate species, each of these taxa will be further subdivided, as they are morphologically polytypic com- 655 1, Abdominal venter of female Pelocoris biimpressus. 2, Abdominal venter of female Pelocoris pared to South American species known to us. The ranges overlap only in Texas and Louisiana as far as is now known, but future collecting will undoubtedly increase the known range of both species. Distribution.—P. biimpressus occurs as far south as Guatemala (Polhemus, in prep.); Montandon (1898) reported biimpressus from Guatemala, Mexico and Uruguay, but his diagnosis was based primarily on color thus his records are potentially unreliable. P. poeyi, as defined by Nieser (1975), occurs in northern South America; Sites (1990) has recently reported that it ranges from Cuba through the Antilles into South America to Ecuador, therefore it is quite possible that the ranges of these two species overlap. Un- fortunately the species are difficult to iden- tify. Although there is a slight difference in the male genitalia, the females are very sim- ilar in appearance, including the shape of female sternum VI. Discussion.—Montandon (1898) and Champion (1901) treated biimpressus as a variety of femoratus, and did not discuss the locality of any type material. Van Duzee 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (916, 1917), Hungerford (1920), La Rivers 948, 1971) and Polhemus and Polhemus (1988) treated biimpressus as a valid species without significant comment. La Rivers (1948) briefly discussed the species in con- nection with his proposal of P. shoshone from warm springs in Nevada, but in spite of the realization that P. biimpressus was a North American species, he made no at- tempt to recognize it or compare it with his P. shoshone, and as a consequence described a synonym. During a survey of the collections of Eu- rope, JTP noted a syntype series of Pelocoris biimpressus consisting of two specimens in the Natural History Museum at Stockholm. This is Stal’s (1876) manuscript species re- ferred to by Montandon (1898) and Cham- pion (1901). Through the kindness of Dr. Per Lindskog we have been able to study these two females. The first of these bears the following labels: “Mexico”; ‘“‘Sallé’’; “‘biimpressus Stal” handwritten; “Typus”’ on red card. Without a previous lectotype designation, this is not a holotype, but only a syntype; we here designate this female as lectotype. The second female bears the la- bels: ““Mexico”’; “‘Stal’’; ““Paratypus” on red card; this specimen is designated a paralec- totype. Material examined.— UNITED STATES: Arizona: Santa Cruz Co.: 1 male, | female, cattle pond | mi. W. of Pena Blanca L., 17- X-78, C. Olson (JTPC). California: San Ber- nardino Co.: many males and females, Sar- atoga Springs, CL 278, 21-II-64, J. T. Pol- hemus (JTPC); 4 males, 2 females, Saratoga Springs, 27-I-57, Menke & Stange (JTPC); 13 males, 8 females, Death Valley Nat. Mon., Saratoga Springs, 22-XI-90, J. A. Back (UMC); 6 males, 2 females, Saratoga Springs, Death Valley Nat. Mon., El. 61 m, water temp. 28°C, CL 2902, 22-VII-92, J. T. & D. A. Polhemus. (JTPC). Inyo Co:: many males and females, warm spring near Tecopa, CL 280 22-II-64, J. T. Polhemus (JERE); 1. male; 1 female, Tecopas Hot Springs; El. _533'm, water temp; 37°C, CL 2903, 29-VI-93, J. T. & D. A. Polhemus (JTPC). Louisiana: Iberville Par.: 1 female, St. Gabriel Exp. Sta., 26-IX-84, C. B. Barr (JTPC); Tangipahoa Par.: 1 female, 1 1-III- 77,J.E. Barr Sr. (WDSC). Nevada: Nye Co.: many males and females, stream below Point of Rock Springs, Ash Meadows, CL 269, 20-II-64, J. T. Polhemus (JTPC); 3 males, 3 females, | nymph, Purgatory Well, Ash Meadows NWR, El. 716 m, water temp. 33,9°C, CL. 2718; 22-Vil-92) Jn Téa Dy a Polhemus (JTPC); 3 males, Mary Scott Spring, Ash Meadows NWR, El. 716 m, water temp. 28°C, CL 2719, 22-VII-92, J. T. & D. A. Polhemus (JTPC); 1 female, Shaft/Chalk Spring, Ash Meadows NWR, no date, D. Threloff (JTPC); 2 females, Claypit Spring, Ash Meadows NWR, no date, D. Threloff (JTPC). Texas: Victoria Co.: | female, det. Pelocoris biimpressus, Montandon 1909, Victoria, J. D. Mitchell (USNM). GUATEMALA: Progreso: 2 males, | female, Jutiapa, pond, # 632, 23- VI-1957, D. R. Lauck (JTPC). MEXICO: Jalisco: 1 female, Atentique, 5-XII-48, E. S. Ross (JTPC); 2 males, S. of Guadalajara, pond, # 671, 11-VIII-1957, D. R. Lauck (JTPC). Sonora: 1 male, | female, Rio So- nora, E. of Ures, CL 714, 3-11-1975, 7.0. Polhemus (JTPC). Veracruz: 3 males, 4 fe- males, Rio Blanco, La Tinaja, CL 504, 4-I- 1971. J: Died. Me S: Polhemuss(GP@) 23 males, 2 females, S. of Guiterrez Zamora, CL 519, 7-1-1971, J. T. & M. S. Polhemus (JTPC). State unknown: | female, Sallé (lec- totype), | female (paralectotype) (SMNH) (see discussion above). Pelocoris biimpressus shoshone La Rivers, NEw COMBINATION Pelocoris shoshone La Rivers 1948. Ann. Entomol. Soc. Am. 61: 371. Pelocoris shoshone shoshone La Rivers 1956. Wasmann J. Biol. 14: 155. The subspecies P. shoshone shoshone was characterized by La Rivers as being smaller, of lighter color, and with a lighter colored yellowish scutellum in comparison to P. shoshone amargosus (now P. biimpressus VOLUME 97, NUMBER 3 biimpressus), which is dark in coloration and typically has a brownish black scutellum ex- cept the yellowish tip. These differences have not been seen to intergrade in the popula- tions studied so far, therefore the subspecies status is retained for the present. However, the variation in size and color over the range of P. biimpressus in Mexico and Mesoam- erica encompasses both forms described by La Rivers under P. shoshone, and the vari- ation shows no clear geographic separation except for the consistently lighter colored scutellum of P. b. shoshone, thus the sub- species status of these taxa may eventually be suppressed. In California and Nevada, La Rivers’ subspecies amargosus is restricted to springs in the Amargosa River system proper, which begins in the Bullfrog Hills and on Pahute Mesa northwest of Beatty, Nevada and flows (during wet periods) into the southern end of Death Valley, whereas at all other local- ities, including Grapevine Springs at the north end of Death Valley, the White River system, and Railroad Valley, the only form so far seen is the subspecies shoshone. The Grapevine Springs drain into the same en- dorheic basin as the Amargosa River, but are separated by a highly saline sink in cen- tral Death Valley. The presence of P. biim- pressus biimpressus at the south end of the valley, and P. biimpressus shoshone at the north end indicates that in the past these areas were connected to separate drainage systems, but more recent tectonic down- warping has caused them to flow to a com- mon terminus. The localities given below are only for P. biimpressus shoshone in the United States. Material examined.— UNITED STATES: California: Inyo Co.: 1 female, Grapevine Springs, nr. Scotty’s Castle, Death Valley Nat. Mon., D. Threloff, II-93 (JTPC); 1 male, | female, Grapevine Springs, nr. Scot- ty’s Castle, El. 838 m, water temp. 35°C, CL 2901, 28-VI-93, J. T. & D. A. Polhemus (JTPC). Nevada: Nye Co.: 7 males, 5 fe- males, 10 nymphs, Railroad Valley, Duck- water, Big Warm Spring, El. 1768 m, water 657 temp/33°G;CL 2893;,27-V1-93,J. T. & D: A. Polhemus (JTPC); many males and fe- males, Railroad Valley, Duckwater, Little Warm Spring, El. 1768 m, water temp. 33°C, CL 2894, 27-VI-93, J.T. & D. A. Polhemus (JTPC). Lincoln Co.: 6 males, 1 female, Ash Warm Springs, elev. 1143 m, water temp. 36°C, CL 27 a2 lewii-92> Jt Teed DisA; Polhemus (JTPC); 2 males, 4 females, 6 nymphs, Ash Springs, 26-27-VIII-89, algal mats in slow water, J. A. Back (UMC). Clark Co.: 12 males, Moapa Warm Springs, N. of Moapa, El. 550 m, water temp. 32°C, CL 423, 16-IV-68, J. T. Polhemus (JTPC); 4 males, 14 nymphs, Warm Springs, 32°C, al- gal mats in slow water, 27-VIII-89, J. A. Back (UMC); 2 males, 2 females, 13 mi. S. Overton, Blue Point Spring, 12-XII-86, W. D. Shepard (WDSC, JTPC). Pelocoris femoratus femoratus (Palisot de Beauvois) Naucoris femorata Palisot de Beauvois 1820. Ins. Afr. Amer. 14: 237, pl. 20, fig. 4. Type-locality “Etats-Unis d’Amerique.” Neotype, male, New Jersey, here desig- nated. Pelocoris femoratus, Stal 1876. Svenska Ve- tensk. Akad. Handl. 14: 144. Naucoris poeyi, Uhler 1876. Bull. U.S. Geol. Surv. Terr. 1: 71. Misidentification. Pelocoris femoratus, Montandon 1898. Bull. Soc. Sci. Buc.-Roum. 7: 284-286. Diag- nosis, synonymy. Pelocoris carolinensis, Hungerford 1927. Bull. Brook. Entomol. Soc. 22: 77. Life history. Misidentification. Distribution. —Pelocoris femoratus ap- parently is restricted to the eastern part of the United States and Canada. We have not yet seen specimens from Mexico, although it is common in eastern Texas (Sites and Polhemus, in press). The distribution given by Polhemus and Polhemus (1988) includ- ed extralimital records from the literature that must now be reconfirmed. Montandon (1898) cited localities in the United States and Panama; the latter record certainly per- 658 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tains to another species. The infraspecific Pelocoris femoratus balius La Rivers is known from only Florida. A neotype is here designated in order to stabilize the nomenclature of Naucoris fe- moratus Palisot de Beauvois, as the type material has never been located and is pre- sumed lost. It is a male from New Jersey, Somerset Co., Basking Ridge, CL 384, 31 May 1967, J. T. Polhemus, in USNM. The type locality is here restricted to New Jersey. Distributional data for this species was given by Polhemus & Polhemus (1988), and amplified by Sites and Polhemus (in press) for the state of Texas where the ranges of P. biimpressus and P. femoratus overlap, therefore these data are not repeated here. ACKNOWLEDGMENTS We are indebted to the following for the loan or gift of material: Dr. Paul Arnaud, California Academy of Sciences, San Fran- cisco (CAS); Dr. R. L. Froeschner, Smith- sonian Institution, Washington, D. C. (USNM); Dr. Per Lindskog, Swedish Mu- seum of Natural History (SMNH); Drs. C. B. Barr and W. D. Shepard, California State University, Sacramento (WDSC). Speci- mens in the Polhemus Collection are marked (JTPC) and those in the Wilbur R. Enns Entomology Museum, University of Mis- souri—Columbia are marked (UMC). Fund- ing was provided in part for RWS by Uni- versity of Missouri project #PSSLO232. This is Missouri Agricultural Experiment Station Journal Series paper 12, 109. LITERATURE CITED Champion, G. C. 1897-1901. Insecta: Rhynchota (Hemiptera-Heteroptera). Vol. II. Jn Godman, F. D. & O. Salvin, eds., Biologia Centrali-Americana. London, xvi + 416 pp., 22 pls. (i-xvi, pp. 345- 416, 1901). Hungerford, H. B. 1920. The biology and ecology of aquatic and semiaquatic Hemiptera. University of Kansas Science Bulletin 11: 1-328, col. pls. 1-3, pls. 1-30. . 1927. Life history of the creeping water bug, Pelocoris carolinensis Bueno. Bulletin of the Brooklyn Entomological Society 22: 77-82, | pl. La Rivers, I. 1948. A new species of Pelocoris from Nevada, with notes on the genus in the United States (Hemiptera: Naucoridae). Annals of the En- tomological Society of America 61: 371-376. . 1956. Anew subspecies of Pelocoris shoshone from the Death Valley drainage (Hemiptera: Nau- coridae). Wasmann Journal of Biology 14: 155- 158. 1971. Studies of Naucoridae (Hemiptera). Catalogue of taxa described in the family Nau- coridae. Biological Society of Nevada Memoire 2: 65-120. Montandon, A. L. 1898. Hemiptera cryptocerata. Notes et descriptions d’especes nouvelles. Bulletin de la Société des Sciences de Bucarest-Roumanie 7: 282-290. Nieser, N. 1975. The water bugs (Heteroptera: Ne- pomorpha) of the Guiana region. Studies on the fauna of Suriname and other Guyanas: No. 59, 310 pp., 24 pls. Palisot de Beauvois, A. M. F. J. 1805-1821. Insectes recueillis en Afrique et en Amérique, dans les roy- aumes d’Oware et de Benin, a Saint Dominique et dans les Etats Unis, pendant les années 1786- 1797. Levrault, Paris, xvi + 276 pp. (Aquatic Het- eroptera in Livr. 14, pp. 234-237, pl. 20, 1820.) Polhemus, D. A. and J. T. Polhemus. 1988. Family Naucoridae Leach, 1815. The creeping water bugs, pp. 521-527. In Henry, T. J. & R. C. Froeschner, eds., Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden, xix + 958 pp. Sites, R. W. 1990. Naucorid records from Amazo- nian Ecuador (Heteroptera: Naucoridae). Florida Entomologist 73: 334-335. Sites, R. W. and J. T. Polhemus. The Pelocoris (He- miptera: Naucoridae) fauna of Texas. Southwest- ern Naturalist (in press). Stal, C. 1876. Enumeratio Hemipterorum. Bidrag till en forteckning 6fver alla hittills kinda Hemiptera, jemte systematiska meddelanden. 5. Kunglica Svenska Vetenskapsakademiens Handlingar 14: 1-162 Uhler, P.R. 1876. List of the Hemiptera of the region west of the Mississippi River, including those col- leted during the Hayden Explorations of 1873. Bulletin of the U.S. Geological and Geographical Survey of the Territories 1: 269-361, pls. 19-21. Van Duzee, E. P. 1916. Check List of the Hemiptera (Excepting the Aphidae, Aleurodidae and Cocci- dae) of America North of Mexico. New York En- tomological Society, New York, xii + 111 pp. . 1917. Catalogue of the Hemiptera of America north of Mexico, excepting the Aphidae, Aleurod- idae and Coccidae. University of California Pub- lications, Technical Bulletins, Entomology, 2: i- xvi + 1-902. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 659-665 THE CHEWING LOUSE GENUS AOTIELLA (PHTHIRAPTERA: GYROPIDAE) FROM SOUTH AMERICAN NIGHT MONKEYS, AOTUS (PRIMATES: CEBIDAE) ROGER D. PRICE AND ROBERT M. TIMM (RDP) Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108 (Current address) 4622 Kinkead Avenue, Fort Smith, Arkansas 72903; (RMT) Museum of Natural History and Department of Systematics and Ecology, University of Kansas, Lawrence, Kansas 66045. Abstract.—The single previously recognized species of Aotiella Eichler, A. aotophilus (Ewing), and a second species, 4. hershkovitzi new species, are described and illustrated. The type host for the former is a red-necked night monkey, Aotus azarai (Humboldt), from Bolivia and Argentina, and for the latter a gray-necked night monkey, A. trivirgatus (Humboldt), from Venezuela. The host and louse distributions are discussed. Key Words: The only previously recognized species of the chewing louse genus Aotiella Eichler (Phthiraptera: Amblycera: Gyropidae) is 4. aotophilus (Ewing), which was described by Ewing (1924:25) as a Tetragyropus Ewing from a single immature specimen collected offa monkey “‘Aotes boliviensis . . . taken in Bolivia by W. E. Moore some time before 1859.” Not only was this species based on a single individual thought by Ewing to be the “last nymph,” but that specimen sub- sequently was determined to be only a sec- ond-instar nymph. Ewing justified the new species description from a nymph because of the uniqueness of its being from a primate and the certainty of its breeding on that host. Subsequent collecting from night mon- keys (genus Aotus Illiger) has confirmed that they are indeed hosts for this louse genus. Werneck (1936) presented a detailed de- scription of both sexes and of nymphal stages of what he presumed to be Gyropus aoto- philus based on specimens from 4otus tri- virgatus (Humboldt) from Brazil. Werneck even sent Ewing a nymph to compare with Phthiraptera, Gyropidae, Aotiella, Cebidae, Aotus the holotype and Ewing replied that Wer- neck’s lice were indeed JT. (= Gyropus) aoto- Dhilus. Eichler (1949) seized upon the unique- ness of this louse taxon and described the new genus Aoftie//a for it, although he almost certainly did not actually examine speci- mens. He was especially impressed with the characteristic chaetotaxy of the female and, no doubt, with the unusual primate host. Hopkins and Clay (1952) accepted the va- lidity of the genus Aoftiel/a, but erroneously listed Gyropus Nitzsch as the original genus instead of the correct one, Tetragyropus. Emerson and Price (1975) perpetuated the error of Gyropus as the original genus as given by Hopkins and Clay (1952). They did, however, provide excellent illustrations of both sexes of what they assumed to be A. aotophilus based on abundant material from seven specimens of Aotus trivirgatus captured in Venezuela. All previous workers apparently had been so fixated on the existence of a single species of Aotiella on night monkeys of the genus fy 60 (otus that they failed to examine critically the available material. Had they done so, they would perhaps have found what we have—mainly, that there are two distinctly different species of Aotiella on these hosts, one which occurs south of the Amazon and one north of the Amazon. For this reason, we are herewith describing the adult of Aotiella aotophilus for the first time and naming and describing a second species of this genus. In the following descriptions, all mea- surements are in millimeters. The scientific names of the hosts follow those of Hersh- kovitz (1983) and updates by Groves (1993). The holotype of the new species will be de- posited in the U.S. National Museum of Natural History (Washington, D.C.) and paratypes will be located in the collections of that museum and those of the University of Minnesota (St. Paul) and Oklahoma State University (Stillwater). Acronyms desig- nating museum collections where speci- mens of the host are deposited are as fol- lows: AMNH = American Museum of Nat- ural History, New York; USNM = USS. Na- tional Museum of Natural History. Aotiella aotophilus (Ewing 1924) Figs. 1-5 Tetragyropus aotophilus Ewing 1924: 23. Type host: Aotus boliviensis Elliot = A. azarai (Humboldt). Female.—As in Fig. 1. Head with dorsal temple having 8 long to very long setae; few minute to short dorsal setae, numerous ven- tral setae; head width, 0.39-0.41; head length, 0.28-0.31. Pronotum with 4 long setae posteriorly, no central setae; proster- nal plate with 12 or so medium to long setae; prothorax width, 0.32-0.34. Pteronotum with 5-6 long posterior setae, 2 long median setae, and shorter setae as shown; meso- sternal and metasternal plates with chae- totaxy as in Fig. 1; pteronotum width, 0.44—- 0.50. Legs with tarsus on I much smaller than those on II-III, with latter bearing re- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ceptacle on base of femur for insertion of tarsus. Abdomen with terga II-IV having clusters of long setae, with total number of long setae being: ITI, 4-7; III, 11-18; IV, 16- 20. Terga V—VIII each with pair of median long setae. Total number of other setae on these segments: IT, 5-14; HI-IV, 5-10; V- VI, 7-12; VII, 6-8; VIII, 4-8. Pleura II-III each with single long seta, [V—VII each with single very long seta, and VIII with pair of very long setae, in addition to sparse short setae. Spiracles on segments IJJ-VIII. Ab- domen width at V, 1.01—1.05. Last segment dorsally with pair of very long setae on each side, row of short setae between them. Ster- nal setae, with lengths as shown: II, 7; III- IV, 12-18; V-VI, 11-15; VII, 14-16; VIII, 22-27. Anus (Fig. 2) ventrally with 5-10 longer setae on each side and 4 short median setae, dorsally with 8-14 longer setae on each side and 4 short median setae. Internal genital chamber structure as in Fig. 2, with evident spination on wall limited to pos- terior portion, as shown. Total body length, 2.06-2.15. Male.—As in Fig. 5. Head and thorax much as for female. Tarsus I (Fig. 4) with smooth posterior margin. Head width, 0.40- 0.41; head length, 0.26-0.29. Prothorax width, 0.29-0.33. Pterothorax width, 0.44—- 0.48. Abdominal terga with chaetotaxy of intermixed short to long setae: IJ, 11-17; III, 17-26; IV, 20-26; V—VIII, 14-22. Ab- domen width at V, 0.81-0.85. Last tergum with 2 very long lateral setae on each side and row of 10-12 shorter setae between them. Sternal setae with lengths much as for female: II, 7-8; III, 12-15; IV, 15-16; V- VIII, 10-17. Last sternum with group of very long posterior setae on each side. Gen- italia as in Fig. 3; parameres slender, gently curved outward; mesomere broadly tapered to point; sac without conspicuous larger sp1- nation; width, 0.19-0.22; length, 0.49-0.51. Total body length, 1.77-1.88. Discussion.— This species is readily rec- ognizable by the combination of only up to 20 long setae on each of female abdominal 661 VOLUME 97, NUMBER 3 Figs. 1-5. Aotiella aotophilus. 1, Female. 2, Female genital chamber and anus. 3, Male genitalia. 4, Male first tarsus. 5, Male. 662 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON -rga III-IV (Fig. 1), shape and spination of ie female genital chamber (Fig. 2), small size and features of the male genitalia (Fig. 3), male first tarsus with a smooth posterior margin (Fig. 4), and both sexes with only up to six long posterior setae on the pter- onotum. Since Ewing (1924) based the original de- scription only on a single nymph and the adults described and illustrated by Werneck (1936) and Emerson and Price (1975) were based on what they mistakenly held to be Aotiella aotophilus, our description of A. aotophilus represents the first for adults of this species. Only the single species of Aotus, A. azarai (A. boliviensis is a junior syn- onym), occurs in Bolivia and there is no reason to suspect that Ewing’s nymph was not from that host taxon. Material examined.— Holotype, 2nd in- star, ex Aotus azarai (USNM 3335), Boliv- ia: no specific locality. Additional material: 2 2, 3 46, ex A. azarai (AMNH 211460), Bo- livia: Beni, Mamore River; 3 9, 2 4, ex A. azarai, Bolivia: no specific locality; 2 9, 2 6, ex A. azarai (Brit. Mus. 1976-263), Ar- gentina: Formosa: Pirané and Grande Guardia. Aotiella hershkovitzi Price and Timm, NEw SPECIES Figs. 6-10 Type host: Aotus trivirgatus (Humboldt). Female. — Head and thorax essentially as for A. aotophilus (Fig. 1). Head width, 0.41- 0.44; head length, 0.28-0.32. Prothorax width, 0.32—0.38. Pteronotum with at least 8 long setae posteriorly, in addition to short setae; pteronotum width, 0.48-0.55. Ab- domen (Fig. 6) close to that of A. aotophilus, except as follows. With more long setae on terga II-IV: II, 8-12; II, 30-39; IV, 33-47. Total of other shorter setae on these seg- ments: II, 12-18; I-IV, 5—9; V—VI, 4-8; VII, 2-4; VIII, 2. Pleura III each with single very long seta. Abdomen width at V, 1.09- 1.20. Sternal setae, with lengths as shown: II, 6-9; HI-IV, 15-19; V—VI, 14-17; VII, 17-26; VIII, 25-37. Anus (Fig. 7) ventrally with 9-11 longer setae on each side and 4 short median setae, dorsally with 12-17 longer setae on each side and 4 short median setae. Internal genital chamber structure as in Fig. 7, with evident spination covering entire wall, as shown. Total body length, 2.09-2.25. Male.— Head and thorax much as for A. aotophilus (Fig. 5). Tarsus I (Fig. 8) with serrated posterior margin. Head width, 0.45- 0.47; head length, 0.28-0.31. Prothorax width, 0.35-0.43. Pterothorax width, 0.50- 0.55. Abdomen as in Fig. 10. Terga with chaetotaxy of intermixed short to long setae: II, 19-23; TI, 28-35; IV, 27-32; V-VIII, 15-20. Abdomen width at V, 0.94-1.04. Pleura II each with long to very long seta, III with very long seta. Last tergum with 2 very long lateral setae on each side and row of 11-17 short to long setae between them. Sternal setae with lengths as in Fig. 10: II, 8-9: III, 17-22; IV, 19-22; V—VIII, 14-21. Genitalia as in Fig. 9; parameres relatively broad, abruptly curved outward; mesomere narrowly tapered to point; sac with con- spicuous pair of large spines; width, 0.27- 0.31; length, 0.63-0.71. Total body length, 2.05-2.19. Discussion.—This distinctive species 1s readily separable from Aotiella aotophilus by having at least 30 long setae on each of female abdominal terga HI-IV (Fig. 6), a different shape and extensive spination of the female genital chamber (Fig. 7), the unique structure and considerably larger di- mensions of the male genitalia (Fig. 9), the first male tarsus with a serrate posterior margin (Fig. 8), and both sexes with at least eight long posterior setae on the pterono- tum. Prior to our recognition and description of Aotiella hershkovitzi, only a single species of the genus had been recognized and it was believed to be widely distributed across all taxa of Aotus. In fact, this is likely what led Werneck (1936) and Emerson and Price (1975) to describe and illustrate improperly VOLUME 97, NUMBER 3 l i thes p/ / le ’ an l Hee. ,/ a hea a ris i & MIs nema SS Vag Tes \ ] SS ee J 1 y TON YY Ha \) \ i A NMI 663 Figs. 6-10. Aotiella hershkovitzi. 6, Female posterior pteronotum and abdomen. 7, Female genital chamber and anus. 8, Male first tarsus. 9, Male genitalia. 10, Male posterior pteronotum and abdomen. identified lice as A. aotophilus. A large num- ber of excellent characters separating these two louse species were overlooked by pre- vious workers. We have seen only a single female Aotiella from Brazil. We note its similarity to our Venezuelan type series and tentatively in- clude it in the material examined for A. hershkovitzi, but, given the potential diver- sity of the host taxa, we are reluctant to conclude that it is definitely conspecific with that species. Material examined.— Holotype 2, ex Aotus trivirgatus, Venezuela: Amazonas, Rio Manapiare, San Juan, 155 m, 24 July 1967, Tuttle Team, #28550. Paratypes: 6 9, 2 4, same data as holotype; 18 9°, 9 4, same, ex- cept 5 July 1967, #19989, 19990, or 19966; 664 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2 , same, except 12 July 1967, #26214; _ 11 6, same, except Raya, 32 km SSE Puerto Ayacucho, 135 m, 11 Oct 1967, #31686; 2 6, same, except Yaracuy, Cara- bobo, 19 km NW Urama, 525 m, 27 Oct 1965, Peterson Team, #1998. Additional material: 1 2, ex A. trivirgatus, Brazil: Para, 1935, F. L. Werneck. Etymology.—This species is named for Philip Hershkovitz, Curator Emeritus of Mammals, Field Museum of Natural His- tory, Chicago, in recognition of his many contributions to Neotropical mammalogy. Phil has often created controversy when none existed previously, but he certainly has pushed our science forward and was correct in assessing that the night monkeys are much more complicated than was believed pre- viously. His published contributions, fla- vored by his colorful writing style, and out- standing collections will keep generations of scientists busy. Type species of Aotiella. — Eichler (1949: 11), in erecting the genus Aofiella, stated that it was “... monotypisch auf Gyropus (Tetragyropus) aotophilus Ewg. 1924 sensu Werneck 1936... ab Aotus trivirgatus.”’ He did not indicate having seen any specimens, but likely acted solely on the basis of the description by Werneck (1936). Even though Eichler (1949) referred to the Werneck (1936) material as the type species of Aotiel- la, he did so only because this was the first description of adult specimens and no one suspected that they were anything other than true A. aotophilus. By emphazing the mono- typical nature of the new genus and by his belief that he was referring to 4. aotophilus, we see no reason not to regard this species as the type species of Aotiella. This action is consistent with all citations in the liter- ature and to do otherwise would needlessly confuse matters. If Eichler had any suspi- cion that it was anything other than A. aoto- philus, he most certainly would have de- scribed the new species himself. Host/louse discussion.— Night monkeys are the only cebid primates not naturally susceptible to infection by malaria proto- zoans (Plasmodium), and, hence, are of great interest to medical researchers. Because of this, some aspects of their biology have re- ceived intense study, especially genetics, be- havior, and blood biochemistry. However, there have been few reports of ectoparasites on free-ranging night monkeys, 1n part be- cause they are nocturnal, secretive, and sel- dom observed in the wild, and now also because all populations are protected under international CITES conventions and trea- ties. Our discovery of a new species of chew- ing louse on night monkeys is of interest not only because it documents the species di- versity of night monkey parasites, but also because it provides another independent line of evidence further supporting recent tax- onomic changes within the genus Aotus that suggest the genus is more diverse than was realized previously. Night monkeys are widely distributed in the Amazon Basin of South America, with one species also occurring in Central Amer- ica in Panama and perhaps in Costa Rica (Hershkovitz 1983, Timm 1989). They can be quite common locally, although there are not many specimens in museum collections. Ecology and distribution of night monkeys are poorly known, in part because they are seldom observed clearly and are difficult to study. They are the only truly nocturnal Neotropical monkeys and they are strictly arboreal, foraging only in the canopy, rarely, if ever, coming to the ground. Traditionally, most authors regarded all populations of night monkeys as the single widespread species Aotus trivirgatus. How- ever, in a recent revision, Hershkovitz (1983) recognized nine allopatric species of Aotus, based on differences in chromosome numbers and in pelage coloration. Subse- quently, a tenth species, 4. hershkovitzi Ra- mirez-Cerquera, has been described. Hershkovitz (1983) divided the genus Aotus into two species groups, the gray- necked species, which occur primarily north of the Rio Amazonas, and the red-necked VOLUME 97, NUMBER 3 species, which occur primarily south of the Rio Amazonas. He recognized four species within the gray-necked group: A. brumbacki Hershkovitz in eastern Colombia; 4. /e- murinus (I. Geoffroy) in Panama, Colom- bia, and Ecuador; 4. trivirgatus in Vene- zuela and Brazil; and A. vociferans (Spix) in Colombia, Ecuador, and Brazil. He recog- nized five species within the red-necked group: A. azarai in Bolivia, Paraguay, and northern Argentina (see Groves 1993 for correction of Hershkovitz’s original spell- ing), 4. infulatus in southern Brazil; A. mi- conax Thomas in eastern Peru; 4. nancy- maae Hershkovitz (see Groves 1993 for correction of the original spelling) in eastern Peru; and 4. nigriceps Dollman in western Brazil and adjacent Peru. Aotus hershkovitzi is amember of the gray-necked species group and is known only from the type locality in eastern Colombia. The distributions of the two known spe- cies of Aotiella are such that A. aotophilus is found on the red-necked night monkey species group in Bolivia and Argentina, and A. hershkovitzi is on the gray-necked group in Venezuela and possibly Brazil. Whether these will ultimately prove to be the only species of Aotie/la or whether other species exist must await additional collections of lice from other Aotus. ACKNOWLEDGMENTS We thank Oliver S. Flint, Jr., and Nancy Adams, U.S. National Museum of Natural History; John Chemsak, University of Cal- ifornia, Berkeley; and Christopher H. C. Lyal, The Natural History Museum, for the generous loan of material that contributed 665 to the success of this paper. Our apprecia- tion also goes to Hans-Peter Schultze and Marcus Vieira, University of Kansas, for their assistance in translating, respectively, selected passages of German and Portu- guese literature. This study was partially supported by Project No. Min-17-015, Minnesota Agricultural Experiment Sta- tion, St. Paul, Minnesota 55108, and has been assigned Paper No. 21,179, Scientific Journal Series. LITERATURE CITED Eichler, W. 1949. Phthirapterorum nova genera. Bol- lettino della Societa Entomologica Italiana 79: 1 1- 13: Emerson, K. C. and R. D. Price. 1975. Mallophaga of Venezuelan mammals. Brigham Young Uni- versity Science Bulletin, Biological Series 20(3): 77 pp. Ewing, H. E. 1924. On the taxonomy, biology, and distribution of the biting lice of the family Gy- ropidae. Proceedings of the United States National Museum 63 (Article 20): 42 pp. + 1 pl. Groves, C. P. 1993. Order Primates, pp. 243-277. In Wilson, D. E. and D. M. Reeder, eds., Mammal Species of the World: A Taxonomic and Geo- graphic Reference, 2nd ed. Smithsonian Institu- tion Press, Washington, xviii + 1206 pp. Hershkovitz, P. 1983. Two new species of night mon- keys, genus Aotus (Cebidae, Platyrrhini): A prelim- inary report on Aotus taxonomy. American Jour- nal of Primatology 4: 209-243. Hopkins, G. H. E. and T. Clay. 1952. A Check List of the Genera & Species of Mallophaga. British Museum (Natural History), London. 362 pp. Timm, R. M. 1989. A review and reappraisal of the night monkey, Aotus lemurinus (Primates: Cebi- dae), in Costa Rica. Revista de Biologia Tropical 36(2B)(1988): 537-539. Werneck, F. L. 1936. Contribuicao ao conhecimento dos Mallophagos encontrados nos mammiferos sul- americanos. Memorias do Instituto Oswaldo Cruz 31: 391-589 + 1 pl. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 666-688 A REVISION OF THE SHORE-FLY GENUS RHYSOPHORA CRESSON (DIPTERA: EPHYDRIDAE) NAVIN RAO AND WAYNE N. MATHIS Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. Abstract.—The New World genus Rhysophora Cresson, now placed in the tribe Dis- comyzini (subfamily Discomyzinae), is revised. Four species, including R. griseola (type locality: Costa Rica. Guanacaste: Colorado), which is newly described, are included. The phylogenetic relationships of the species of Rhysophora are reanalyzed, and brief com- ments are provided on the relationships of the genus within the tribe. A key to the genera of the tribe is included. Key Words: The subfamily Discomyzinae, presently comprising two tribes, 22 genera, and nearly 200 species (Zatwarnicki 1992, Mathis and Zatwarnicki, in preparation), is one of the smaller subfamilies of shore flies. One of the tribes, Discomyzini, with half of the gen- era (11) but only one-fourth of the species (ca. 50), includes the genus Rhysophora Cresson, the subject of this revision. Cresson (1924) described Rhysophora as a monotypic genus with R. robusta Cresson as its type species. Cresson (1946) later transferred two previously described spe- cies, Psilopa umbrosa Loew and P. caeru- leiventris Loew, to Rhysophora. Mathis and Wirth (1977), however, placed the latter two species in Nesopsilopa Mathis and Wirth, which is now recognized as a subgenus of Guttipsilopa Wirth (Mathis and Zatwar- nicki 1990). Since the original description of Rhysophora, most references to the genus have been part of generalized studies, such as keys to genera (Cresson 1942, Sturtevant and Wheeler 1954, Wirth and Stone 1956) or regional catalogs (Wirth 1965, 1968). An exception is Mathis’ (1977) revision of the genus, which included two new species, R. ardeoceras and R. liropus, and provided the Diptera, Ephydridae, shore flies, Rhysophora, phylogeny first phylogeny for the species. Mathis’ phy- logenetic analysis resulted in two basal lin- eages that were recognized as species groups: the ardeoceras group (R. ardeoceras) and the robusta group (R. robusta and R. liropus) as sister groups. A sister group to Rhysophora was not identified. The phylogenetic relationships Rhyso- phora has with related genera within Dis- comyzini remain largely unresolved, al- though Zatwarnicki (in litt.) is now inves- tigating them. Previously, Zatwarnicki (1992) accorded subfamilial status to Dis- comyzinae and reconstituted the two in- cluded tribes with Rhysophora and 10 other genera being moved from Psilopini to Dis- comyzini. Information on the natural history and distribution of Rhysophora is meager. Adults are collected infrequently, resulting in a paucity of specimens and locality records. Some species are associated with plants of the family Pontederiaceae, perhaps as her- bivores. Virtually nothing is known about the larval stages, behavior, or life history of Rhysophora. The species of Rhysophora are known thus far only from the New World. The purpose of this study is to revise the VOLUME 97, NUMBER 3 species, including the description of a new species, and to reanalyze the phylogeny of the species. We also discuss, albeit briefly, the position of Rhysophora within the tribe Discomyzini. Methods.—The descriptive terminology, with the exceptions noted in Mathis (1986) and below, follows that published in the Manual of Nearctic Diptera (McAlpine 1981). The species descriptions are com- posite and not based solely on the holo- types. We have discovered or reinterpreted a majority of the characters used in this study, especially those from the male post- abdomen, and for that reason we are pro- viding descriptions of all species. Four head and two venational ratios used in the de- scriptions are defined below (all ratios are averages of three specimens (the largest, smallest, and one other), except R. ardeo- ceras for which two specimens were mea- sured): 1. Frons width-to-length ratio is the frons width divided by the frons length. The length is measured from the anterior margin of the frons to the posterior mar- gin of the posterior ocelli. Width is mea- sured at the level of the anterior ocellus. 2. Face width-to-height ratio is the narrow- est width between the eyes divided by the height. 3. Gena-to-Eye ratio is the genal height measured at the maximum eye height divided by the eye height. In previous publications, this ratio was designated as the “‘eye-to-cheek ratio.” 4. Eye width-to-height ratio is the eye width divided by the eye height, where both measurements are the longest distances taken with the eye oriented laterally. 5. Costal vein ratio is the straight line dis- tance between the apices of R,,, and R,,; divided by the distance between the api- ces Ol Rand Ro 4 6. M vein ratio is the straight line distance along M between crossvein dm-cu and 667 r-m divided by the distance apicad of crossvein dm-cu. The phylogenetic analysis was performed with the assistance of Hennig86 (Farris 1988), a computerized algorithm that pro- duces cladograms by parsimony. Before analysis, character data were arranged in transformation series and then polarized, primarily using outgroup procedures. Terminology for structures of the male terminalia are provided directly on the il- lustrations of R. ardeoceras and R. griseola (Figs. 22-25, 30-33). Note that the term ““postgonite,” as used in Mathis (1993), is changed to “‘clasper,” as in Zatwarnicki (1992) and Mathis (1994). This structure is part of the surstylur complex. The termi- nology is not repeated for comparable il- lustrations of other species. Although most specimens for this study, including the primary types, are in the Na- tional Museum of Natural History (USNM), additional specimens were borrowed and studied from the Academy of Natural Sci- ences of Philadelphia (ANSP), Pennsylva- nia (Dr. Jon K. Gelhaus, Mr. Don Azuma); American Museum of Natural History (AMNH), New York, New York (Dr. David A. Grimaldi); Canadian National Collec- tion (CNC), Ottawa, Canada (Dr. J. E. O’Hara, Mr. Bruce Cooper); University of Guelph (GUE), Guelph, Ontario, Canada (Dr. S. L. Marshall). ANNOTATED KEY TO GENERA OF THE TRIBE DISCOMYZINI — . Wing spotted, usually white or hyaline spots oniaidarksbackeround sees oe ree 2 — Wing hyaline or with anterior margin dark- ened but not spotted . Supra-alar setae absent tO Trypetomima de Meiere [4 species; Old World (Australasian/ Oceanian and Oriental)] — Supra-alar seta present, size of seta moderate tol well¥developedinmere yas as restntretaee ees oe 3 3. Facial setae 2, dorsal seta well developed; aris- ta bearing 7-8 dorsal rays; tibia or tarsus gen- erally lighter colored than femur .......... 4 668 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Facial setae 4, all well developed; arista bear- ing 14-16 dorsal rays; legs unicolorous, usu- ally: Gack® S2hece7. cot seer Actocetor Becker [11 species; Old World (Afrotropical and Oriental)] 4. Vein R,., basad of crossvein r-m bearing 3- 4 black setulae; tibiae yellowish white with brownibands ers. sao. Eremomusca Mathis [1 species; E. nussbaumi Mathis 1985; Old World (Palearctic); Mathis 1985] — Vein R,,,; basad of crossvein r-m bare; tibiae blackish brown, tarsi yellow with apical tar- SOmMere.GAnK as. Seo oe RA ie elas eke iS oa Guttipsilopa Wirth (subgenus Guttipsilopa) [1 species; G. (G.) haydeni Wirth 1956; New World (Neotropical)] 5. Alula well developed — Alula weakly developed, very narrow 6. Intrafrontal seta present; fronto-orbital setae 4 (anterior 2 setae proclinate, 3rd laterocli- nate, 4th lateroreclinate) ... Paratissa Coquillett [4 species; Old and New Worlds (Afro- tropical, Australasian/Oceanian, and Neotropical); Mathis 1993] — Intrafrontal seta absent; fronto-orbital setae ODEN aR st a Sealant Ne te OAM a Aare 7 7. Arista bearing 9-10 dorsal branches; supra- alar seta well developed, length subequal to postalamsetas- hee Rhysophora Cresson [4 species; New World (Nearctic and Neotropical)] — Arista bearing 3-5 dorsal branches, rarely 6; supra-alar seta moderately well developed, length about half postalar seta 8. Vein R,,, basad of crossvein r-m bearing 3- 4 black setulae; legs unicolorous; maxillary Palpusibrowne sear Hostis Cresson [1 species; H. guamensis Cresson 1945; Old World (Afrotropical and Austral- asian/Oceanian; Mathis 1993] — Vein R,,,; basad of crossvein r-m bare; legs dark with yellow tarsus, apical tarsomere dark; maxillary palpus usually pale colored ...... eee: Guttipsilopa Wirth (subgenus Nesopsilopa Mathis and Wirth) [5 species; New World (Nearctic and Neotropical); Mathis and Wirth 1977] 9. Supra-alar seta present; prescutellar acrosti- chal seta absent; presutural seta minute and poorly developed; katepisternal seta 1; facial setae 3 pairs, all small and poorly developed Bape msg Acordia Set dh tue Discomyza Meigen [9 species; Old and New Worlds (Afro- tropical, Australasian/Oceanian, Nearc- tic, and Palearctic] — Supra-alar seta greatly reduced or absent; prescutellar acrostichal seta present; presu- tural seta well developed; katepisternal setae 2, dorsal seta larger; facial setae 3 pairs, dorsal pair cruciate, ventral pair poorly developed 10 10. Legs dark with yellow tarsi, apical tarsomere dark; alula narrow and bearing long, fine setae along) margin’ 73.00 ss... Mimapsilopa Cresson [5 species; New World (Nearctic and Neotropical); Lizarralde de Grosso 1982] — Legs unicolorous; alula reduced or absent .. 11 11. Arista bearing 6-7 dorsal rays; anepisternal setae 2, ventral seta slightly less than twice length of dorsal seta; femur setulose, setae in LOWS: eee en eat ee Helaeomyia Cresson [2 species; New World (Nearctic and Neotropical); Lizarralde de Grosso 1982] — Arista bearing 8-10 dorsal rays; anepisternal setae 2, ventral seta more than twice the length of the dorsal seta; femur bearing 2-3 promi- nent setae near the apex ....Clasiopella Hendel [2 species; Old World (Afrotropical, Aus- tralasian/Oceanian, Oriental; 1 species introduced to New World (Nearctic and Neotropical); Mathis 1994] Genus Rhysophora Cresson Rhysophora Cresson, 1924: 159. Type spe- cies: Rhysophora robusta Cresson, 1924, monotypy.—Cresson, 1942: 105, 123 [key, review].—Sturtevant and Wheeler, 1954: 160 [key].— Wirth and Stone, 1956: 465 [key].—Wirth, 1965: 743 [Nearctic catalog].—Mathis, 1977: 921-945 [revi- sion]. Diagnosis.— Specimens of Rhysophora are similar to those of Guttipsilopa and other genera of Discomyzini but can be differ- entiated by the following combination of characters: outer vertical seta usually half or less length of inner seta, but never equal; proclinate fronto-orbital setae 1-2, inserted anteriad of median, reclinate seta; intra- frontal setae lacking; arista bearing 8-15 dorsal rays; face generally with prominent antennal grooves (antennal grooves in R. ardeoceras and R. griseola are very shallow); supra-alar seta well developed; wing gen- erally hyaline or faintly infuscate; vein R,;, lacking setulae basad of crossvein r-m; alula well developed; hind basitarsus of male slightly swollen, with a ventral groove that bears a row of angularly recurved, pale se- tae. VOLUME 97, NUMBER 3 669 Figs. 1-6. Rhysophora liropus. 1, Head, lateral view. 2, Same, anterior view. 3, Male genitalia, posterior view. 4, Same, lateral view. 5, Aedeagus and aedeagal apodeme, dorsal view. 6, Same, lateral view. Scale = 0.1 mm. Description. — Moderately small to mod- erately large shore flies, length 2.4 to 4.4 mm; usually blackish brown. Head: Frons wider than long; mesofrons generally triangular, with base towards ver- tex and anterior angle at ptilinal suture; tri- angle shape and size variable among spe- cies; fronto-orbits and mesofrons concol- orous; ocellar triangle slightly raised with ocelli forming equilateral or isosceles tri- angle; pseudopostocellar setae inserted close together just posteromediad of posterior ocelli; ocellar setae proclinate, well devel- oped, inserted wide apart just posterolater- ad of anterior ocellus; ocellar setae greater than twice length of pseudopostocellar se- tae; 2 poorly developed, minute setulae in- serted anteromediad of rear ocelli; intra- frontal setae lacking; inner vertical seta large, usually twice or more length of outer seta; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ostocular setulae minute, inserted poste- r of either vertical seta, extended along »ye posterior margin; fronto-orbital setae 2— 3, posterior seta reclinate, 1-2 proclinate setae inserted anteriad of reclinate seta; me- dian fronto-orbital seta often reduced or missing. Antenna oriented anterolaterad; pedicel sparsely setulose, bearing 4—6 small- er setae along ventral edge with | seta long, prominent; arista inserted on dorsal base of velvety appearing, macropubescent flagel- lomere |. Face higher than wide; presence of antennal grooves and facial carina vari- able; face usually convex in profile, bearing 2-4 inclinate facial setae along lateral mar- gin parallelling parafacials; facial setae in- serted just below midfacial height; genal seta 1, prominent, inclinate; gena and postgena setulose; occipital setae small, running along occiput. Eye higher than wide; gena-to-eye ratio less than 0.25. Mouthparts small; maxillary palpus sparsely setulose. Thorax: Generally unicolorous or light- er colored laterally; scutellum parabolic, slightly convex with disc setulose. Chaeto- taxy as follows: prescutellar acrostichal seta inserted slightly anteriad of larger, laterally displaced dorsocentral seta; postalar seta large, subequal in length to dorsocentral seta; supra-alar seta 1; apical scutellar setae 1; subapical scutellar setae weakly developed; basal scutellar seta 1; notopleural setae 2, both inserted at same level just dorsad of ventral notopleural crease; postpronotal seta 1; anepisternal setae along posterior margin 2, ventral seta longer; katepisternum lightly setulose, bearing | large katepisternal seta, usually curved upward. Legs setulose with small setae; mid femur bearing | prominent spinelike seta on posteroventral margin, lo- cated at apical '; tibia bearing 1 prominent spine at apex, especially prominent on mid leg; hind basitarsomere of male slightly swollen, bearing a row of distinctive, pale, recurved setae arising from ventral groove; coxa with | larger seta along anterior margin and a whorl of small setae, prominent on mid coxa. Wings usually hyaline to yellow- ish brown; halter yellowish white; costal vein ratio variable; M vein ratio usually close to 1; costal margin near basicosta and tegula bearing 2 larger setae; subcostal margin lined with stout, black setae and indented at vein R,; smaller, black setae along costal margin ended at vein R,,5; alula very narrow; vein R,,,; basad of crossvein r-m lacking setulae. Abdomen: Usually dark brown to black, setulose; posterior tergites slightly larger; posterior and lateral margins of tergites usu- ally bearing larger setae; 4th and Sth tergites sometimes bearing semierect, larger setae. Male terminalia: symmetrical; epandrium broadly U-shaped in posterior view, usually becoming wider subventrally in lateral view; cercus with ventral margin gradually merged with membrane; surstylus variable, greatly reduced or prominent, variously shaped de- pending on species; clasper with | or 2 pro- cesses, each longer than wide; pregonite small (could be the fused pre- and postgon- ites), usually closely associated with hypan- drium, bearing 1-2 long setulae; aedeagal apodeme L-shaped to triangular in lateral view; aedeagus variously shaped, depending on species; hypandrium concave or very shallowly depressed, nearly flat. Distribution.—The genus is known thus far only from the New World, with greater diversity in the tropics. Natural history.—Two species, R. liropus and R. robusta, are associated with plants of the family Pontederiaceae, more com- monly know as pickerel weed (species of the genus Pontederia L.) and anchored water hyacinth (species of the genus Eichhornia K. Kunth). These often abundant plants are sometimes considered weeds, and the phy- tophagous species of Rhysophora may offer some measure of biological control. Adult flies are often collected by sweeping species of Pontederiaceae. Discussion.— Although Rhysophora is well characterized and its monophyly rea- sonably well established, we remain rela- tively ignorant about its phylogenetic rela- tionships within the tribe Discomyzini. This VOLUME 97, NUMBER 3 tribe, which was only recently recharacter- ized, has received inadequate attention from the standpoint of phylogenetics, and the available information on relationships is limited to recent revisionary studies that are of limited scope and that do not impinge directly on Rhysophora (Mathis 1993, 1994). Moreover, one of the only synapomorphies that may indicate a sister group for Rhy- sophora is ambiguous. The species of Rhy- sophora, like Guttipsilopa, lack setulae on the dorsum of vein R,,,;. This is apparently a secondary reversal of a synapomorphy for the tribe (Zatwarnicki 1992). Although this character may indicate a sister-group rela- tionship between these two genera, the loss could also have arisen independently in each genus. Zatwarnicki (personal communica- tion) is now investigating these and other issues germane to the phylogeny of Disco- myzini and Psilopini, and for the present we defer to his study and analysis. We did consult with Zatwarnicki (personal com- munication), who suggested, albeit provi- sionally, that we use Helaeomyia as an out- group. Although the evidence for the monophyly of Rhysophora is rather compelling (see ‘**Phylogenetic Considerations’’), the in- cluded species, with the exception of R. J/ir- opus and R. robusta, are comparatively het- erogeneous in external features. Casual ob- servation could easily lead one to wonder if the species are indeed closely related, es- pecially R. ardeoceras, which has many ple- siomorphic external features and is quite similar to other taxa in the tribe. KEY TO SPECIES OF RHYSOPHORA CRESSON 1. Antennal grooves and facial carina weakly dif- ferentiated, grooves very shallowly impressed; ventral margin of face nearly flat; face height- to-width ratio 0.64 to 0.70; mid femur bearing 2-4 setae, including | prominent spinelike seta on posteroventral margin near apical '/; clasper AESIN PIE IPLOCESS) eee eae ieee has cus 2 — Antennal grooves and facial carina between an- tenna distinct; ventral margin of face broadly and deeply emarginate; face height-to-width 671 ratio 0.75 to 0.81; mid femur bearing | prom- inent spinelike setae on posteroventral margin about apical '3; clasper with 2 fingerlike pro- cesses, posterior process of clasper bearing a row of long, conspicuous setae along posterior INALOING AMEE Lee ae de crete tein nes 3 2. General coloration blackish brown, without whitish microtomentum; pseudopostocellar setae well developed; flagellomere | longer than wide, pointed apically; presutural seta present; costal vein ratio 0.58 to 0.65; surstylus small, a narrow, bandlike, transverse process ...... ACRE ee ee es R. ardeoceras Mathis — General coloration blackish brown with whit- ish microtomentum; pseudopostocellar setae greatly reduced or absent; flagellomere | blunt- ly rounded; presutural seta absent; costal vein ratio 0.31 to 0.35; surstylus large, conspicuous in lateral view as a bootlike process sy re steel ee Rea Oara Sr R. griseola, new species 3. General coloration black; antennal grooves very deep and distinct; ventral portion of face ru- gose; flagellomere | black; gena-to-eye ratio 0.09 to 0.11; color of tarsi sexually dimorphic: ¢ tarsi black; 2 mid and hind tarsi yellow, fore tarsusiblackishpers 0-020 «ere R. robusta Cresson — General coloration grayish tan; antennal grooves less well defined; ventral margin of face not rugose; flagellomere | yellowish orange; gena-to-eye ratio 0.20 to 0.21; tarsi of 46 and 2 concolorous, yellowish brown with apical tra- Somerciolack: sae eay. ae ae ie R. liropus Mathis Rhysophora liropus Mathis Figs. 1-6, 26 Rhysophora liropus Mathis, 1977: 931 [Ar- gentina. Buenos Aires: Campana; HT 4, USNM (75359)]. Diagnosis.—This species is closely relat- ed to R. robusta but can be distinguished by its generally grayish tan coloration; yel- lowish orange flagellomere 1; relatively smooth ventral margin of face, not rugose; wide gena (gena-to-eye ratio about 0.20); tarsi of 6 and 2 yellowish brown with apical tarsomere black. Description. —Generally light gray to tan; medium-sized to moderately large sized shore flies, length 3.6-4.2 mm. Head (Figs. 1-2): Frons width-to-length ratio 0.64—0.66; mesofrons rust brown, mi- crotomentose, forming a lighter, raised pa- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-12. Rhysophora robusta. 7, Head, lateral view. 8, Same, anterior view. 9, Male genitalia, posterior view. 10, Same, lateral view. 11, Aedeagus and aedeagal apodeme dorsal view. 12, Same, lateral view. Scale = 0.1 mm. rabola with bluntly arched apex dorsal to parabolic ptilinal suture; frons between 2 raised portions slightly darker, depressed; inner vertical seta inserted close to outer seta, length of inner seta twice or more that of reduced outer seta; fronto-orbital plate narrow, slightly raised, wider at insertions of fronto-orbital setae; fronto-orbital setae 3; reclinate dorsal fronto-orbital seta ap- proximately same length as ocellar setae; median seta slightly displaced laterally, pro- clinate and poorly developed; ventral fron- to-orbital seta proclinate and twice length of median seta. Pedicel same color as frons but darker brown at base; pedicel bearing 2 setae, | proclinate spinelike seta on anter- odorsal edge and | ventral seta slightly larg- er and better developed; flagellomere | yel- lowish orange with darker apex; flagello- mere | short, subequal to combined length of scape and pedicel. Face width-to-height ratio 0.79-0.81; face golden gray and vel- vety; facial carina distinct; face emarginate and arched along ventral margin; lunule re- cedes inward to ptilinal suture at an angle; antennal grooves prominent but not deep; facial carina broad, not clearly defined; dor- sal portion of face between antenna convex; face convex ventrally; parafacial crease clearly defined; facial setae 2-3, with 3-4 VOLUME 97, NUMBER 3 smaller setulae; minute setulae running along lateral margin of face parallel to parafacial;: eye width-to-height ratio 0.71-0.73; gena- to-eye ratio 0.20-0.21; maxillary palpus brown, lighter ventrally. Thorax: Greenish gray; dorsum with 4 dark brown lines faded anterior of trans- verse suture; | presutural seta; anepisternal setae 2; ventral seta twice as long as dorsal seta, both setae oriented slightly away from thorax. Femora and tibiae greenish gray, sometimes dark brown apically; tarsi light brown to yellowish with apical tarsomere darker brown to black; fore femur bearing 2-4 rows of long, fine setae on posterior margin. Wing brownish yellow; anterior margin relatively straight; costal vein ratio 0.61-0.63; M vein ratio 0.96—1.0; subcostal margin posterior to vein R, bearing | prom- inent seta; vein CuA, ends before fusing with margin. Halter brownish yellow. Abdomen: Grayish green to grayish brown; anterior portion of tergites 3-5 brownish; 4th sternite of male completely sclerotized, lacking a membranous center. Male terminalia (Figs. 36) as follows: epan- drium in lateral view becoming slightly wid- er ventrally with ventral margin broadly and bluntly rounded; cercus relatively long, oc- cupying most of height of cercal cavity; sur- stylus greatly reduced, as a horizontal, nar- row, bandlike process just ventrad of ven- tral margin of cerci; clasper well developed, deeply bifurcate a 2 fingerlike processes; posterior process of clasper (Figs. 3—4) nar- rowed medially, with linear dorsal lobe and spatulate ventral lobe, dorsal lobe bearing 8-10 robust, long setae along posterior mar- gin, ventral lobe bearing numerous spine- like, short setae; anterior process of clasper (Fig. 4) nearly straight, relatively wide, with sub-basal enlargement on anterior margin, apex bluntly rounded, bearing numerous short setulae on anterior surface; aedeagal apodeme (Figs. 5-6) with extended keel rounded in lateral view; aedeagus (Figs. 5— 6) about twice as long as wide, L-shaped in lateral view, with short, narrow, postero- 673 dorsal arm, larger arm narrowly rectangular in posterior view with apical margin shal- lowly mucronate; hypandrium concave, broadly rounded, wider than long, shallowly and narrowly emarginate at juncture with aedeagal apodeme. Specimens examined.—ARGENTINA. Buenos Aires: Campana, Frente all astillero (on Pontederia as pupa), A-361, Nov 5 1973 (1 6; USNM); Dique Lujan: Pupas de Dique Lujan (Eichhornia azurea), Acc. A-358c, 6 Nov 1973 (1 9; USNM). Distribution (Fig. 26).— Neotropical: Ar- gentina. Buenos Aires Province. This spe- cies is only known from the type series. Natural history.— The allotype female was collected from anchored water hyacinth (Eichhornia azurea (Sw.) Kunth), and H. A. Cordo (in litt.) reared this species from a species of Pontederia in Argentina. Rhysophora robusta Cresson Figs. 7-15, 19 Rhysophora robusta Cresson, 1924: 159 [USA. Virginia: Fairfax Co., Dyke; HT 4, USNM (56453)]; 1942: 123 [review].— Wirth, 1965: 743 [Nearctic catalog].— Mathis, 1977: 927-931 [revision]. Discocerina magna Coquillett in Johnson, 1910: 806 [nomen nudum].— Wirth, 1965: 743 [synonymy]. Diagnosis.— This species is closely relat- ed to R. /iropus but can be differentiated by its generally black coloration, including fla- gellomere 1; rugose ventral portion of the face; short gena (gena-to-eye ratio about 0.10); and coloration of tarsi sexually di- morphic: 6 black, 2 with mid and hind legs yellow, foreleg black. Description. —Generally black to brown- ish black; moderately small to moderately large shore flies, length 2.6-—4.4 mm. Head (Figs. 7-8): Frons width-to-length ratio 0.48- 0.59; frons black and indented ventrally; mesofrontal triangle lightly microtomen- tose with apex of triangle dorsad of ptilinal suture, ventral portion of triangle bearing a PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 13-18. Scanning electron micrographs of Rhysophora robusta and R. arderoceras. 13, Hind basitarsus, ventral view. 14, Hind basitarsus, lateral view. 15, Hind basitarsus, enlargement of ventral view. R. ardeoceras: 16, Hind basitarsus, ventral view. 17, Hind basitarsus, lateral view. 18, Hind basitarsus, enlargement of ventral view. slight indent; parafrons velvety, minutely microtomentose; pseudopostocellar seta well developed; inner vertical seta inserted close to outer seta, length of inner seta twice or more that of reduced outer seta; fronto-or- bital plate narrow, slightly raised, wider at insertions of fronto-orbital setae; fronto-or- bital setae 3, all inserted close together; large reclinate fronto-orbital seta inserted dor- sally, about same length as outer vertical seta; proclinate fronto-orbital setae as large as pseudopostocellar seta; median fronto- orbital seta proclinate, often reduced or very minute; ventral seta proclinate. Pedicel black, bearing 2 prominent, spinelike, sub- equal setae on dorsal portion; posterior seta lateroclinate, anterior seta proclinate; arista bearing 9-10 long, dorsal rays; flagellomere 1 bluntly rounded at apex; flagellomere | as long as of slightly smaller than combined length of scape and pedicel. Face width-to- height ratio 0.75-0.81; face black, similar to mesofrontal triangle; facial carina dis- tinct; ventral margin of face broadly emar- ginate, arched and strongly rugose; lunule recedes inward almost horizontally to ptil- inal suture; antennal grooves deep with a prominent, raised facial carina; face convex and velvety dorsal to facial carina; black parafacial crease divides velvety whitish parafacial from face; facial setae 4, inserted along lateral margin; 2-3 smaller, facial se- tae inserted anteriad to 4 well-developed, inclinate facial setae. Eye width-to-height ratio 0.65—0.71; gena-to-eye ratio 0.09-0.11; maxillary palpus black. Thorax: Generally black, unicolorous; presutural seta; anepisternal setae 2; dorsal seta less than 2 length of ventral seta, 2-4 smaller setae inserted between and near these VOLUME 97, NUMBER 3 2 seta. Legs black, concolorous; female mid and hind tarsi yellowish, with mid tarsus often brownish yellow; apical tarsomeres 1- 3 of female blackish; male mid and hind tarsus black, sometimes lighter ventrally. Wings yellowish brown but darker dorsally; anterior margin relatively straight; costal vein ratio 0.58-0.62; M vein ratio 0.82- 0.96; 1 stout spine along subcostal margin posterior to vein R,. Halter yellow. Abdomen: Generally black, subshiny; 4th and Sth tergites lacking longer, dorso- oblique setae along posterior margin; 4th sternite of male completely sclerotized, lacking a membranous center. Male termin- alia (Figs. 9-12) as follows: epandrium in lateral view becoming slightly wider ven- trally with ventral margin broadly and bluntly rounded; cercus relatively long, oc- cupying most of height of cercal cavity; sur- stylus greatly reduced, as a horizontal, nar- row, bandlike process just ventrad of ven- tral margin of cerci; clasper well developed, deeply bifurcate as 2 fingerlike processes; posterior process of clasper (Figs. 9-10) somewhat L-shaped, anterior margin an- gulate in lateral view, posterior margin con- cave, rounded in lateral view, bearing nu- merous, long setae in row along concave, posterior margin, ventral apex broadly rounded, bearing numerous, short, spine- like setae apically; anterior process of clasp- er (Fig. 10) slightly angulate to curved and tapered gradually to apex in lateral view, in posterior view digitiform, broadly rounded, parallel sided, bearing numerous, short setulae along anterior surface; aedeagal apo- deme (Figs. 1 1-12) triangular in lateral view; aedeagus (Figs. 11-12) longer than wide, L-shaped in lateral view, with posterodorsal arm shorter, roundly rectangular in poste- rior view with apical margin shallowly and broadly rounded; hypandrium concave, broadly rounded, wider than long, shallowly and widely emarginate at juncture with ae- deagal apodeme. Specimens examined.—CANADA. On- tario: Algonquin Park, 28 Jul, J. Mc- 675 Dunnough (3 6, 3 2¢; CNC, ANSP); Eagle Lake (Pontederia), 21 Jul 1976, S. L. Miller (1 6; GUE); Elgin, Hart Creek (Pontederia), 14-21 Jul 1977, S. L. Miller (1 4, 4 9; GUE); Lake Opinicon (Pontederia), 26 Jun 1977, S. L. Miller (1 4; QUE); Ottawa, 1 Jul 1958, A. L. Melander (4 6; ANSP, USNM)); Spar- row Lake (Pontederia), 19 Jul 1976, S. L. Miller (1 6; GUE). Quebec: Lac Bernard, 7 Aug 1938, G. E. Shewell (1 4, 1 2; CNC); Perkins Mills, 14 Aug 1938, G. E. Shewell (6 6, 3 2; CNC); St. Pierre de Wakefield (on flowers of Pontederia cordata L.), 28 Jul L961; J. R: Vockeroth (Sie, 110F 2 ENE USNM). UNITED STATES. Connecticut: Fairfield Co., Redding, 16-23 Jul 1930, 1932, A. L. Melander (4 4, 4 9; ANSP, USNM). District of Columbia: Analoston Island (Theodore Roosevelt Island), Laittle River (flowers of Pontederia cordata), 15 Jul 1916, H. L. Viereck (1 9; USNM). Florida: Dade Co., Royal Palm Park, 12-18 Apr 1923 (1 6; AMNH). Highlands Co., Archbold Bi- ological Station (8 km W), 15 Apr 1989, W. & D. Mathis (11 4, 10 2; USNM); Highlands Hammock State Park, 20 Mar 1954, H. V. Weems, Jr. (1 2; USNM); Venus, 4 May 1961, H. V. Weems, (1 2; USNM). Marion Cox, 10) May. 1956; H. V.. Weems Jr (12: USNM). Putnam Co., Crescent City, 20 Apr 1908, VanDuzee (1 2; AMNH). Maine: Hancock Co., Bar Harbor, 5 Jul 1930, C. W. Johnson (1 6; USNM). Maryland: Prince - Georges Co., Hyattsville; 1) Sep 1912; FE: Knab, J. R. Malloch (1 6; USNM). Massa- chusetts: Barnstable Co., Pocasset, 26 Jul 1950, A. H. Sturtevant (2 6; USNM). Plym- outh Co., Rochester, 21 Jul 1950, A. H. Sturtevant (1 6; USNM). Michigan: Mecos- ta Co., 24 Jul 1946, R. R. Dreisbach (1 2; USNM). New Jersey: Burlington Co., Riv- erton; Aug) 1917,0C.. W: Johnson *¢li rc: USNM). New Hampshire: Cheshire Co., Keene, 4 Aug 1956, A. H. Sturtevant (1 4, 2 2; USNM). New York: Richmond Co., Staten Island, 1923 (1 6; USNM). Suffolk Co., Long Island, Riverhead, 4-16 Jun 1951, Roy Latham (4 2; USNM); Long Island, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ent, 2 Sep 1954, Roy Latham (1 92; (JSNM); Long Island, Montauk, 11 Sep 1954, Roy Latham (1 6; USNM). West Nyack, 11 Jul 1920 (1 6; ANSP). Texas: Bastrop Co., Bastrop, 11 May 1954, L. D. Beamer (1 6, 1 2; USNM). Virginia: Fairfax Co., Mt. Vernon, 27 Jun 1915, W. L. McAtee (1 4, USNM); Dyke Marsh, 16 Jul 1916, W. L. McAtee (2 2; ANSP, USNM); Alexan- dria, 29 Jun 1952, W. W. Wirth (1 92; USNM). New Kent Co., Lanexa at Chick- ahominy River, 9 Jun 1984, W. E. Steiner, J. E. Lowry, A. G. Gerberich, D. S. Bogar (3 6, 1 2; USNM). Distribution (Fig. 19).—Nearctic: Cana- da (ON, QB), USA (CT, DC, FL, ME, MD, MA, MI, NH, NJ, NY, TX, VA). Remarks.—The coloration of the tarsi in this species is sexually dimorphic, with the tarsi of males being dark colored, mostly blackish, whereas the mid and hind tarsi of females are mostly yellow. Rhysophora ardeoceras Mathis Figs. 16-18, 20-26 Rhysophora ardeoceras Mathis, 1977: 936 [Costa Rica. Guanacaste: Tilaran (14 km NE); HT 4, USNM (75360)]. Diagnosis.— This species is similar to R. griseola but is distinguished by the generally blackish brown coloration; fronto-orbital setae 2 (2nd proclinate seta lacking); elon- gate and pointed flagellomere 1; well-de- veloped presutural seta; and short vein Rj, ; (costal vein ratio about 0.60). Description.— Generally blackish brown; moderately small to medium-sized shore flies, body length 2.4—3.3 mm. Head: Frons width-to-length ratio 0.50- 0.55; frons black and slightly indented ven- trally; mesofrontal triangle subshiny with slightly arched vertex at straight ptilinal su- ture; ventral portion of triangle near vertex slightly shallow and lightly indented; fron- to-orbital plate wide and slightly raised above level of triangle; dorsal fronto-orbital plate wide and fused with base of triangle; parafrons velvety and minutely microto- mentose between triangle and fronto-orbital plate, gently inclined dorsally; paler along ventral portions, bearing minute setulae; in- ner vertical seta inserted close to outer seta, relatively long, about 3 times longer than outer seta; fronto-orbital setae 2—3, inserted moderately close together; larger dorsal fronto-orbital seta reclinate, slightly larger than ocellar seta; median seta reduced or absent; ventral fronto-orbital seta procli- nate, less well developed, ' length of dorsal seta. Antenna oriented laterally; pedicel blackish brown with 2 prominent spinelike setae on dorsal surface; posterior seta small and slightly lateroclinate, '2 the length of anterior seta; anterior seta slightly smaller than enlarged ventral seta, well developed and proclinate; black arista bearing 12-15 long dorsal rays; flagellomere | yellowish orange near base, darker along margins; apex very acutely rounded, almost pointed; fla- gellomere 1 nearly twice as long as com- bined pedicel and scape. Face width-to- height ratio 0.64-0.70; face black, lighter and velvety ventrally; face lightly emargin- ate along ventral margin; lunule receded in- ward to ptilinal suture at a slight angle; an- tennal grooves very reduced or absent; fa- cial carina absent; face convex between an- tenna; face wide and convex centrally; whitish parafacials on either side of raised face sloping ventrad to eye; facial setae 3; dorsal seta well developed and cruciate; ventral pair small, reduced; line of minute setulae running dorsal to black parafacial margin; eye width-to-height ratio 0.71; gena- to-eye ratio 0.10-0.11; gena whitish and heavily setulose but darker near post gena; clypeus and clypeal membrane brown; brown maxillary palpus long and slender; mouthparts brownish but reddish brown and lighter ventrally. Thorax: Generally subshiny, blackish brown; presutural setae |, about 2 the length of supra-alar seta; anepisternal setae 2; dor- sal seta oriented along thorax, ventral seta about three times as large as dorsal seta. VOLUME 97, NUMBER 3 Fig. 19. Distribution map of Rhysophora robusta. Legs brownish black with yellowish tarsi; apical tarsomere blackish brown; mid fe- mur bearing 3-4 prominent spinelike setae on posteroventral margin with | usually very well developed. Wings infumate and yel- lowish brown; costal vein ratio 0.58—0.65; M vein ratio 0.85—0.88; subcostal margin posterior to vein R, bearing | stout spine. Halter pale yellow. 677 Abdomen: Generally dark brown to black, subshiny; tergites 4 and 5 bearing several large dorso-obliquely oriented setae along posterior margin; 4th sternite of male with a membranous oval in center. Male terminalia (Figs. 22-25) as follows: epan- drium in lateral view parallel sided, slightly tapered on ventral ‘4 to rounded ventral margin; cerci relatively short, occupying 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 21 eagus aedeagal apodeme Figs. 20-25. \ epandrium ie KH \ cercus SAS N Tin eee % clasper—~~\_( clasper hypandrium Rhysophora ardeoceras. 20, Head, lateral view. 21, Same, anterior view. 22, Male genitalia, posterior view. 23, Same, lateral view. 24, Aedeagus and aedeagal apodeme, dorsal view. 25, Same, lateral view. Scale = 0.1 mm. about half height of cercal cavity; surstylus greatly reduced, horizontal, narrow, band- like process, situated just ventrad of cerci; clasper well developed (Figs. 22-23) rela- tively complex, bilobed with small cleft pos- teriorly, smaller dorsal lobe truncate pos- teriorly with posterior margin bearing nu- merous, short, spinelike setae; ventral lobe of clasper much larger, rounded anteriorly and ventrally, posterior margin truncate, with dorsal half bear several short, pale setulae in a comblike row, also bearing few setulae at posteroventral angle, especially along median surface; pregonite barlike, bearing | long seta, inserted medially; ae- deagal apodeme (Figs. 24—25) relatively flat, lacking a conspicuous keel but with long lateral extensions; aedeagus (Figs. 24-25) tubular with dorsal slit, longer than wide, broadly rounded apically, with an apical, curved, narrow process oriented postero- ventrally, aedeagus in posterior view nar- row, helmet shaped dorsally; hypandrium concave, broadly rounded, wider than long, shallowly and widely emarginate with a short projection at apex of each side of emargi- nation, bearing a median, short keel along midline on internal surface. Specimens examined.—COLOMBIA. Buenaventura, 2 Nov 1950, A. E. Michel- VOLUME 97, NUMBER 3 bacher, E. S. Ross (2 6; USNM). COSTA RICA. Guanacaste: Tilaran (14 km NE), 5 jun 1973, 1. c. Erwin, G. H. Hevel (2 2, 1 2; USNM). EL SALVADOR. Santa Tecla (12 km NW; also known as Nueva San Sal- vador), Oct 1953, W. B. Heed (1 6; USNM). MEXICO. Jalisco: Barranquillas, 3 Feb 1964, E. I. Schlinger (1 2; USNM). Chiapas: Rio Izapa (near Tapachula), 21 Apr 1983, W. N. Mathis (1 4, 1 9; USNM). SURI- NAM. Paramaribo, Feb 1968, F. D. Ben- nett, H. Zwolfer (1 2; USNM). Distribution (Fig. 26).— Neotropical: Su- riname and Mexico (CHI, JAL, TEP) south through El Salvador and Costa Rica to Co- lombia. Remarks. — Externally this species differs rather markedly from its congeners (see di- agnosis and key). Despite its heterogeneity, itis apparently the sister group of the lineage giving rise to R. liropus and R. robusta (see “Phylogeny Considerations’’). Rhysophora griseola Rao and Mathis, NEw SPECIES Figs. 26-33 Diagnosis.—Specimens of R. griseola are similar to those of R. ardeoceras but are distinguished by the generally blackish brown coloration with some whitish micro- tomentum; short and slightly rounded fla- gellomere 1; presutural seta lacking; and vein R,,,; long (costal vein ratio about 0.33). Description.— Generally grayish black to black; medium-sized shore flies, length 3.3 to 3.9 mm. Head: Frons width-to-length ratio 0.48- 0.49; black frons deeply indented ventrally; mesofrons triangular, subshiny black with rounded vertex at crescent-shaped ptilinal suture; ventral portion of triangle near ver- tex shallow and indented: fronto-orbital plate wide and raised above mesofrons; dor- sal portion of fronto-orbital plate wider, fused with base of mesofrons; parafrons ap- pearing velvety and minutely microtomen- tose between mesofrons and fronto-orbital plate; pseudopostocellar seta greatly re- 679 duced or lacking (a pair of narrowly latero- proclinate setulae present, subequal in length to setulae within ocellar triangle); in- ner vertical seta inserted moderately close to outer vertical seta; outer vertical seta rel- atively long, about 74 length of inner seta: fronto-orbital setae 3; reclinate fronto-or- bital seta slightly shorter than outer vertical seta; median proclinate fronto-orbital seta minute and inserted slightly laterad; ventral proclinate fronto-orbital seta about '2 length of reclinate seta. Antenna extended outward laterally; pedicel black with 2 prominent spinelike setae, posterior seta smaller, slightly lateroclinate, stronger anterior seta proclinate; arista black, bearing 8—10 dorsal rays; flagellomere | lighter along margins with apex bluntly rounded; flagellomere 1 slightly longer than combined length of scape and pedicel. Face width-to-height ratio 0.58- 0.60; face with whitish microtomentum ventrally, dorsal areas near lunule and sometimes central areas blackish brown and microtomentose; ventral margin of face shallowly emarginate but mostly flat; lunule flat, receded inward to ptilinal suture at a slight angle; antennal groves very reduced or absent; facial carina absent; face slightly convex between antenna; face broadly con- vex centrally; white parafacials slanted ven- trad, paralleling eye on either side of raised face, bearing setulae, becoming more dense- ly, whitish microtomentose ventrally; facial setae 2-5, sometimes asymmetrical and dis- placed dorsally; eye width-to-height ratio 0.67-0.70; gena-to-eye ratio 0.11-0.15; gena with whitish microtomentose, especially along posterior margin; clypeus and clypeal membrane swollen; maxillary palpus black- ish brown. Thorax (Fig. 29): Blackish brown; scu- tellum with lighter triangular area, base at scutoscutellar suture and vertex near scu- tellar apex; pleurae grayish and velvety; anepisternum, katepisternum, and parts of notopleuron covered by white microtomen- tum; presutural setae lacking; prescutellar acrostichal setae about half that of dorso- 680 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 26. Distribution map of RAysophora liropus (diamond), R. ardeoceras (triangle), and R. griseola (dots). central setae; notopleuron bearing 2 small- almost yellowish along ventral margin; mid er, weakly developed setulae dorsally. Legs femur bearing 3—6 prominent spinelike se- blackish brown; fore coxa with whitish mi- tae along posteroventral margin, usually crotomentum; microtomentum of fore fe- with 1 seta enlarged. Wing with anterior mur and tibia lightly whitish; tarsi lighter, margin relatively flat; yellowish brown and VOLUME 97, NUMBER 3 681 Ly AY Wy \ A AN ) ow) wep pee us ra) oes ONSe teat sat i) Avi, UNE SAM) ab Figs. 27-29. Rhysophora griseola. 27, Head, anterior view. 28, Same, lateral view. 29, Thorax, dorsal view. Scale = 0.5 mm. ‘lightly darker dorsally; halter white; costal n ratio 0.31-0.35; M vein ratio 0.94—- 38: vein CuA, ends before margin. Abdomen: Generally dark brown, sub- shiny; 4th sternite of male completely scler- otized, lacking a membranous center. Male terminalia (Figs. 30-33) as follows: epan- drium in lateral view becoming slightly wid- er subventrally, broadly and bluntly round- ed ventrally, bearing a prominent seta along posterior margin just below midheight; cer- ci relatively long, occupying most of height of cercal cavity; surstylus well developed (Figs. 30-31) essentially bare of setae, shape angulate at about a right angle with ventral footlike portion enlarged and oriented pos- terolaterally, with a small, pointed tooth on mediodorsal surface; pregonite papillalike, bearing 2 long, apical setulae; clasper (Fig. 31) bare of setae, extended anteroventrally, tapered ventrally with apical 4 enlarged, curved posteroventrally, and pointed api- cally; aedeagal apodeme (Figs. 32-33); ae- deagus (Figs. 32-33) as wide as long, apical margin with 3 points, 2 lateral projections short, median point more prominent in pos- terior view; hypandrium very shallowly ~ concave, appearing almost flat. Type material.—The holotype male is la- beled ““COSTA RICA[.] Guanacaste Prov. Colorado[,] 31 March 1988/W. E. Steiner[,] J. M. Hillf,] J. M. Swearingen[,] J. M. Mitch- ell/HOLOTYPE ¢ Rhysophora griseola Rao & Mathis USNM [red; species name and gender symbol handwritten].”’ The allotype female and 14 other paratypes (3 4, 11 2; USNM) bear the same label data as the ho- lotype. Other paratypes are as follows: MEXICO. Chiapas: Rio Izapa (near Tapa- chula), 21 Apr 1983, W. N. Mathis (1 4, 1 2; USNM). The holotype is double mounted (paper triangle), is in good condition, and is deposited in the USNM. Other specimens examined.—ECUA- DOR. H. A. Parrish, 1914 (2 2; USNM). VENEZUELA. Anzoategue: Bergantin swamp near creek (river rocks), 7 May 1988, S. A. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Marshall (2 6; GUE). Federal District: Ca- racas, 3 Sep 1942, D. A. Iriarte (1 ¢; USNM). Distribution (Fig. 26).—Neotropical: Mexico (CHI) south through Costa Rica to Venezuela and Ecuador. Etymology.—The specific epithet, griseo- la, is a Latinized adjective that refers to the whitish gray microtomentum, especially on the lateral margins of the face, post gena, and fore coxa. Remarks.—This species is the basal lin- eage of Rhysophora according to the results of our phylogenetic analysis, differing from the remaining species in several characters. PHYLOGENETIC CONSIDERATIONS Rhysophora is one of 11 genera that com- prise the tribe Discomyzini (Zatwarnicki 1992: Clanoneurum Becker, which was in- cluded in Discomyzini, has been returned to Psilopini (Zatrwarnicki, personal com- munication)). Potential outgroups for phy- logenetic analysis and the search for a sister group for Rhysophora were limited to the other 10 genera. We used four outgroups in the various analyses (Guttipsilopa, Helaeo- myia, Discomyza, and Mimapsilopa,) and in the final analysis decided upon the former two, in part based on Zatwarnicki’s rec- ommendation (personal communication). These two genera are similar and probably closely related to Rhysophora. In our anal- ysis, Guttipsilopa is indicated to be the sister group of Rhysophora, although that rela- tionship must be considered tentative, as the only supportive synapomorphy is the secondary loss of setulae on vein R,,; which may be homoplasious. Regardless of the outgroup used, however, our analysis and the topology of the cladogram indicate that Rhysophora is monophyletic. Before proceeding with the analysis of species relationships within Rhysophora, it is important to establish the monophyly of the genus. Unlike the unresolved sister- group relationships, the monophyly of Rhy- sophora is well established with synapo- VOLUME 97, NUMBER 3 surstylus 32 Figs. 30-33. hypandrium aedeagal apodeme ie. 683 4 @~ surstylus clasper 33 7—— aedeagus— Rhysophora griseola. 30, Male genitalia, posterior view. 31, Same, lateral view. 32, Aedeagus and aedeagal apodeme, dorsal view. 33, Same, lateral view. Scale = 0.1 mm. morphies as follows: hind basitarsus slightly swollen; hind basitarsus of male with a ven- tral groove that bears a distinctive row of pale, recurved setulae; outer vertical seta shorter than inner seta, usually less than one-half the length of the latter (R. griseola is an exception, with the outer seta about *4 the length of the inner seta); and pseudo- postocellar setulae lateroreclinate. With the monophyly of Rhysophora established, we now focus on the phylogenetic relationships among the included species. In the presentation on species relation- ships that follows, the characters used in the analysis are noted first. Each character is immediately followed by a discussion of its states or any qualifying comments. After the presentation of information on characters, an hypothesis of the cladistic relationships is presented and discussed. The cladograms (Figs. 34-35) are the primary mode for con- veying these relationships; the discussion 1s a supplement to the cladogram and is in- tended only to complement the latter. In the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON HH HH i HH ices Lr ancestor Helaeomyia Gluttipsilopa R. ardeoceras +_HlK Hah R. griseola R. liropus R. robusta Length of 23 steps; Consistency Index 0.86; Retention Index 0.88. Charactems, =. 2 38 7445 6G! 7 8 \9e 10 Tia iSs aloe slomsig, Sicps: eae. ge vod SS ele eee Cl: 50 100 100 100 100 100 100 100 100 100 100 100 100 50100100 50 RI: 50 100 100 100 100 100 100 100 100 100 100 100 100 50100100 50 34 ancestor Helaeomyia Gluttipsilopa R. griseola R. ardeoceras R. liropus | | 12°73 45° 6 07 (8° 9 TOM 12°13 14) 15 16 17 18) 19 720°2122523 24 IP R. robusta Length of 23 steps; Consistency Index 0.86; Retention Index 0.88. Gharacter® “1 32: eS 74) 95 60 27 Bis STOP Ta eal2a Sie Saale Stepsii: ©. ab 2 We nSw eg 2a thege 2, Ghali lg le 2 ieee ee Cl: 100 100 100 66 50100100100 100 100 100 100 100 50 100 100 100 RI: 100 100 100 50 50100100100 100 100 100 100 100 50 100 100 100 35 Figs. 34-35. Cladograms depicting hypothetical phylogenetic relationships among species of the genus Rhy- sophora (with analysis and statistics of cladogram and characters). 34, With R. griseola as the sister group to R. liropus and R. robusta. 35, With R. ardeoceras as the sister group to R. liropus and R. robusta. CI = Consistency Index, RI = Retention Index. discussion of character data, a “0” indicates derived states. The coding for nonadditive the more plesiomorphic state (blank bars on the cladograms), with “1” (black bars on the cladogram) and ‘‘2” (hatched bars on the cladograms) indicating respectively more characters is reviewed on a character by character basis as indicated in the text. The numbers used in the presentation are the same as those on the cladogram (Fig. 34), VOLUME 97, NUMBER 3 and the sequence is the same as noted in the character matrix (Table 1). CHARACTERS USED IN THE PHYLOGENETIC ANALYSIS Head li: Comparative lengths of inner and outer vertical setae: (0) vertical setae subequal in length, frequently with the outer seta slightly shorter; (1) outer vertical seta much shorter than inner seta, usually half or less. In most shore flies, including members of Discomyzini, the outer seta is slightly shorter than the inner seta, and the greatly reduced outer seta, as found in three of the species of Rhysophora, represents a synapomorphy. Orientation of pseudopostocellar setae: (0) lateroclinate; (1) lateroreclinate; (2) lateroproclinate. The more typical con- dition found in Discomyzini and Psilo- pini is for the pseudopostocellar setae to be reduced in length and widely latero- clinate. In species of Rhysophora, these setae are usually well developed, and with a narrowly lateroclinate orientation. In specimens of R. griseola these setae are greatly reduced, with a slightly diver- gent, proclinate orientation, or lacking. . Shape of ptilinal suture: (0) somewhat flattened medially with curved lateral margins; (1) strongly arched, almost crescent shaped. Shape of flagellomere 1: (0) apex nar- rowly pointed; (1) apex rounded; (2) apex bluntly rounded. The first flagellomere is generally conspicuously pointed, as found in R. ardeoceras. The rounded and bluntly rounded states are synapomor- phies. Lacking an indication as to the transformation series for this character, we treated it as a nonadditive character. . Comparative lengths of flagellomere 1 and combined length of the scape and pedicel: (0) flagellomere | larger than the combined length of the scape and pedi- cel; (1) flagellomere 1 equal or slightly Table 1. 685 Matrix of characters and taxa used in the phylogenetic analysis (numbers for characters corre- spond with those used in text). Characters Taxa 12345 67890 12345 67 Ancestor 00000 00000 00000 00 Helaeomyia 00000 00000 01000 00 Guttipsilopa 00010 00000 02000 00 R. ardeoceras 11100 00011 02010 O1 R. griseola 02121 01011 02000 00 R. liropus A TEN Seat R. robusta WHT bo WPA itil shorter than the combined length of the scape and pedicel. . Antennal grooves and facial carina: (0) weakly developed or absent; (1) distinct, with relatively deep grooves and a well- defined carina. The generalized condi- tion in the tribe is either a relatively flat face or with very shallow antennal grooves. The relatively deeply formed grooves of R. /iropus and R. robusta are a synapomorphy. . Size and orientation of dorsal facial se- tae: (0) setae large, at least dorsalmost pair cruciate; (1) setae reduced in size, usually not cruciate. The generalized condition is for one, frequently the dor- salmost seta, or more of the facial setae to be cruciate. Although the facial setae in most species of Rhysophora are incli- nate, they usually do not cross. . Shape of ventral facial margin: (0) nearly flat or very shallowly emarginate; (1) deeply emarginate. Thorax 2 10. Row of distinctive, pale, curved setae inserted along anteroventral surface of male hind basitarsus: (0) setae absent; (1) setae present. This is a sexually di- morphic character and one of the pri- mary synapomorphies to indicate the monophyly of the genus. Size of male hind tarsus: (0) hind tarsus equal in size to mid and fore tarsi; (1) 686 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON hind tasu slightly swollen. Like the pre- ceding character, this is a synapomor- phy of the genus. |i. Spinelike setae along posteroventral margin of mid femur: (0) mid femur bearing 3-6 prominent setae; (1) mid femur bearing | prominent seta. 12. Setulae on dorsum of vein R,, ; basad of crossvein r-m: (0) setulae absent; (1) 3-4 setulae present; (2) setulae second- arily lost. The presence of setulae on the base of R,,, 1s a synapomorphy for the tribe, and Zatwarnicki (1992) has interpreted their absence in Rhyso- phora and Guttipsilopa to be a second- ary development. 13. Anterior wing margin: (0) shallowly arched, giving the overall appearance of a somewhat ovate wing; (1) some- what flattened with anterior and pos- terior margins largely parallel. Abdomen 14. Shape of surstylus: (0) well developed, although shape variable; (1) greatly re- duced as a horizontal, narrow, bandlike process near the ventral margin of the cercl. 15. Shape of clasper: (0) simple, primarily a single process; (1) bifurcate, frequent- ly deeply, resulting in 2 processes. 16. Setation of posterior process of the clasper: (0) a few setulae and at most a few setae present; (1) a row of long, con- spicuous setae along posterior margin. The derived state is apparently unique to R. liropus and R. robusta. 17. Shape and setation of pregonite: (0) papillalike, bearing 2 long, apical setu- lae; (1) barlike, bearing | long seta, in- serted medially. AUTAPOMORPHIES Not USED IN THE PHYLOGENETIC ANALYSIS 18. Development of pseudopostocellar se- tae: (0) pseudopostocellar setae well de- veloped; (1) pseudopostocellar setae re- duced and poorly developed. The de- rived state is unique to R. griseola. 19. Sculpturing of ventral portion of face: (0) similar to dorsal portions of face, generally relatively smooth; (1) rugose. 20. Presutural seta: (0) strongly developed; (1) seta absent. Most members of the tribe have a well developed presutural seta, and its apparent absence (or great reduction) in specimens of R. griseola is unique to that species. 21. Tarsal coloration: (0) males and fe- males similarly colored; (1) sexually di- morphic, female mid and hind tarsi yel- low, male mid and hind tarsi dark brown, concolorous with tarsus of fore- leg. 22. Setae along posterior margin of tergites 4 & 5: (1) setae oriented along abdo- men; (1) setae dorso-oblique. 23. Fourth male sternum: (0) 4th sternum entirely and evenly sclerotized; (1) 4th sternum with median, oval, membra- nous area. 24. Dorsal rays of arista: (0) 8-10 dorsal rays; (1) 12-15 dorsal rays. Table | is the character matrix. From this matrix and using the implicit enumeration (ie) option of Hennig86, two trees (Fig. 34) of equal, ““most parsimonious” length re- sulted (length of 23 steps, overall consisten- cy index of 0.86, and a retention index of 0.88). Using the successive weighing tech- nique (xsteps w, ie, cc), we further analyzed the characters to determine the goodness of fit for the two tree topologies. After succes- siving weighing the same two trees resulted, and each character had a weight of 10. Other statistics of each character are cited with the cladograms (Figs. 34, 35). The cladograms for the species of Rhy- sophora indicate that the genus is mono- phyletic (Synapomorphies 2, 3, 9, 10, and 12). They also corroborate the monophyly of R. robusta and R. liropus (synapomor- phies 6,°8, I; 12, 13; 1d; and 16)! which was formerly recognized as a species group, VOLUME 97, NUMBER 3 as a well-established monophyletic group that is the most derived lineage within the genus. Within the latter lineage, the species R. robusta is characterized by two autapo- morphies (19, 21). Our analysis indicates that the sister group giving rise to R. Jiropus and R. robusta is ambiguous, with either R. ardeoceras (syn- apomorphies 1, 14, 17), or R. griseola (syn- apomorphies 5, 7) being equal candidates. Of the two possibilities, we advocate R. ar- deoceras (Fig. 34) as the better choice. The synapomorphies that demonstrate this re- lationship are based on structures of the male genitalia and are more reliable in our opin- ion than the shape of flagellomere 1, which is known to vary somewhat within the sub- family Discomyzinae. The species R. ar- deoceras is distinguished by three autapo- morphies (22, 23, and 24), and R. griseola is distinguished by two autapomorphies (18, 20). ACKNOWLEDGMENTS We thank Elaine R. S. Hodges for ren- dering the head, thorax, and genitalic illus- trations of R. griseola, and for touching up the other illustrations, and George L. Ven- able for producing the final version of the cladograms. For reviewing an early draft of this paper, we thank Tadeusz Zatwarnick1, Amnon Freidberg, Allen L. Norrbom, and Norman E. Woodley. T. Zatwarnicki was particularly helpful in interpreting the struc- tures of the male terminalia. To our col- leagues and their institutions (listed previ- ously) who loaned specimens, we express our sincere thanks. LITERATURE CITED Cresson, E. T., Jr. 1924. Descriptions of new genera and species of the dipterous family Ephydridae. Paper VI. Entomological News 35(5): 159-164. 1942. Synopses of North American Ephyd- ridae (Diptera). I. The Subfamily Psilopinae, with descriptions of new species. Transactions of the American Entomological Society 68: 101-128. 1945. A systematic annotated arrangement of the genera and species of the Indo-Australian 687 Ephydridae (Diptera). I. The subfamily Psilopi- nae. Transactions of the American Entomological Society 71: 47-75. 1946. A systematic annotated arrangement of the genera and species of the Neotropical Ephydridae (Diptera). I. The subfamily Psilopi- nae. Transactions of the American Entomological Society 71: 129-163. Farris, J. S. 1988. Hennig86, Version 1.5. Port Jef- ferson Station, New York. Johnson, C. W. 1910. Order Diptera, pp. 703-814. In Smith, J. B., ed. The Insects of New Jersey. New York State Museum, Annual Report 1909: 11-888. Lizarralde de Grosso, M.S. 1982. Redescription del géenero Helaeomyia Cresson y reivindicacion de Mimapsilopa Cresson (Diptera, Ephydridae). Physis (Buenos Aires) 40(99): 121-128. Mathis, W. N. 1977. A Revision of the genus Rhy- sophora Cresson with a key to related genera (Dip- tera: Ephydridae). Proceedings of the Biological Society of Washington 90(4): 921-945, 29 figures, 1 table. . 1985. Anew psilopine genus and species from Israel with a recharacterization of the tribe and key to other psilopine genera of the Middle East (Diptera: Ephydridae). Proceedings of the Ento- mological Society of Washington 87(2): 375-380. . 1986. Studies of Psilopinae (Diptera: Ephyd- ridae), I: A revision of the shore fly genus Placop- sidella Kertész. Smithsonian Contributions to Zo- ology 430: iv + 30 pp. . 1993. A Revision of the shore-fly genera Hos- tis Cresson and Paratissa Coquillett (Diptera: Ephydridae). Proceedings of the Entomological Society of Washington 95(1): 21-47. 1994. A revision of the genus Clasiopella Hendel (Diptera: Ephydridae). Proceedings of the Entomological Society of Washington 96(3): 454— 465. Mathis, W. N. and W. W. Wirth. 1977. A new genus of Psilopine flies (Diptera: Ephydridae) with notes on its relationships. Proceedings of the Entomo- logical Society of Washington 79(1): 63-74. Mathis, W. N. and T. Zatwarnicki. 1990. Taxonomic notes on Ephydridae (Diptera). Proceedings of the Biological Society of Washington 103(4): 891-906. McAlpine, J. F. 1981. Morphology and terminology- adults, pp. 9-63. In McAlpine, J. F., et al., eds., Manual of Nearctic Diptera, Vol. 1. Ottawa, vi + 674 pp. [Volume | is Monograph 27 of Research Branch Agriculture Canada.] Sturtevant, A. H. and M. R. Wheeler. 1954. Synopses of Nearctic Ephydridae (Diptera). Transactions of the American Entomological Society 79: 151-257. Wirth, W. W. 1956. The Ephydridae (Diptera) of the Bahama Islands. American Museum Novitates 1817: 1-20. 588 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1965. Family Ephydridae, pp. 734-759. In Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson, eds., A Catalog of the Diptera North of Mexico. U. S. D. A. Handbook, 276, Washington, D.C., 1696 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. Depar- tamento de Zoologia, Secretaria de Agricultura, Sado Paulo. Wirth, W. W.andA. Stone. 1956. Chapter 14. Aquat- ic Diptera, pp. 372-482. Jn Usinger, R. L., ed., Aquatic Insects of California, With Keys to North American Genera and California Species. Uni- versity of California, Berkeley, viii + 508 pp. Zatwarnicki, T. 1992. A new classification of Ephyd- ridae based on phylogenetic reconstruction (Dip- tera: Cyclorrhapha). Genus 3(2): 65-119. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 689-694 A NEW SPECIES OF TOWNSENDIA WILLISTON (DIPTERA: ASILIDAE) FROM FLORIDA WITH NOTES ON ITS ASSOCIATION WITH PERDITA GRAENICHERI TIMBERLAKE (HYMENOPTERA: ANDRENIDAE) AUBREY G. SCARBROUGH, BETH B. NORDEN, AND KARL V. KROMBEIN (AGS) Department of Biological Sciences, Towson State University, Towson, Maryland 21204; (BBN, KVK) Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. Abstract.—A new species of asilid fly belonging to the genus Townsendia is described and figured. Flies were collected in association with a species of ground-nesting bee, Perdita graenicheri Timberlake at the Archbold Biological Station, Highland County, Florida. Behavioral observations of flies and speculation on their occurence near bee burrows are given. Key Words: A detailed study of the small (4-5 mm), gregarious, ground-nesting bee, Perdita graenicheri Timberlake, was conducted at the Archbold Biological Station (ABS), Highlands County, Florida in 1989 (Norden et al. 1992). During that study, a very small species of Townsendia (Diptera: Asilidae) was often observed perching on vegetation near bee nest openings. The flies appeared to be watching the bees as they entered or exited their burrows. Specimens of the fly were collected, and later were determined to be a new species. During a second trip to the Station, 10-17 August 1992, additional flies were collected and observations were made on their behavior. MATERIALS AND METHODS Study site.—Nesting Perdita graenicheri were found at several ABS sites. Deyrup (pers. comm.) noted that Townsendia ar- enicola, n. sp. is the most common asilid in the scrub and nearby open xeric areas. We found that, the fly was often abundant in Asilidae, Andrenidae, Townsendia, Perdita, ground-nesting association with the bees on the sunny, ex- posed sand at the northeastern end of Lake Annie. All of our (BNN & KVK) observa- tions and collections were made at this site. Behavioral observations.—Flies were found only on sunny days and were ob- served throughout the day (0800-1700 h, EST) during both still and breezy condi- tions. Sand surface temperatures ranged be- tween 38-52°C. Notes were taken of perch selection, behavior of the flies at perches and flight behavior. Selected blades of grass and twigs (perches) were occasionally re- moved from the area and the subsequent changes in asilid behavior were recorded. Observation time totaled ca. five hours and included approximately 30 different flies. RESULTS Flies used a series of perches having one or more bee nest openings located roughly in their center. Flights from these perches were short (ca. 5-15 cm) in distance, and about 1-5 cm above the sand. Flights were 690 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON »re hovering than darting in nature. Flies ways positioned themselves atop their perches in a horizontal position with the head facing in the direction of a bee nest. Removal of perches seemed to temporarily confuse flies who would attempt to land where a perch had previously been located. Only occasionally would flies land on the sand near bee nest openings. Movement be- tween perches appeared to increase during particularly windy days. Since flies were much smaller than the stockier bees, we doubted that adults preyed on bees but we watched closely for any for- aging behavior. We observed no obvious attempt by the flies to chase or to attack bees or other insects in the vicinity. How- ever, one specimen in the ABS collection was pinned with its presumptive prey, a spe- cies of frit fly (Diptera: Chloropidae). Small ants were occasionally followed for a short distance from the perch, as were other 7. arenicola that flew near a perched individ- ual. Most of the flies observed were females and they appeared to be watching the Per- dita burrow openings. They turned their heads from side to side, as if to scan the area around the burrow. In 1992, two fe- males were observed entering burrows only seconds after the female bees exited. Since little is known about the biology of these flies, we (Norden et al. 1992) suggested that these fly larvae might feed upon the bee larvae and that adult flies were entering nests to oviposit. Though information on the di- etary habits of asilid larvae are sparse (Woods 1981), observations suggest that they are predaceous on soil-dwelling in- sects. However, the bees were still provisioning their nests during these observations, and bee larvae probably were not present yet. Melin (1923) suggested that some asilid lar- vae may feed on non-insect diets. If this is so then a pollen mass of P. graenicheri could provide sufficient food for one or more 7. arenicola larvae. It is also plausible that the larvae are general opportunists, feeding on both pollen and bee larvae. Though we are uncertain of the signifi- cance of the association of 7. arenicola with bee burrows, three other asilids sometimes frequent vertebrate burrows. Lavigne (1968) reported (as Asilus) Machimus formosus (Hine) and M. gilvipes (Hine) in badger and ground squirrel burrows, and Bullington and Beck (1991) reported M. polyphemi Bul- lington & Beck in gopher tortoise burrows. Large burrows of such vertebrates probably serve as important retreats from harsh en- vironmental conditions, oviposition and larval developmental sites, and/or mating sites for these flies. Mating pairs of M. gil- vipes often alight and rest on the walls of burrows (Lavigne 1968). Perhaps females of T. arenicola select perches near burrows or other land marks to attract mates. Townsendia Williston Townsendia Williston, 1895: 107. Town- sendia minuta Williston. Type Locality, Mexico. Diagnosis.—Small flies, 3-4 mm long, head much wider than high, broad in lateral view; eyes with anterior facets unusually large; face narrow, vertex shallow and wide, front strongly divergent; wings basally nar- row with only four posterior cells, anal angle and alulae absent, and vein M; absent with only four posterior cells. The genus Jownsendia Williston is re- stricted to the New World, with eight spe- cies from Mexico (Martin 1966), two from the United States (Williston 1895, Back 1909), and one from Puerto Rico (Curran 1926). Both species found in the United States were described from a single female (Back 1909). The type locality for 7. nigra Back is New Jersey and that for 7. pul- cherrima Back is Texas. Martin (1966) re- ferred to two undescribed species from the United States, one from Arizona and the VOLUME 97, NUMBER 3 hd i ] rE YY Va ye / “ —— pe a Pa f- ith. Peg a fh ; es) b Nas ry ‘Ja 1 igeale 2 Right antenna of male Townsendia arenicola, n. sp., a. dorsal view, b. lateral view. Scale = 0.2 mm. Fig. 2. Right wing of Townsendia arenicola, n. sp. Scale = 0.4 mm. other from Florida. The following new spe- cies from Florida is probably the one re- ferred to by Martin. DESCRIPTION OF SPECIES Townsendia arenicola Scarbrough, NEw SPECIES Figs. 1-5 Male.— Head and thorax dark brown to black; abdomen dark brown with a slight tint of orange. Length, body 2.7-3.4 mm; wing 2.1—-2.4 mm. Face, front, and occiput gray to slightly yellowish-gray tomentose; a small, light brownish tomentose spot pres- ent adjacent to each posterior ocellus. Oral margin of face with a row of six to seven short, thick, yellowish bristles. Palpus and short proboscis with short, fine yellowish vestiture. Front narrow basally, wide above with strongly divergent sides, one to two dark brown setae present; basally, three long grooves present, with the outer two diver- gent beyond the sides of ocellar tubercle, the middle groove about twice as deep as the other two. Antenna with brown vestiture, basal two segments combined about two- thirds as long as flagellum; flagellum flat lat- erally with a median constriction anteriorly, basal width slightly greater than apical width, ventral margin slightly narrowed to apex; stylus one-half as long as and two-thirds as wide as flagellum, with a short, lateroapical spine, the latter one-fourth to one-third as long as stylus. Vertex shallow, three to four times wider than front basally. Occiput dor- sally with six to seven short, brown post- ocular bristles, laterally and ventrally with shorter, thinner, yellow bristles. Thorax mostly gray to slightly yellowish gray tomentose; scutum medially with a wide brown stripe that extends posteriorly to su- pra-alar bristles. A large lateral brown spot and sometimes a much smaller, less distinct brownish tomentose spot present on each side of median stripe. Vestiture on thoracic dorsum sparse, of short, thin, brown bris- tles; posteriorly dorsum with one, rarely two pairs of long dorsocentral bristles and three pairs of thick lateral bristles. Scutellum flat, margin with four to six long, brown setae, each alternating with one to two shorter se- tae. Pleuron usually with two thick, brown tergosternal bristles, sometimes a weaker pale seta also present. Halter basally brown, knob entirely and stalk partly yellowish. Wing iridescent, often largely yellowish, with dense microtrichiae. Veins R, and R, widely divergent with R, ending before and R, well behind wing apex. Apically cell d and basally cell m, unusually narrow; cell m, at wing margin contrastingly wide, with veins M, and M, strongly divergent. Coxae usually dark brown, sometimes PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mii apices narrowly yellowish, with yellowish vestiture. Trochanters, fore- and midfe- mora, fore- and midtibiae, and basal three to four tarsomeres of all tarsi yellow to slightly brownish yellow; fore- and midfe- mora ventrally with a long, thin, yellowish seta, seta below forefemora thickest and longest; all femora dorsoapically with two to three brown bristles. Hindfemora and hindtibiae largely yellow-brown to brown, usually yellow basally; hindfemora only slightly clavate with several long, thin, brownish or yellowish setae posteroven- trally. Tibiae and tarsi with several thick, yellowish bristles; hindtibiae strongly cla- vate, sometimes posteriorly with two or three brown bristles. Apical segment of each tarsus brown; basotarsomere of hind tarsi swollen, remaining tarsomeres more slen- der: Abdomen shiny, basal two and apical two to three segments dark reddish brown or orangish brown, with mostly brown vesti- ture; tergite 1 mostly and tergites 2-6 api- cally with a dense gray tomentose band, each band in succession from tergite 2 to 6 de- creasing in width, making band on tergite 2 the widest and that on tergite 6 narrowest; sternites sparsely gray tomentose. Sternite 8 apically with abundant black setae. Terminalia dark brown to black. Epan- drium undivided dorsally, sides dome-like and covering most of terminalia laterally, apical corners strongly pointed. Gonostylus flat, long, digitate. Hypandrium medially with a deep emargination and abundant, short, pale setae. Aedeagus as in Figure Id. Female.— Differs from the male as fol- lows. Length, body 3.2-4.3 mm; wing 2.6- 2.7 mm. Hind legs yellow to light brownish yellow; hind tibiae only slightly clavate, api- cal three tarsomeres of all tarsi yellowish brown to brown with the apical segment — Figs. 3-4. 3. Left femur and left tibia of male Town- sendia arenicola, n. sp. Scale = 0.4 mm. 4. Abdomen of Townsendia arenicola, n. sp. Scale = 0.4 mm. VOLUME 97, NUMBER 3 5a 5c Fig. 5: lateral. Scale = 0.4 mm. Abbr. Ae = Aedeagus, Ep = Epandrium, Ce = Cercus, Hy = Hypandrium, Gs = Gonostylus. dark brown. Wings almost entirely irides- cent yellow. Abdomen shiny with all of seg- ments | and 9, narrow base of segment 2, and narrow apex of segment 8, dark brown to black or sooty blackish yellow, remaining segments contrastingly lighter, often par- tially sooty, usually brownish yellow to yel- low. Sternite 8 narrowly divided medially with incision reaching apical third. Sper- matheca long, extending posteriorly to api- cal margin of segment 3; apically each sper- 693 Ep 5d Male terminalia of Townsendia arenicola, n. sp., a. lateral, b. dorsal, and c. ventral views; d. aedeagus, matheca tubular, darkly sclerotized, loosely coiled one and one-half times, darkly scler- otized, with apex slightly tapered; base of each ductus separate, not fused, and about as wide in diameter as spermatheca. Ter- minalia apically with eight black acantho- phorite spines. Holotype ¢ FLORIDA: Archbold Biol. Sta., LK Placid, Highlands Co., Rosemary shrub, 21.VI.1988, M. Deyrup; allotype °, same data except 18.VI.1992. Paratypes, 23 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2, same location, along various trails, frelanes, sand surfaces, and on a flowering Paronychia chartacea, | with chloropid prey, 4. VI-8.1X.89-92, B. Norden/M. Deyrup. Voucher material.—Holotype ¢ & allo- type 2, deposited in the National Museum of Natural History, Smithsonian Institution (USNM); paratypes in the National Muse- um of Natural History, the Archbold Bio- logical Station, and the Towson State Mu- seum of Zoology. Etymology.—Latin arenicola, meaning frequenting or living in sandy areas, refer- ring to the habitat of the species. Remarks.— Townsendia arenicola, n. sp. is easily separated from other known species in this genus by the wide gray tomentose bands on the apical margins of abdominal tergites 2—6, and by the dense gray and brown tomentose pattern on the scutum. The gen- eral color of the abdomen of females of 7. arenicola and T. pulcherrima are essentially identical, but the latter species lacks the gray tomentose bands. In addition, the antennal spine of T. pulcherrima is subapical, short, and hidden in the long pubescence, whereas this spine in T. arenicola is apical, long and exposed. In females of T. niger, the abdo- men is wholly blackish with the sides be- yond the middle strongly tapered to a point- ed apex. In 7. arenicola, the abdomen is much lighter in color, largely brownish yel- low to yellow, with parallel sides and a broad, rounded apex. It also differs from 7. niger in having only sparse, short vestiture on the front, ocellarium and scutum. In both 7. pulcherrima and T. niger, the vestiture of the head and scutum is much longer and more abundant than in 7. arenicola. The former species have three to five prominent tergosternal bristles whereas T. arenicola has only two such bristles. ACKNOWLEDGMENTS We (BBN, K VK) thank the personnel of the Archbold Biological Station (ABS), Lake Placid, Florida, for accommodations that facilitated our work. Mark Deyrup, Assis- tant Research Biologist ABS, kindly dis- cussed his behavioral observations of Townsendia, and loaned us specimens from the ABS collection. We are also grateful to Mark Deyrup, Arnold Norden and an anon- ymous reviewer for critically reading and providing helpful comments on the manu- script. LITERATURE CITED Back, E. A. 1909. The robber-flies of America, north of Mexico, belonging to the subfamilies Lepto- gasterinae and Dasypogoninae. Transactions of the American Entomological Society 35: 174-178. Bullington, S. W. and A. F. Beck. 1991. Anew species of machimus Loew (Diptera: Asilidae) from bur- rows of Gopherus polyphemu (Testudines: Testu- dinidae). Annals of the Entomological Society of America. 84(6): 590-959. Curran, C. H. 1926. New Diptera from the West Indies. American Museum Novitates. 220: 1-14. Lavigne, R. J. 1968. Notes on two species of Asilus (Diptera: Asilidae) associated with animal bur- rows, with a redescription of Asilus gilvipes Hine. Journal of the Kansas Entomological Society. 41: 334-339. Martin, C.H. 1966. The genus Townsendia Williston in Mexico (Diptera: Asilidae). Journal of the Kan- sas Entomological Society. 39: 542-551. Melin, D. 1923. Contributions to the knowledge of the biology, metamorphosis and distribution of the Swedish asilids in relation to the whole family of asilids. Uppsala Universitet Zoologiska Bidrag. 8: 1-317. Norden, B. B., K. V. Krombein, and B. N. Danforth. 1992. Taxonomic and bionomic observations on a Floridian panurgine bee, Perdita (Hexaperdita) graenicheri Timberlake (Hymenoptera: Andreni- dae). Journal of Hymenoptera Research. 1(1): 107— 118. Wood, G.C. 1981. Asilidae, pp. 549-573. Jn Manual of Nearctic Diptera. Biosystematic Research In- stitute, Agriculture Canada, Ottawa. Monograph #22. Williston, S. W. 1895. Two remarkable new genera of Diptera. Kansas University Quarterly. 4(2): 107— 109. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 695-700 RESPONSE OF GYPSY MOTH (LEPIDOPTERA: LYMANTRIIDAE) LARVAE TO STICKY BARRIER BANDS ON SIMULATED TREES RALPH E. WEBB, GEOFFREY B. WHITE, AND KEVIN W. THORPE Insect Biocontrol Laboratory, USDA-ARS, Bldg. 402, BARC-East, Beltsville, MD 2070S. Abstract.—The behavioral response of gypsy moth, Lymantria dispar (L.), unfed first instars to intact and defective sticky barrier bands was determined. The barrier bands were installed on simulated trees in a laboratory arena. Defects incorporated into the barrier bands were: gaps in the sticky material (2 mm, 4 mm, and 8 mm width), tunnels under the band (3 mm and 6 mm height) and bridges across the sticky material (cross sections of | mm? and 2 mm diameter). The defects allowing the highest percentage of larvae to cross were the 2 mm diameter bridge (55.3%) and the 6 mm high tunnel (46.6%). Other defects also allowed crossing. These findings indicate the importance of frequent inspection and repair of sticky barrier bands that are used for gypsy moth control. Key Words: Sticky barrier bands are used to protect trees against various pests, including the gypsy moth, Lymantria dispar (L.). Origi- nally, barrier bands were made by applying sticky substances formulated with materials which included coal tar derivatives and ros- in oil. These sticky substances were applied directly to the bark (Burgess 1930, Collins and Hood 1920, Forbush and Fernald 1896). Recently, other types of sticky materials have been applied on top of tape bands on tree boles (Blumenthal 1983, Blumenthal and Hoover 1986, Webb and Boyd 1983). Such barriers are still used by homeowners and managers of small tracts of land (Miller and Lindsay 1993). Success of this method depends upon the ability of the barrier to prevent the return of larvae to the canopy after ballooning or silking down (Burgess 1930, Leonard 1971, McManus 1973). In the field, barrier band efficacy may be re- duced by gaps in the sticky material from uneven application, tunnels under the tape caused by texture or contour of the tree bark, and dirt, debris or leaves that become stuck Lymantria dispar, barrier bands, behavior in the material creating bridges. Blumenthal (1983) made useful observations on the re- sponse of gypsy moth larvae when they en- countered barrier bands, however, these ob- servations were generally of older larvae, and did not include responses to damaged or otherwise compromised bands. Thorpe et al. (1993) found barrier bands reduced larval numbers in groups of trees, but only by about 28%. The present study involved the quantitative analysis of larval response to intact and compromised sticky bands through controlled laboratory experiments. Results lead to a better understanding of possible causes of variable performance of these devices (Blumenthal and Hoover 1986, Thorpe et al. 1993). METHODS Test arena, bole, and band treatments. — A corrugated cardboard carton was used to form the walls of the arena (60 cm?) (Fig. 1). The box was lined on the inside with white drawing paper to reduce directional bias due to light sources and extraneous sil- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SIDE VIEW TOP VIEW | | 60 CM | | | | | (NOT TO SCALE) Fig.. 1’. II® (Schutter et al. 1988). houettes, both of which influence larval movements (Ludwig and Schneider-Hem- pel 1954, Roden etal. 1992, Zanforlin 1970). The simulated bole was constructed of a cardboard mailing tube 92 mm outside di- ameter and 597 mm height (Alperstein Bros., Inc., Silver Spring, MD). The yellow paper sheathing on the tube was peeled off to ex- pose the unbleached, undyed cardboard. The TANGLEFOOT BAND ———> ‘RECO TANGLEFOOT Test arena with artificial bole (“tree”) assembly in place. Graphics were generated with MacDraw tube was attached to a corrugated cardboard base (approximately 58 cm?) by anchoring it with two straight pins and sealing the seam with acrylic latex caulk plus silicone (DAP Inc., Dayton, OH). Caulk was allowed to dry at ambient room temperature and rel- ative humidity for a minimum of 24 hrs. before the apparatus was used for testing. Placement of the artificial bole was approx- VOLUME 97, NUMBER 3 imately | cm from the edge of one side and on center relative to that side. A band of Tanglefoot® (Tanglefoot Co., Grand Rap- ids, MI) was applied in a circle (approxi- mately 50 cm diameter) on the base to pre- vent larvae from leaving the test arena. This containment band bisected the bole longi- tudinally so that larvae were restricted to one half of the bole facing the center of the arena. New bole and base assemblies were used for each replicate for each treatment in order to eliminate any possible influence from silk trails and chemical cues from pre- vious tests. The barrier band was constructed of duct tape (ServiStar®, SSR20340, SERVISTAR Corp., Butler, PA) 5.08 cm wide that was wrapped around the bole with the lower edge of the tape approximately 43 cm from the base. In all treatments, a sticky layer was formed by masking a | cm band in the cen- ter of the duct tape with Scotch® Magic Tape (3M Commercial Office Supply Di- vision, St. Paul, MN), applying Tanglefoot along the unmasked area and then smooth- ing the Tanglefoot with a glass microscope slide to give a thin, uniform layer that was 1 cm wide after removing the tape. All treat- ments were modifications of this basic con- figuration (Fig. 2). Treatments using gaps in the sticky layer were formed by masking with a piece of Magic® Tape of the appropriate size. Three widths of gaps were tested: 2 mm (approx- imately 2x the width of a first instar head capsule), 4 mm, and 8 mm. Treatments us- ing tunnels were formed by placing a plastic form on the duct tape band and securing it with another layer of duct tape; the Tangle- foot band was then applied on the top layer of duct tape. Two sizes of tunnels were test- ed. The form for the smaller tunnel (low tunnel) was made from a plastic soda straw that was bisected lengthwise and trimmed to 5.08 cm (duct tape width); the tunnel space was 3 mm wide at the base and 3 mm high at the center. The larger tunnel (high tunnel) was made from a piece cut from a 697 polystyrene container (thickness: 0.3-0.5 mm) and bent to form a tunnel with a base width of 10 mm and center height of 6 mm. Two sizes of bridges were tested. One type (1 mm bridge) was formed from a flat wood- en toothpick (Forster Mfg. Co., P.O. Box 657, Wilton, ME) that was trimmed to the dimensions 1.22 + 0.02 mm x 1.25 + 0.04 mm x 30 mm. The other bridge (2 mm bridge) was a 30 mm section of a wooden applicator stick (Fisher Brand®, Cat. # 01- 340) (diameter: 2.18 + 0.01 mm). The 1 cm band of Tanglefoot® was sufficient to hold the bridges in place. Test insects.—Gypsy moth egg masses were provided by the Otis Methods Devel- opment Center, US Department of Agri- culture, APHIS, Otis ANG, MA. They were from generations F37, F38, and F40 of the New Jersey Standard Strain (NJSS). They were stored at 4°C until three to five days before testing. They were then placed in a sealed plastic bag at ambient room temper- ature (20-—25°C) with moist paper towels to maintain high humidity. Egg masses were checked daily. When the larvae began mov- ing off the egg mass, that egg mass was used for tests. Larvae were introduced to the are- na by placing the entire egg mass in the center of the base allowing approximately 50-100 individuals to crawl off (Fig. 1), and then promptly removing the egg mass with the remaining larvae. All test insects were unfed first instars. Validation of test arena. —A series of pre- liminary tests were conducted to determine if test insects behaved in the arena in a man- ner consistent with that described in the lit- erature. Tropism and behavior of larval gypsy moths are well documented (Doan & Leonard 1975, Roden et al. 1992, Weseloh 1990, Zanforlin 1970) and it was desirable to verify that behavior in the arena was sim- ilar to that previously reported. A variety of images were presented to larvae in the arena, including a solid black panel covering one side of the arena, both black and white vertical bars on contrasting backgrounds, 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON CARDBOARD TUBE (9.2 cM DIA) DUCT TAPE (5.1 CM WIDTH) lilt fest sie We We (itt Vill TANGLEFOOT TUBE CARDBOARD TUBE DUCT TAPE TANGLEFOOT CAROBOARD TUBE BRIDGE Wy Nitrest. ODUCT TAPE YY, eae TANGLEFOOT d Fig. 2. Barrier band treatments as installed on artificial boles. In the one hour laboratory tests treatments included a) duct tape band with an unbroken sticky layer b) three sizes of gaps tested and c), d) two sizes each for the tunnels and bridges. Graphics were generated with Desk Paint® (Gariepy 1988). and cardboard mailing tubes of the type lat- er used in the barrier band tests. Groups of 50 to 100 larvae were observed for periods of 30 minutes or longer and their move- ments were recorded on video tape. The initial observations and later review of the video tapes indicated that the larvae were strongly attracted to black vertical bars of various widths on a white background. This is consistent with earlier reports of this be- VOLUME 97, NUMBER 3 havior (Roden et al. 1992, Zanforlin 1970). Larvae also readily moved toward and climbed the cardboard mailing tubes placed in the arena. Barrier band tests.— Tests were conduct- ed for | hour, which began at the time that the egg mass was removed from the center of the arena. The test was observed contin- uously, and every 10 minutes the following information was recorded: number of larvae that crossed the barrier band, total number of larvae on the bole, number that walked back down below barrier band, and number that silked down past the barrier or simply dropped. The first series of tests consisted of seven treatments: 1) duct tape banding with no Tanglefoot, 2) a barrier band with a contin- uous band of Tanglefoot, gaps in Tanglefoot of 3) 2 mm, 4) 4 mm, and 5) 8 mm, and the two different sizes of tunnels that were 6) 3 mm and 7) 6 mm high in the center. The second series of tests consisted of four treatments: 1) duct tape without Tanglefoot, 2) and 8 mm gap, and two sizes of bridges that were 3) approximately | mm? x 30 mm and 4) approximately 2 mm diameter x 30 mm. Each series was analyzed as a Latin Square design, with artificial boles and time periods as separate effects (Cochran and Cox 1957, SAS Institute 1987). RESULTS AND DISCUSSION The percentages of larvae that actually found and climbed on the bole during the 1 hour tests were 87.7% for series | and 78.1% for series 2. Analyses indicated no effects of treatment, bole, or time period for finding the bole. Numbers that either walked down or silked down below the barrier band during the test period were very low (typi- cally three or less out of 50-100) and typi- cally occurred in the last 10-20 minutes of the test period; these categories were not statistically analyzed. Notable exceptions occurred in two trials, both control treat- ments in series 2. In these trials, 28 and 18 larvae moved below the duct tape-only 699 Table 1. Percentage of unfed first instar larvae crossing barrier bands. Series 2 Treatment Series | Tape only 93.5a 82.4a High tunnel 46.6b — 8 mm gap SG! PLONE Low tunnel 58d — 4 mm gap 28d — 2 mm gap Od - Unbroken band of Tanglefoot Od a 2 mm bridge = Ds) Wo 1 mm bridge _ ZIEDIC (SE = 2.5) (SE=5.3) ' Numbers within a column followed by the same letter are not significantly different at a 0.05 compar- ison-wise error rate (LSD test). bands by either walking or silking down. We suspect this behavior was promoted, at least in part, by crowding around the top edge of the bole. Apparently many of these larvae re-crossed the band walking upward which resulted in reporting of greater than 100% crossing during the test. Because of this anomaly, these two data points were entered as 100% crossing for purposes of analysis. The frequency with which larvae were able to cross the band was calculated as a pro- portion of the numbers that actually found the bole during the tests (Table 1). There were significant treatment effects observed in both series of test (series 1: F = 194.1; df = 6,30; P = 0.0001; series 2: F = 26.7: df = 3,6; P = 0.0007). No bole effects were seen in either series. No time effect was seen in series 1, however the time effect was sig- nificant in series 2. This may have been due to some environmental factor or difference in handling of the group of egg masses used for tests. The high tunnel and the 8 mm gap both allowed substantial numbers of larvae to cross the barriers (46.6% and 31.4%, re- spectively) (Table 1). In contrast, no larvae successfully passed through the 2 mm gap, which, based on measurements of the lar- vae, is wide enough to allow passage without touching the sticky material. Some larvae were observed entering the 2 mm gap, 100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sometimes as far as half their body length, and then backing out of the gap. The comparison of bridges and the 8 mm gap indicated that a bridge of approximately | mm width reduced the effectiveness of the barrier bands as much as an 8 mm gap in the sticky material (Table 1). Doubling of the bridge width in this test allowed a two- fold increase in the proportion of larvae that crossed the barrier. Bridges were more detrimental to the ef- ficacy of barrier bands than were gaps and tunnels of similar size, based on those sizes tested in these experiments (Table 1). The 2 mm bridge allowed strikingly more neo- nates to cross than did the 2 mm gap and the low tunnel. It should be remembered that these test periods were limited to | hour. Since more time 1s available to larvae in the field it is reasonable to expect more indi- viduals to cross bands having defects. This indicates the importance of frequent in- spection and repair of these devices to op- timize their performance. ACKNOWLEDGMENTS The authors thank Gary Bernon and the other personnel at Otis Methods Develop- ment Center, USDA, APHIS, Otis ANG, Massachussetts for providing gypsy moth egg masses from the New Jersey Standard Strain. Also, thanks to Stephen P. Cook, Thomas C. Elden, Michael J. Raupp, and anonymous reviewers for their comments. LITERATURE CITED Blumenthal, E.M. 1983. Gypsy moth defoliation re- duction using mechanical barrier devices. Mels- heimer Entomological Series 33: 21-30. Blumenthal, E. M. and C. R. Hoover. 1986. Gypsy moth (Lepidoptera: Lymantriidae) population control using mechanical barriers and contact in- secticides applied to tree stems. Journal of Eco- nomic Entomology 79: 1394-1396. Burgess, A. F. 1930. The Gipsy Moth and the Brown- tail Moth. USDA Farmer’s Bulletin No. 1623. 33 pp. Cochran, W. G. and G. M. Cox. 1957. Experimental Designs, 2nd ed. John Wiley & Sons, New York. 611 pp. Collins, C. W. and C. E. Hood. 1920. Gypsy moth tree banding material: How to make, use, and ap- ply it. USDA Bull. 899. Doane, C. C. and D. E. Leonard. 1975. Orientation and dispersal of late-stage larvae of Porthetria dis- par (Lepidoptera: Lymantriidae). Canadian En- tomologist 107: 1333-1338. Forbush, E. H. and C. H. Fernald. 1896. The Gypsy Moth. Porthetria dispar (Linn.). Wright & Potter Printing Co., Boston. 495 pp. Gariepy, A.R. 1988. Desk Paint®, version 2.01. Zed- cor, Inc. Leonard, D. E. 1971. Air-borne dispersal of larvae of the gypsy moth and its influence on concepts of control. Journal of Economic Entomology 64(3): 638-641. Ludwig, W. and I. Schneider-Hempel. 1954. [Con- stancy of lower animals in experiments with one or two lights. III. Experiments with blinded ani- mals and under various light gradients.] Zoolo- gische Jahrbucher Abteilung fiir allgemeine Zoolo- gie und Physiologie der Tiere 65(1): 126-140. Bi- ological Abstracts 30, No. 24775. McManus, M. L. 1973. The role of behavior in the dispersal of newly hatched gypsy moth larvae. USDA Forest Service Research Paper NE-267. 10 pp. Miller, J. D. and B. E. Lindsay. 1993. Influences on individual initiative to use gypsy moth control in New Hampshire, USA. Environmental Manage- ment 17: 765-772. Roden, D. B., J. R. Miller, and G. A. Simmons. 1992. Visual stimuli influencing orientation by larval gypsy moth, Lymantria dispar (L.). Canadian En- tomologist 124: 287-304. SAS Institute. 1987. Release 6.04. PROC GLM. Schutter, G., A. Goldsmith, M. Kaptanoglu, J. Speigel, and A. Wagner. 1988. MacDraw II® 1.0v2, Clar- is® Corporation. Thorpe, K. W., R. E. Webb, R. L. Ridgway, L. Ven- ables, and K. M. Tatman. 1993. Sticky barrier bands affect density of gypsy moth (Lepidoptera: Lymantriidae) and damage in oak canopies. Jour- nal of Economic Entomology 86(5): 1497-1501. Webb, R. E. and V. K. Boyd. 1983. Evaluation of barrier bands and insecticidal strips for impeding intraplant movement of gypsy moth caterpillars. Melsheimer Entomological Series 33: 15-20. Weseloh, R. 1990. Simulation of litter residence times of young gypsy moth larvae and implications for predation by ants. Entomologica Experimentalis et Applicata 57: 215-221. Zanforlin, M. 1970. The inhibition of light orientat- ing reactions in caterpillars of Lymantriidae, Ly- mantria dispar (L.) and Orgyia antiqua (L.) (Lep- idoptera). Monitore Zoologico Italiano (N.S.) 4: 1-19. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 701-716 COCCINELLA NOVEMNOTATA IN NORTHEASTERN NORTH AMERICA: HISTORICAL OCCURRENCE AND CURRENT STATUS (COLEOPTERA: COCCINELLIDAE) A. G. WHEELER, JR. AND E. RICHARD HOEBEKE (AGW) Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Pennsylvania 17110; (ERH) Department of Entomology, Cornell University, Ithaca, New York 14853. Abstract. —A review of the literature documents that the native lady beetle Coccinella novemnotata Herbst (C9) was once common in the northeastern United States and Canada. Despite extensive recent fieldwork and surveys for coccinellids, only five collection records of C9 in the Northeast have been located since the mid-1980s. Its apparent decline in numbers and possible local extirpation could be the result of factors such as changes in land-use and cropping patterns, decline in aphid populations, parasitism, or disease. The factor most often suggested is possible adverse effects from the Old World C. septem- punctata L. (C7), whose establishment in North America was detected in 1973. New World populations of C7 may have resulted from previous releases for the biological control of aphids or an unintentional importation with commerce. Without a cause-and- effect relationship having been established, proposed detrimental impacts of C7 on native coccinellids are based solely on anecdotal evidence and speculation. Even though C7 was extensively recolonized in North America by biological control specialists, the C7 project does not typify classical biological control. If C7 has had a negative effect on C9, it is more appropriately considered displacement of an indigenous species by a polyphagous nonindigenous species than an example of unintended effects of classical biological control. Key Words: For several years we have realized that the native lady beetle Coccinella novem- notata Herbst (hereafter C9) has become in- creasingly difficult to find in the Northeast and may even be locally extirpated. One hypothesis proposed to account for its ap- parent decline in numbers is adverse effects from the establishment in North America of a more aggressive congener, the Old World C. septempunctata L. (hereafter C7). In this paper we will demonstrate that C. novemnotata was once a widespread and sometimes abundant coccinellid in north- eastern North America (Delaware, Mary- land, and New Jersey north to Ontario and Insecta, lady beetles, biological control, faunal change Quebec) and that it has been collected only once during surveys and extensive fieldwork in the Northeast in the 1990s. Our intent is (1) to document what others have suspect- ed: that C9 is no longer a common species in the Northeast; (2) to stimulate entomol- ogists, ecologists, and conservation biolo- gists to search for C9 populations in the Northeast and to deposit voucher speci- mens in major entomological collections; (3) to encourage workers in western North America, where C7 has become established more recently, to begin, or continue, to monitor populations of C9; and (4) to in- crease interest in documenting the effects of 0 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON iventive species— whether intentionally or intentionally introduced—on indigenous species. C7 IN NoRTH AMERICA The first U.S. releases of this well-known Old World coccinellid (e.g. Hodek 1973) were made by the U.S. Department of Ag- riculture (USDA) in California in 1957, based on material from India. Adults were recovered after a month but no further in- dividuals were collected, and no eggs or lar- vae were observed. From 1958 to 1973, beetles from India, France, Italy, Norway, and Sweden were released (nearly 150,000) for aphid control in eastern and western states by the USDA (Angalet et al. 1979, Schaefer et al. 1987). In eastern Canada small numbers were released in New Bruns- wick during 1959-1960 (Clark et al. 1971, Schooley et al. 1984). A particular effort was made to establish C7 in Maine for suppres- sion of potato-infesting aphids; about 80,000 individuals were released in test plots at Presque Isle (Shands et al. 1972). Although the Fl generation was recovered at several localities, these releases apparently did not lead to permanent colonization in the Unit- ed States or Canada (Angalet and Jacques 1975, Angalet et al. 1979, Schaefer et al. 1987). C7’s detection in New Jersey (Angalet and Jacques 1975) and Quebec (Larochelle and Lariviére 1979) in 1973 renewed interest in this predator and led to redistribution ef- forts during 1974-1978 (Angalet et al. 1979). More than 500,000 beetles were released in 20 states and the District of Columbia (Schaefer et al. 1987), and in Canada they were redistributed in Nova Scotia (Kelleher 1984). The beetle’s natural dispersal from the area of detection in New Jersey was con- : sidered slow: by 1975 records were avail- able only for 10 counties in Connecticut, New Jersey, and New York; its presence in Delaware was considered the result of re- colonization (Angalet et al. 1979). But C7’s rapid spread began to be documented (Hoe- beke and Wheeler 1980). Natural spread, aided by successful recolonization in Geor- gia, Ohio, and Oklahoma, resulted in re- coveries from 15 states by the end of 1979 (Schaefer et al. 1987 and references therein). By 1986, C7 was established in 34 eastern and central states and in 5 Canadian prov- inces (Schaefer et al. 1987). In Iowa and Missouri this recently established predator was still less abundant than any of the native coccinellids found in croplands, abandoned fields, and roadsides (Obrycki et al. 1987). Additional releases were planned for the western two-thirds of the United States (Comis and Heppner 1986). Detection of the Russian wheat aphid, Diuraphis noxia (Mordvilko), in Texas in 1986 (Stoetzel 1987) further emphasized the need to re- distribute C7 in the West (Olkowski et al. 1990, Gordon and Vandenberg 1991). This project was led by the USDA-Animal & Plant Health Inspection Service’s National Biological Control Laboratory in Niles, Michigan, and redistribution of C7 in four western states was initiated in 1989 (Flan- ders et al. 1993). It is now the most com- monly collected species of Coccinella east of the Rocky Mountains (Gordon and Van- denberg 1991), one of the dominant cocci- nellids of agricultural crops in Michigan (Maredia et al. 1992), and is known from all 48 contiguous states (Prokrym et al. 1992; see also Rice 1992). In western Canada it occurs in the Prairie Provinces and in cen- tral British Columbia (Humble 1991, Mc- Namara 1991). Gene diversity of North American populations is similar to that among Eurasian C7, such a broad genetic basis being characteristic of other successful adventive insects in the New World (Kraf- sur et al. 1992). The origin of North American popula- tions of C7 remains in doubt. It was first suspected that the beetles found at the Hackensack Meadowlands in New Jersey were associated with disposal of trash at sites near Kennedy International Airport (An- galet and Jacques 1975). C7’s collection in VOLUME 97, NUMBER 3 Quebec, however, suggested separate intro- ductions with transoceanic commerce— along the St. Lawrence Seaway in Quebec and the Upper Bay of the Hudson River and at New Jersey ports of entry (Schaefer et al. 1987). Although an unintentional in- troduction with ship trafic remains a strong possibility (Schaefer and Dysart 1988, Day et al. 1994), an alternative hypothesis should be considered: that classical biological con- trol releases of C7 made during 1958 to 1973 led to its establishment in the East (Schaefer et al. 1987). Studies of genetic diversity in North American populations have not dis- criminated between these two hypotheses (Krafsur et al. 1992). C9 In NorRTH AMERICA Widespread in North America, this na- tive lady beetle ranges from Maine, Ontario, and Quebec to Florida and west to British Columbia and southern California (Dob- zhansky 1931, Gordon 1985) and occurs in all northeastern states (Dobzhansky 1931, Procter 1946, Belicek 1976). In Canada it is found north to 46° in Quebec, 45°30’ in Ontario, and 62° in the Northwest Terri- tories (Brown 1962). Recorded prey includes numerous aphid species (Thompson and Simmonds 1965), and in laboratory evaluations C9 fed more on aphids than on spider mites or lepidop- teran eggs (Putman 1957). Larvae and adults similarly preferred aphids to larvae of the alfalfa weevil, Hypera postica (Gyllenhal), or leafhopper nymphs (Yadava and Shaw 1968). Although C9 has been considered an important natural enemy of the European corn borer, Ostrinia nubilalis (Hiibner) (Sparks et al. 1966), more recent work in- dicates that predation on corn borer eggs is unimportant (Andow 1990). Feeding has also been observed at extrafloral nectaries (Putman 1964, Pemberton and Vandenberg 1993; see also van den Bosch and Telford 1964: Fig. 92). C9 is especially common in gardens and 703 other cultivated lands (Leng 1903, Blatchley 1910, Stehr 1930, Chapin 1974, Belicek 1976), occurring in field crops such as alfalfa (Fluke 1925, Goodarzy and Davis 1958, McMullen 1967a, Neuenschwander et al. 1975: see also Hodek 1973), clover (Folsom 1909, Smith 1958), corn (Everly 1938, Bar- tholomai 1954, Smith 1971, Wright and Laing 1980), cotton (Bell and Whitcomb 1964, Whitcomb and Bell 1964, van den Bosch and Hagen 1966), potatoes (Leonard 1963, Day 1965, Mack and Smilowitz 1980), small grains (Kirk 1970, Shade et al. 1970, Bernal et al. 1993), and soybeans (Blick- enstaff and Huggans 1962, Dumas et al. 1964, Tugwell et al. 1973). In Connecticut (Britton 1914) and Minnesota (Stehr 1930), C9 has been ranked as one of the coccinel- lids of greatest economic importance. C9 can also be collected on weeds in disturbed areas (e.g. McMullen 1967a, Richerson and DeLoach 1973, Dailey et al. 1978, Lago and Mann 1987, Maredia et al. 1992), as well as on apple (Smith 1958, LeRoux 1960, Oatman et al. 1964, Hagley 1975, Travis et al. 1978, Carroll and Hoyt 1984), peach (Putman 1964, Kirk 1970), conifers, and hardwood trees (Felt 1906, Smith 1958, Gagné and Martin 1968, Drooz 1985). Detailed life history studies of C9 have not been conducted, although Burgess (1903) and Palmer (1914) provided information on fecundity and duration of life stages in the laboratory. Data on preoviposition period, fecundity, and longevity were obtained by McMullen (1967b) and duration of egg and larval stages by Gagné and Martin (1968). Studies on the sex ratio, weight, and size of adults have also been conducted (Smith 1966). Its relative abundance, seasonal his- tory, adult behavior, and factors inducing diapause were studied in California (McMullen 1967a, b). This predator is ap- parently univoltine in Ontario (Gagné and Martin 1968) and bivoltine in Colorado (Palmer 1914). An important mortality fac- tor may be parasitism by the Holarctic brac- onid Dinocampus coccinellae (Schrank) 704 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hudon 1959, Richerson and DeLoach Ad > Je | Historical occurrence before establish- ment of C7.—C9 is referred to as frequent throughout Indiana (Blatchley 1910), one of the most abundant coccinellids in Oregon (Ewing 1913), common in Iowa prairies (Hendrickson 1930), and the most common species of Coccinella found in Minnesota and the Upper Mississippi Valley (Stehr 1930, Wingo 1952). Occurring statewide in North Carolina (Brimley 1938), South Da- kota (Kirk and Balsbaugh 1975), and sev- eral northeastern states (see Table 1), C9 is included in many field guides and general references (e.g. Lutz 1948, Jaques 1951, Zim and Cottam 1956, Dillon and Dillon 1961, Swan 1964, Borror and White 1970, Swan and Papp 1972, Milne and Milne 1980, Ar- nett and Jacques 1981, Arnett 1985, Boyd 1991). In fact, C9 is used as the lead illus- tration for the coccinellid sections in Arnett (1968) and Borror and White (1970) and in 1989 was designated the official state insect of New York (see Hoffmann and Frodsham 1993). In addition, the USDA’s Cooperative Economic Insect (later Plant Pest) Report contains numerous references to C9 on var- ious crops. Examples of these state reports include: “Extremely active” on crimson clo- ver, vetch, and cotton in Mobile Co., Ala- bama (Seibels et al. 1963); “extremely heavy feeding” on woolly apple aphid, Eriosoma lanigerum (Hausmann), on apple in Talla- poosa Co., Alabama (Webb et al. 1965); and the most abundant coccinellid in alfalfa (75- 100/100 sweeps) in Lafayette Co., Arkansas (Boyer 1970). At one time C9 was routinely collected in the Northeast (Table 1; see also locality records in Dobzhansky 1931). C9 in the Northeast, 1973-1985.—After C7’s detection in North America, C9 con- tinued to be collected in insect surveys in the Northeast during 1973 to 1985. From 1974 to 1978, C9 was scarce at the Hack- ensack Meadowlands, where C7 had be- come the dominant coccinellid, and it was also much less numerous than C7 at a near- by site in Connecticut in 1978 (Angalet et al. 1979; see also Table 2). When C7’s es- tablishment was first reported in Pennsyl- vania, similar numbers of both coccinellids were obtained in limited sweepnet samples (Hoebeke and Wheeler 1980; Table 2). Two years earlier, large numbers of C9 had been found on apple in Pennsylvania (Travis et al. 1978; Table 2). C9 was not found in surveys of managed, abandoned, and “organic”’ apple orchards in New York, Pennsylvania, Virginia, and West Virginia during 1983 and 1984, al- though C7 was present (Brown et al. 1988). Three specimens of C9, however, were col- lected in Jefferson Co., West Virginia, in 1982, 1984, and 1985 (2 in blacklight traps, 1 in an unsprayed orchard); this species has not since been collected on apple in Jeffer- son Co. (M. W. Brown, pers. comm. 1994). C9 also was not among the 10 coccinellid species, including C7, collected during a survey of corn insects in the Connecticut Valley of Massachusetts from 1982 to 1984 (Eaton 1984). C9 in the Northeast since 1985.—We are aware of only five collections of C9 since 1985 (Table 3). One adult was collected at each of two localities in Maryland in 1986 during a survey of coccinellids associated with nursery stock from 1986-1988; C7, however, was taken at 87 of the 186 loca- tions and was the most abundant of the 28 species collected (Staines et al. 1990). C9 has not been seen in Maryland since 1986 (C.L. Staines, Jr., pers. comm. 1994). Pop- ulations of the aphid Cinara pilicornis (Har- tig) were monitored on spruce (Picea spp.) seedlings and transplants in a southcentral Pennsylvania nursery during 1987-1988, and two C9 adults were observed on 13 May 1987. But it was C7 that was the most abun- dant coccinellid associated with aphid-in- fested spruce (Wheeler 1989 and unpubl. data). In Delaware during a census of over- wintering Coccinellidae in a 0.5 ha plot con- taining >1000 grass clumps, 27 C9 adults were found at Delaware City in winter 1987- VOLUME 97, NUMBER 3 705 Table 1. Examples of historical occurrence of Coccinella novemnotata (C9) in Northeast before establishment of C. septempunctata (C7). State/Province Remarks Reference Connecticut Common throughout state; of >30 coccinellid spp. re- Britton 1914 corded, among the 12 considered most economically important Very common at Meriden Johnson 1915 Maine Scarce in Mount Desert Region Procter 1946 Massachusetts On white birch at Malden, feeding on aphid eggs, Burgess 1903 1898 Common on Nantucket Island Johnson 1930 Present in cranberry bogs Franklin 1950 New Jersey Common throughout state Smith 1910 Larvae, adults on Hibiscus moscheutos L. Weiss and Dickerson 1919 On Oenothera biennis L., feeding on aphids Dickerson and Weiss 1920 On Asclepias syriaca L., A. tuberosa L. Weiss and Dickerson 1921 New York Abundant on Pinus rigida Mill. at Karner (Albany Felt 1906 Pine Bush) A common species on Staten Island Leng and Davis 1924 4th in abundance of 8 coccinellid spp. on alfalfa at Pack 1925 Ithaca: 8 adults collected during 10 min. of sam- pling; one of most abundant coccinellids at Ithaca More localities listed than for any of the other 66 coc- Leonard 1928 cinellids recorded from state Present in small numbers on collards at Ithaca, 1957— Pimentel 1961, Root 1973 1958; 1966-1968 2nd most numerous coccinellid on potatoes on Long Leonard 1963, Day 1965 Island, 1956-1958 (19% of family); sometimes abundant late June-early July On cruciferous crops on Long Island, 1960-1963 Sutherland 1966 Present in low densities on alfalfa at Ithaca, 1966- Pimentel and Wheeler 1973 1969 Present in 100-sweep samples of alfalfa in Columbia, A.G.W., unpublished data Orange, and Steuben counties, May—Aug.; largest number/100 sweeps = 12 in Columbia Co., 15 July 1970 Ontario Ranked 3rd in abundance among 9 most numerous Smith 1958 coccinellid spp. in survey of forages, weeds, and trees; 13% of 2300 specimens collected, 1957 Ranked 6th of 10 coccinellid spp. collected, 1948- Putman 1964 1960, on peach; 2.9% of 888 specimens identified Represented 19.1% of coccinellids associated with es- | Gagné and Martin 1968 tablishment stage of red pine, 1964; numbers dropped drastically in 1965 Represented 20% of coccinellids (6 spp.) on corn, Smith 1971 1963 Pennsylvania Very abundant on oak, feeding on Archips semiferana _ Nichols 1971 (Walker) Quebec On apple at Rougemont, 1951-1955 and 1955-1957; LeRoux 1960, Parent 1967 predacious on aphids Known historically from 24 localities Larochelle 1979 Vermont Known historically from 25 collections Parker et al. 1976 706 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2 ent of C. septempunctata (C7). State/Province Remarks 2. Examples of occurrence of Coccinella novemnotata (C9) in Northeast, 1973-1985, after establish- Reference Connecticut Hammonasset, Aug. 1978 16 adults collected (vs. 175 for C7) during survey at Angalet et al. 1979 New Jersey Scarce at Hackensack Meadowlands, 1974-1978; Angalet et al. 1979 C7 the most abundant of 17 coccinellid spp. oc- curring at site Common in Pine Barrens Boyd and Marucci 1979 Ontario On apple at Vineland, 1969-1974 Hagley 1975 On Onopordum acanthium L., 1976 Judd 1978 ‘“‘Never numerous” in corn, 1977-1978 High population levels on apple, 1977; aided signif- icantly in reducing aphid numbers Occasionally common on Evonymus alatus Pennsylvania Wright and Laing 1980 Travis et al. 1978 Wheeler and Stimmel 1979 (Thunb.) Sieb. infested with Aphis fabae Scopoli, 1977 10 adults/300 sweeps of weeds in ruderal site (vs. 12 for C7), Harrisburg, 1979 Collected 4 times (vs. 5 for C7) on soybeans, 1980- 1982 Larva reared from aphid-infested shoots of Physo- Hoebeke and Wheeler 1980 Wheeler and Stimmel 1983 Wheeler and Hoebeke 1985 carpus opulifolius (L.) Maxim., 1979 Quebec Vermont 1988; this survey yielded 3000 C7 adults (P. W. Schaefer, pers. comm. 1993). In a study in Maine, C9 was the second most abundant coccinellid (C7 was most abun- dant) found in potato plots and the second most numerous lady beetle in barley plots adjacent to potato in 1992. C9 was not en- countered in either crop in 1993 (F. A. Drummond, pers. comm. 1994), which could reflect normal year-to-year fluctua- tions in density rather than an actual decline in numbers. No detections of C9 were made in the Northeast during 1993 coccinellid surveys conducted as part of the USDA’s Cooper- ative Agricultural Pest Survey program. In Connecticut, an alfalfa field in each of 4 counties was sampled 6 times (400 sweeps/ visit) from 10 June to 19 August and once in September; supplemental 200-sweep samples were taken at 16 sites in 4 counties during June to September (D. Ellis, pers. comm. 1993). New York surveys consisted of 1500 sweeps taken 4—5 times in alfalfa Collected at 5 localities (vs. 43 for C7), 1979 On apple in Chittenden Co., 1973-1974 Larochelle and Lariviére 1980 Hauschild 1975 or clover fields in each of 4 counties (1 was sampled only 3 times) from 15 July to 9 September; additional samples from forages and goldenrod were taken in 5 counties dur- ing August and September (J. J. Knodel, pers. comm. 1993). In Pennsylvania, 3 al- falfa fields in 3 counties were each moni- tored 6 times (400 sweeps/visit) from 18 June to 1 September; 52 additional fields or disturbed weedy sites were surveyed in 18 counties from June to August. Similar coc- cinellid surveys in various disturbed habi- tats in Pennsylvania (136 sites in 23 eastern counties) were also negative for C9 in 1994 (A.G.W., unpubl. data). Moreover, C9 was not observed during an extensive survey for the adventive Hip- podamia variegata (Goeze) in the Northeast in 1992. Nearly 600 adults of 8 coccinellid species, including 66 C7, were collected in 8 states (Wheeler 1993). We have not seen C9 in general collecting since 1985 or in our surveys of disturbed and relatively undis- turbed habitats ranging from urban vacant VOLUME 97, NUMBER 3 707 Table 3. Known records of Coccinella novemnotata (C9) in Northeast since 1985; see text for additional information on collections. State Remarks Delaware 27 adults at Delaware City, winter 1987-1988 Jet Maine Common on barley and potatoes in study plots at Ee Presque Isle, 1992 Maryland Two collections during 1986-1988: 1 adult on nurs- Reference W. Schaefer, pers. comm. A. Drummond, pers. comm. Staines et al. 1990 ery stock in Allegany Co., another in Carroll Co. Pennsylvania Co. nursery, May 1987 lots to pitch pine-scrub oak barrens, ser- pentine barrens, and shale barrens. No re- cent records of C9 were available in the in- sect collections that were checked: Ameri- can Museum of Natural History, New York; Canadian National Collection, Ottawa; Carnegie Museum of Natural History, Pitts- burgh; Cornell University, Ithaca, N.Y.; Florida State Collection of Arthropods, Gainesville; National Museum of Natural History, Washington, D.C.; Ohio State Uni- versity, Columbus; Pennsylvania State Uni- versity, University Park; University of Maine, Orono; and University of New Hampshire, Durham. DISCUSSION Adverse effects from the establishment of C7 is only one possible explanation for the apparent decline in C9 populations. Other factors that could be involved are changes in land-use and cropping patterns, decline in aphid densities, parasitism, disease, or even global warming. It is C7, however, that has been proposed most often as the likely cause of C9’s decline. Soon after C7’s establishment in eastern North America, its possible detrimental ef- fects on native coccinellids began to be not- ed. There was no evidence for C7’s replace- ment of native coccinellids in Georgia with- in three years of its release and establish- ment for suppression of pecan aphids (Tedders and Angalet 1981), but by the ear- ly 1980s a possible “‘antagonistic relation- ship with C. novemnotata appeared to be Two adults on spruce transplants in Cumberland Wheeler 1989, unpubl. data developing” in Ontario (W. Y. Watson, let- ter to A.G.W., 11 Feb. 1983). The need to evaluate the effects of the rapidly spreading C7 on the native coccinellid fauna became apparent (Schaefer et al. 1987). When field surveys were conducted during a three-year period in Maryland (186 localities), the once “very common” C9 was collected only twice. Competitive displacement by C7 was sug- gested as a reason for the apparent dimin- ished numbers of C9 in Maryland (Staines et al. 1990). Ehler (1990) also emphasized C7’s potential for affecting nontarget species (see also Evans 1991, Tedders 1992, Elliott et al. 1993, and Wheeler 1993). The sev- enspotted lady beetle’s possible effects on endangered lycaenid butterflies have re- cently been evaluated in Ohio. Although their population declines coincided with in- creases in C7 and this coccinellid fed on lycaenid eggs in the laboratory (Horn 1991), no field data are available to substantiate any adverse effect of C7 on these endan- gered lepidopterans. Populations of C9 in the Northeast seem to have declined sharply during the 1980s and 1990s, a period when the Old World C7 was undergoing rapid range expansion. Our after-the-fact evidence for the adverse effects of C7 on C9 must be considered cir- cumstantial. Populations of C9 were not monitored systematically during the time when C7 was becoming established in the Northeast and assuming dominance among coccinellids in disturbed and relatively un- disturbed communities. Coccinellid densi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ies often fluctuate widely from year to year (e.g. Foott 1974, Elliott and Kieckhefer 1990, Kieckhefer and Elliott 1990, Elliott et al. 1993). Quantitative data from sam- pling at several sites over a 10- to 15-year period—beginning even before the estab- lishment of C7 in the local fauna—would therefore be needed to assess accurately C7’s role among various other factors that may be responsible for a decline in populations of C9. Except in classical biological control of weeds programs, rarely are such quantified data available assessing the effects of ad- ventive species on indigenous organisms; however, there are incomplete baseline data documenting the presence and abundance of C9. Literature references adequately sup- port the view that C9 was once routinely collected or observed in the Northeast, of- ten in considerable numbers. If C9 were still relatively common, it should have been de- tected during recent surveys for Old World coccinellids in the Northeast or in our ex- tensive fieldwork involving agricultural crops, herbaceous weeds, shrubs, and trees since 1987. Adults of C9 are conspicuous because of their size (Britton 1914). Even though adults somewhat resemble those of C7, the two species are easily recognized. The head of C9 has a solid white rectangle instead of two white dots, and the anterior pronotal margin has a narrow white border, which is lacking in C7. C9’s recognition in the East is not complicated by the pro- nounced color polymorphism that charac- terizes certain native or naturalized lady beetles in our fauna; only the fully maculate nine-spotted morph occurs in eastern North America (Belicek 1976, Gordon and Van- denberg 1991). As evidence for a recent de- cline in C9 populations, data from extensive fieldwork probably should be weighted more heavily than the absence of new material in collections (museum accessions sometimes are not processed for several years). The sevenspotted lady beetle is a vora- cious, nearly ubiquitous aphidophage in the Old World that can also be characterized as eurytopic, polyphagous, and ecologically plastic (Hodek 1973). The aggressive adults (see Miller 1992) will attack early-instar chrysopid larvae, even when other prey are available (Sengonca and Frings 1985). For a summary of studies on its foraging be- havior, see Kareiva (1986) and Andersen and Kareiva (1993). C7 is an active flier and an aggressive colonizer that has become es- tablished on Sable Island, Nova Scotia, which is isolated in the Atlantic about 175 km from the nearest land (Schaefer et al. 1987), and also at high elevations (nearly 3500 m) in the Rocky Mountains (Rice 1992). The only evidence available that C9 might be susceptible to interspecific competition is the possibility that its gradual disappear- ance from establishment-stage red pine in Ontario was the result of competition from other coccinellids (Gagné and Martin 1968). Competitive displacement and interspecific predation by C7, as well as pesticide use and changes in land management, could con- tribute to declining populations of C9 in the Northeast. Determining the precise nature of C7’s detrimental effects on C9 would prove difficult. C7’s explosive colonization of North America provides an opportunity for eval- uating the effects of an aggressive polyph- agous predator on nontarget organisms. Eh- ler (1990) found “‘it difficult to believe that this introduction will not have an impact on non-target species in the United States,” and Evans (1991) suggested that C7 “may have profound impact on the ladybeetle fauna native to North America through complex interactions of Old and New World ladybeetles.”” Indeed, uncommon coccinel- lids temporarily disappeared from crop- fields in South Dakota following the inva- sion and establishment of C7 (Elliott et al. 1993), although C7 was not shown to be a direct cause of their decline; factors other than C7’s establishment could be involved. C9 is still common in parts of western VOLUME 97, NUMBER 3 North America. In fact, its relatively high densities (and those of Hippodamia conver- gens Guérin-Méneville) may have hindered or delayed C7’s establishment in California (Flanders et al. 1993). At most, C9 may be only locally extirpated in the East. Workers in the Northeast and even the Southeast (C9 appears at least to have declined in Alabama and Mississippi during the past five years; P. M. Estes and R. L. Brown, pers. comm. 1994, 1995) are encouraged to look for this native species. Workers in areas where C7 iS a more recent invader may want to begin, or continue (Elliott et al. 1993), to monitor its potential effects on C9 and other native coccinellids, as well as document a decline in populations of injurious aphids (see Kauffman and Schwalbe 1991). There is also a need to determine if other recently estab- lished Eurasian lady beetles—Harmonia axyridis (Pallas), Hippodamia variegata, and Propylea quatuordecimpunctata L.—are af- fecting native coccinellids in the East (see Day et al. 1994). Most people will not be concerned if an introduced predator having superior attri- butes reduces pest populations more than do indigenous natural enemies. So many plant and animal species have been affected by the needs of human society (e.g. Soulé 1990) that a decline in numbers of one or a few native predators will be viewed as inconsequential. Local extirpation or ex- tinction of C9 would elicit more concern. It is likely that C9 will find habitats in which it can coexist with C7; such popu- lations, as yet undetected, may exist in the Northeast. Its populations may again build to sizable levels. The current low densities of C9 in the Northeast may actually be sim- ilar to those that existed before the advent of agroecosystems, which facilitated in- creases in aphid numbers, allowing C9 per- haps to reach population levels greater than before human intervention. The benefits of C7’s presence in the Ne- arctic fauna may outweigh any costs to the environment. Use of pesticides against an 709 important crop pest may actually pose more of an environmental threat than does the release of some biological control agent (e.g. Nechols et al. 1992). The establishment of C7 can be viewed as representing a contin- uum of ecological changes. We have dis- cussed C7 as a principal factor contributing to a decline in populations of C9, although the evidence is speculative and anecdotal. But C7 may even be having positive effects on other nontarget organisms. The importation of any biological control agent 1s an experiment. Most introduced species fail to become established, relatively few provide substantial suppression of tar- get pests, and fewer still cause serious en- vironmental problems (e.g. Hall and Ehler 1979, Hallet al. 1980, Samways 1988, Ehler 1990). Whether any environmental disrup- tion associated with C7’s establishment is considered acceptable should await the re- sults of long-term ecological monitoring, preferably at sites where C7 and C9 do and do not co-occur, and critical evaluation of the accumulated quantitative data. Even then, an evaluation of the program to re- colonize C7 in North America will be influ- enced by one’s environmental, political, and social views. SUMMARY AND CONCLUSIONS Classical biological control has tradition- ally been favorably received by conserva- tionists and environmentalists (Samways 1988) but is under increasing pressure from such groups (Howarth 1983, 1991, Nechols and Kauffman 1992, Simberloff 1992, Lockwood 1993, Miller and Aplet 1993, U.S. Congress 1993). We realize the intro- duction of biological control agents has been viewed as part of the larger problem of en- vironmental disruption resulting from in- vasion of nonindigenous species (Howarth 1983, 1991, Samways 1988, Ehler 1990, 1991, Miller and Aplet 1993). Moreover, we support the need to analyze and evaluate biological control projects, but the data used and conclusions reached should have a 710 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ind scientific basis and avoid inference ind speculation. We cannot document a cause-and-effect relationship between the establishment of C7 and the decline of C9. What can be stated with reasonable certainty is that C7 has in- creased and, at the same time and in some of the same places, C9 has decreased. Pro- posed adverse effects of C7 on native coc- cinellids, such as Hippodamia convergens (Tedders 1992) and C. transversoguttata ri- chardsoni Brown, are now based entirely on speculation. That all forms of pest control— biological as well as chemical—pose some environ- mental risk is well known (Taylor 1955, El- ton 1958, Turnbull and Chant 1961, Ehler and van den Bosch 1974, Beirne 1975, Pi- mentel et al. 1984, Carruthers and Onsager 1993, Drea 1993, Miller and Aplet 1993). Consequently, biological control practition- ers generally, especially those involved with weeds, have attempted to minimize poten- tial environmental problems (e.g. Zw6lfer and Harris 1971, Harris 1973, Goeden 1983, Klingman and Coulson 1983). It is tempting to look at the case of C7 and C9 within the context of classical bio- logical control. Even though C7 was reco- lonized extensively by biological control specialists, this project does not typify clas- sical biological control. Initially there was no target pest (e.g. Comis and Heppner 1986); only later was the Russian wheat aphid identified as the target aphid for re- distribution. It is also uncertain whether the successful invasive genotype of C7 should be attributed to intentional releases in North America or to an accidental introduction with commerce. If C7 has indeed adversely affected C9, that interaction is more appro- priately viewed as displacement of an in- digenous species by a polyphagous, aggres- sive nonindigenous species. It should not be cited as an example of negative effects of classical biological control. Instead it reem- phasizes the continuing need to assess host (and prey) specificity of all agents consid- ered for release in classical biological con- trol programs. ACKNOWLEDGMENTS We are grateful to M. W. Brown, F. A. Drummond, D. Ellis, P. M. Estes, J. J. Kno- del, R. L. Brown, P. W. Schaefer, C. L. Staines, Jr., and W. Y. Watson for allowing us to refer to their unpublished data or cor- respondence; E. C. Becker, S. W. Bullington, D. S. Chandler, J. R. Coulson, W. H. Day, L. E. Ehler, R. D. Gordon, K. I. Hauschild, L. A. Hull, W. C. 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Abstract.—The life history of Hydrometra hungerfordi Torre-Bueno was investigated in southern Illinois during April-November 1992 and 1993, and the immature stages were described. The bug was reared from egg to adult under controlled laboratory con- ditions. This apparently bivoltine species occurred along the stream banks on the water surface, root mats, and floating detritus. Adults overwintered in leaf litter and became active in April. Eggs were found in May, June, and July above the water level in root mats, tree snags, and associated detritus along the stream bank. Seasonal occurrences of adults and nymphs are discussed. Adults were last observed in early November. This species was reared under a 16L:8D photoperiod and 28 + 1°C. The incubation period averaged 8.81 days. Durations of the five nymphal stadia averaged 2.52, 2.48, 2.68, 3.39, and 4.68 days, respectively. Key Words: Water measurer, Hydrometridae, southern Illinois, life history, laboratory rearing, immature stages, descriptions The family Hydrometridae, which con- tains about 110 species and seven genera (Smith 1988) in three subfamilies (China and Usinger 1949, Smith 1988), occurs worldwide, although most species are found in the tropics and subtropics (Smith 1988). The largest of the subfamilies, Hydrome- trinae, also occurs worldwide and includes the genus Hydrometra, the only genus found in America north of Mexico (hereafter re- ferred to as North America). Hydrometra is represented in North America by seven to nine species; H. australis Say may be con- ' Part of a thesis submitted to Southern Illinois Uni- versity at Carbondale by the senior author in partial fulfillment of the requirements for the M.S. degree in zoology. specific with H. martini Kirkaldy, and the supposed occurrence of H. /illianis Torre- Bueno, reported only from California, may be in error (Polhemus and Chapman 1979). Hydrometrids, commonly called marsh treaders or water measurers, are fragile-ap- pearing stiltlike insects with long threadlike legs and antennae, slender bodies, and elon- gate heads. In North America, they are gen- erally found on or near emergent or floating vegetation associated with still or slowly moving bodies of freshwater, but may move onto open water if disturbed. H/. australis has occasionally been found in saltwater tide pools in Florida (Herring 1949) and in brackish ponds in Texas (Polhemus and Chapman 1979). Marsh treaders are pre- daceous, feeding on dead or dying insects PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1d other microinvertebrates on or within the water film or on floating vegetation (Smith 1988). Most of the North American species have a limited distribution, the exceptions being H. martini, H. australis, H. wileyae Hun- gerford, and H. hungerfordi Torre-Bueno. Not surprisingly, the species with the widest distribution, H. martini, is also the species that has been most intensively studied (Pol- hemus and Chapman 1979). H. martini occurs from Quebec and Maine south to Florida and west to British Colum- bia, Oregon, and Arizona (Smith 1988). If H. australis is conspecific, then it has also been collected in California, Mexico to Cen- tral America, and the West Indies (Smith 1988). The most thorough studies of H. martini are those of Hungerford (1920) and Sprague (1956). Sprague (1956), Polhemus and Chapman (1979), and Bennett and Cook (1981) provide excellent reviews of the ear- lier literature. Adults of H. martini overwinter in trash surrounding the water, and emerge in early spring (Hungerford 1920). The eggs are about 2 mm long, slender, spindle shaped, sculptured, and stalked. They are laid singly on almost any object above the water sur- face, and each is affixed at the base of the stalk with a sticky secretion. Eggs are white at oviposition but soon darken. There are five instars, as in most Heter- optera. Hungerford (1920) stated that the complete life cycle from egg to egg, under the most favorable conditions, was 15 days. Torre-Bueno (1905) and Polhemus and Chapman (1979) stated, respectively, that 25 to 35 days and 21 to 35 days was typical. Adults generally live seven to nine months under laboratory conditions (Sprague 1956), although Torre-Bueno (1905) reported adults in his laboratory lived up to one year, and one lived 15 months. H. martini has been reared in the labo- ratory under unspecified conditions (Martin 1900, Torre-Bueno 1905, Hungerford 1920, Sprague 1956). The egg was first described and illustrated by Martin (1900); it and the instars were subsequently described and il- lustrated by Hungerford (1920) and Sprague (1956). H. hungerfordi, as noted above, is also widely distributed. It occurs from Maine south to Florida and west to Kansas and Louisiana (Smith 1988). However little is known about its biology. It has been found in heavily shaded, cool, clear streams with little or no aquatic vegetation (Gonsoulin 1973, Harp 1985) and in slow moving streams, rivers, and swamps (Bobb 1974). Adults have been collected from May through August (Bobb 1974). This paper presents information on the field life history in southern Ilinois and lab- oratory rearing of H. hungerfordi and in- cludes descriptions of the immature stages. MATERIALS AND METHODS Study site.—In summer 1991, a popula- tion of H. hungerfordi was discovered on Cooper’s Creek in Alexander Co., IL. The numbers observed and accessibility of the site suggested a life history study was pos- sible. Therefore, a study was conducted from spring 1992 to fall 1993. The study site is located in the Mill Creek quadrangle 7.5’ topographic (T145, R1IW, N'% of N'% of NW of Sec. 6). Cooper’s Creek at this point is classified as a third order stream (Horton 1945). The substrate consists of cobble and gravel with some sand. The width and depth at the center of the channel during spring 1993, when flow was greatest, averaged 6.0 and 0.6 m, respec- tively. During midsummer, as the water level receded, the stream was reduced to isolated pools. The banks were generally steep sided and undercut, with exposed root mats. Very little emergent vegetation was present. The site was bounded by a riparian zone containing Acer negundo L., Betula ni- gra L., Celtis occidentalis L., Juglans nigra L., Ostrya virginiana (Miller), Prunus ser- otina Ehrhart, Carya cordiformis (Wangen- VOLUME 97, NUMBER 3 heim), Staphylea sp., Ranunculus sp., Po- lygonum sp., Plantago sp., Cirsium sp., and Dicentra sp. Much of this zone was limited to a width of 10-15 m on either side because of agricultural use of the surrounding land. Life history.—Samples of 10 individuals (adults and/or nymphs) were taken weekly from early spring through early fall. Speci- mens were hand picked or, for early instars, collected with an aspirator. The few eggs collected (11) were not included in the counts. Adults were identified and released. Immatures were preserved in 75% EtOH and taken to the laboratory for closer ex- amination and to determine the instars. Confirmation of instars was based on com- parisons with laboratory reared specimens because published keys were not available for identification. Prey items were also pre- served in 75% EtOH. Laboratory rearing.—Ten pairs of adults were collected from the field and placed in 1 qt Mason jars (1 2, | °/jar). Each jar con- tained about 200 ml of distilled water and a section of Styrofoam (2 x 1.5 x 1 cm); this floating square provided an additional substrate for walking and served as an ovi- position site. The jar was closed with wire screen and secured with the band of the 2-piece mason jar lid. Each pair was fed S— 6 frozen Drosophila melanogaster Meigen daily, and the carcasses were removed the following day. The squares were examined daily for eggs. Eggs were removed and placed in covered petri dishes (about 9 cm diam, 2 cm deep) lined with moistened filter paper on the bot- tom; distilled water was added as needed to keep the paper moist. Each first instar was placed in a round plastic container (4 cm diam, 5 cm deep). Each container was filled with about 2 cm of distilled water. A wedge of Styrofoam (4 x 3 x 2.cm, 30° angle) was added to serve as an additional substrate for walking. The container was closed with standard fiberglass screening secured with a rubber band. As the immatures developed, molts were recorded and the exuviae re- 719 moved. First through third instars were fed 2 fruit flies, and fourth through fifth instars 3-4 fruit flies, daily. The containers were kept in incubators maintained at 28 + 1°C and a photoperiod of 16L:8D (3 florescent “daylight” lamps, about 2800 lux). Distilled water was added to the mason jars and con- tainers as needed to maintain water levels. Descriptions of immature stages.—The description of each stage is based on 10 in- dividuals. Eggs were selected from those de- posited in the laboratory by field collected adults, first through fifth instars from field collected individuals. All had been pre- served in 75% EtOH. Drawings were made with the aid of a camera lucida, measure- ments with an ocular micrometer. Dimen- sions are expressed in mm as X + SE (SE < 0.005 listed as 0.00). RESULTS AND DISCUSSION Life history. —H. hungerfordi overwinters as adults. The one adult (female) collected during the winter was found in leaf litter about 2 m from the water edge in early De- cember 1992. Adults emerged in mid-April (Fig. 1A, B); air temperature when they first were Observed (11 April 1992), based on data collected at the Anna, Alexander Co., weather station, ranged between 14 and 27°C during the 24 h. They were found contin- uously during the season near the banks on the detritus and root mats associated with the banks and undercut areas. They and nymphs were never observed in the open channel. Copulating adults were found throughout the season. Precopulatory behavior was noted on eight occasions; at no time was the male observed to antennate the female. He usually approached her from the side until he was almost in contact and then reorient- ed until his body was parallel to hers. Upon contact, he would mount her from the side, grasp her with his pro- and mesothoracic legs, extend his genital capsule, and lower the tip of his abdomen to make contact with her genitalia. Pairs were observed in copulo PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1ST INSTAR (N = 18) 2ND INSTAR (N = 18) 3RD INSTAR (N = 33) 4TH INSTAR (N = 28) PERCENT OF INDIVIDUALS 5TH INSTAR 1ST INSTAR 2ND INSTAR 3RD INSTAR 4TH INSTAR PERCENT OF INDIVIDUALS 5TH INSTAR Fig. 1. Field life cycle of Hydrometra hungerfordi. A, Percent of individuals of each stage per sample during 1992 season in Alexander Co., IL. B, Percent in each sample of total individuals of same stage collected during the 1992 season in Alexander Co., IL. VOLUME 97, NUMBER 3 up to 5-6 minutes. During this time, fe- males would often move about and feed if prey were located. On one occasion, a cop- ulating male was physically dislodged by an- other male. The second male then attempt- ed, repeatedly, to mount the female. She prevented his attempts by dislodging him with her legs and finally left the area. Eggs were found in early May (4), late June (2) and mid-July (5). They were laid on root mats, tree snags, and associated de- tritus. Generally, they were laid 6-10 cm above the water surface in dark, moist areas and always affixed to the substrate by the base of the egg. First instars were found from mid-May to mid-August, second instars from late May to late August, third instars from late May to mid-September, fourth instars from early June to late September, and fifth instars from mid-June to early October (Fig. 1A, B). First and second instars were often found in the back reaches of the undercut bank. If they were disturbed when in more exposed areas, they would invariably flee beneath the bank. Third, fourth, and fifth instars were generally found in the more open ar- eas, but never far from the bank. H. hungerfordi, as evidenced by the peaks in abundance of the adults and nymphs, is apparently bivoltine in southern Illinois (Fig. 1A, B). Several prey items were noted during this study (Table 1). The hydrometrids were ob- served to actively capture small prey (i.e., collembolans), but only noted feeding on larger prey. H. hungerfordi adults and nymphs were occasionally observed attempting to feed on nymphs. In one instance, an adult was noted antennating a third or fourth instar. It then extended its beak and attempted to probe the nymph, which then fled. As H. hungerfordi walked across the sur- face film and floating detritus, it moved its antennae from side to side. When it en- countered potential prey, it positioned its antennae over the specimen. If the speci- 721 Table 1. Feeding records of Hydrometra hunger- fordi from a stream in Alexander Co., IL, in 1992 and 1993. No. of Taxon specimens Collembola Isotomidae Isotomurus palustroides Folsom# 2D, Entomobryidae Entomobrya socia Denis* 1 Tomocerus flavescens Tullberg* Ephemeroptera Leptophlebiidae Paraleptophlebia sp.* 1 Coleoptera Dytiscidae Hydaticus sp.” 1 Diptera Culicidae Anopheles crucians Wiedemann? 1 Chironomidae Polypedilum sp.” 1 Cyprinodontiformes Fundulidae Fundulus sp.° 1 a Adult. > Larva. ° Egg. men was acceptable, the beak was extended and used to probe and then pierce the prey. If the prey was small, the hydrometrid lifted the now impaled prey and, generally, car- ried it near or onto the stream bank. The use of the antennae to locate prey has also been reported in other hydrometrid species by Andersen (1982). He noted an invagination at the apex of the fourth an- tennal segment bordered by modified hairs and suggested this area may have an olfac- tory function. An encounter between an adult H. hun- gerfordi female and a lycosid spider was ob- served on one occasion. The spider grabbed and pulled the female towards its mouth and then immediately flung it away. The female was observed for about 10 minutes and appeared unharmed by the encounter. Sprague (1956) felt there probably was little or no predation on adult H. martini because of the hard exoskeleton, fusion of 122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Duration (in days) of each immature stage of Hydrometra hungerfordi under controlled laboratory conditions. Number Cumulative Stage completing stadium Range 39.25 Sia mean age Egg* 181 8-10 Seslee=tOLOS 8.81 Nymph Ist instar 178 1-4 IV) SE (0055 11.33 2nd instar 178 1-6 2.48 + 0.05 13.81 3rd instar 143 2-4 2.68 + 0.06 16.49 4th instar 103 2-5 3.39 + 0.08 20.88 Sth instar 41 3-7 4.68 + 0.13 25.56 a 200 eggs were laid. body parts, and lack of thin intersegmental membranes. H. hungerfordi usually flees when dis- turbed. If it is unable to do so, it often will drop to the surfaces of the detritus and root mats with the prothoracic legs and antennae directed anteriorly and the meso- and me- tathoracic legs directed posteriorly, all ap- pendages in line with the body, and remain motionless. This behavior was noted on seven occasions. During these times, the an- imal was difficult to see. Laboratory rearing.—Eggs were laid 2-3 cm above the water surface on the walls of the mason jars and on the upper surface of the Styrofoam. They were laid singly with the base of the stalk attached to the surface. The female would touch the tip of her ab- domen to the surface and then slowly move away as she extruded the egg. The egg was white when first laid, but darkened to light brown within eight hrs. Eye spots were visible within five days. The egg burster was visible one to two days later. The incubation period averaged 8.81 days (Table 2). The first instar emerged through a lon- gitudinal slit along one side at the cephalic end of the egg. Emerging nymphs were transparent except for their red eyes. They were observed feeding within three to four hours. The first through fifth stadia averaged 2.52, 2.48, 2.68, 3.39, and 4.68 days, re- spectively (Table 2). The total developmen- tal period averaged 25.56 days. Highest mortality ocurred during the fifth stadium and resulted primarily from incomplete ec- dysis. Descriptions of immature stages.—Egg (Fig. 2A—D): Length, 1.90 + 0.02; width, 0.28 + 0.01. Egg fusiform; comprised of distal micropylar projection, central region, and basal pedicel. Eggs laid singly, glued at base of pedicel to living or dead plant ma- terial; white at oviposition, becoming gold to brown during maturation. Chorion of mi- cropylar projection and of most of pedicel with scalelike sculptering; each scale appar- ently with several pores. Apex of micropylar projection with single micropyle. Central region with alternating ridges and furrows. Nymphal instars: The first instar is de- scribed in detail, but only major changes from previous instars are described for sub- sequent instars. Length is measured from tip of head to tip of abdomen, width across pronotum. Additional measurments are given in Table 3. First instar (Fig. 3A): Length, 1.70 = 0.07; width, 0.24 + 0.01. Body slender, elongate, greatest width at prothorax; gen- eral ground color of head and thorax light brown dorsally and laterally, yellowish to white ventrally; ground color of abdomen usually yellowish to white on all surfaces; ecdysial line evident dorsomedially, yellow- ish white, arising at posterior margin of head, bifurcating near anterior margin of eyes; body sparsely setose dorsally and laterally, VOLUME 97, NUMBER 3 100 Fig. 2. Scanning electron micrographs of egg of Hydrometra hungerfordi. A, Egg. B, Micropylar region. C, Central region, D, Basal region. more heavily setose ventrally except on head _ teocular area subrectangular, constricted just posteroventral to eyes where setae are ab- anterior to eyes, area about 1.4 length of sent. postocular area; postocular area narrowing Head scelerotized, elongate, narrow; an- anteriorly. Anteclypeus rectangular, sub- Table 3. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Measurements (mm)? of Hydrometra hungerfordi instars?. Nymph Ist instar 2nd instar 3rd instar 4th instar 5th instar Body lengths 1.70 + 0.07 eM ae (OI) 3.54 + 0.13 46210515 12/22 WENO Head length* 0.61 + 0.01 0.85 + 0.02 ET OVON 167 + 0:03 2.45 + 0.03 Anteocular 0.28 + 0.01 0.43 + 0.01 0.63 + 0.01 0.95 + 0.02 1.47 + 0.02 Postocular 0.20 + 0.00 0.28 + 0.01 0.44 + 0.02 0.54 + 0.01 0.76 + 0.01 Width across eyes 0.23 + 0.00 OSE 0:01 0.30 + 0.01 O35) =) 0201 0.43 + 0.01 Synthlipsis 0.09 + 0.00 0.09 + 0.00 0.10 + 0.00 OTL = O01 0.12 + 0.01 Antennal segs. Ist 0.14 + 0.00 0.18 + 0.00 0:22-0:00 0.30 + 0.00 0.38 + 0.01 2nd 0.19 + 0.00 0.29 + 0.01 0.39 + 0.01 0.56 + 0.01 0.80 + 0.01 3rd 0.73 + 0.02 1.06 + 0.01 1.45 + 0.02 1.88 + 0.02 2.42 + 0.00 4th 0.86 + 0.01 0.99 + 0.01 Le13)-£/ 0/01 1E267== OO 1.41 + 0.01 Notal lengths* Pronotum Ons 0/01 0.20 + 0.01 0.28 + 0.00 0.44 + 0.00 0.74 + 0.01 Mesonotum 0.14 + 0.01 0:21 2:10:01 0.30 + 0.01 0.40 + 0.01 0.42 + 0.01 Metanotum 0.07 + 0.00 0.09 + 0.00 0.10 + 0.01 0.23 + 0.01 OF527==) O01 Width at pronotum 0.24 + 0.01 0.26 + 0.01 0.28 + 0.01 0.34 + 0.01 0.40 + 0.01 Abd. length 0.71 + 0.06 0.96 + 0.10 las) e= (OLN? DD, =O ale 3.84 + 0.09 Leg lengths Profemur 0.56 + 0.01 0.79 + 0.01 1.09 + 0.01 1.48 + 0.03 2-16) 40:03 Protibia 0.74 + 0.01 1.04 + 0.01 1.43 + 0.02 1.93 + 0.02 2.74 + 0.02 Protarsus 0.27 + 0.00 0.30 + 0.01 0.36 + 0.01 0.44 + 0.01 0.59 + 0.01 Mesofemur 0.58 + 0.01 0.85 + 0.01 Nk se (OXON 1:66 = 0103 2.45 + 0.02 Mesotibia 0.78 + 0.01 5 ex (OO 12567==7 0:0) Dal2s==) O03 3.02 + 0.02 Mesotarsus 0.27 + 0.00 0.31 + 0.00 0.39 + 0.01 0.48 + 0.01 0.64 + 0.01 Metafemur 0.78 + 0.01 LalSe== 10102 1495 On 2.19 + 0.04 3:14 70103 Metatibia 1.08 + 0.02 1.60 + 0.03 2235+ 0:02 3.07 + 0.05 4.33 + 0.03 Metatarsus 0.27 + 0.00 OFS 10/01 0.38 + 0.01 0.47 + 0.01 0.62 + 0.01 a3%e-t SE: > Based on 10 individuals per instar. © Measured at midline. truncate apically, extending beyond bases of antennae. Area ventrolateral of antecly- peus and anterior to antennal sockets not readily divisible into maxillary plate and ventral lobe. Labrum narrow, transverse, located just anterior to anteclypeus. Beak 4-segmented; segments | and 2 short, nar- row, subequal in length; segment 3 longest of segments, about 6.0 x length of segment 1 and 2 combined; segment 4 about 0.4 x length of segment 3. Eyes red, granular, lo- cated laterally about midway along length of head. Three pairs of trichobothria; first pair at base of anteclypeus close to midline, second pair dorsolateral and posterior to an- tennal bases; third pair dorsolateral, near posterior margin of head; each trichoboth- rium arising from tubercle. Antennae 4-segmented, filiform, segment 4 slightly fu- siform distally; segment | shortest; ratio of antennal segment lengths about 1:1.3:5.1:6. Thoracic nota scelerotized; middorsal, longitudinal ecdysial line continuous with that of head. Pronotum quadrangular; an- terior margin overlapping posterior margin of head; posterior margin straight laterally, convex medially. Mesonotum at midline about 0.8 x length of pronotum; subquad- rangular; posterior margin slightly convex. Metanotum at midline about 0.5 length of mesonotum; rectangular; posterior mar- gin slightly concave. Thoracic pleura visa- VOLUME 97, NUMBER 3 725 Fig. 3. Nymphal stages of Hydrometra hungerfordi. A, First instar. B, Second instar. C, Third instar. D, Fourth instar. E, Fifth instar. 726 e in dorsal view, particularly those of neso- and metathoraces; each pleuron di- ided by suture into episternum and epi- meron. Thoracic sterna lightly sclerotized; sclerotization best developed on proster- num; prosternum narrowest of thoracic sterna. Thoracic spiracles present; spiracle | present on proepisternum adjacent to in- tersegmental line; spiracle 2 present on me- soepisternum near intersegmental line. Legs light brown; coxae subconical; tro- chanters cylindrical; femora, tibiae, and tar- si slender, elongate, each increasing in length on pro-, meso-, and metathoraces, respec- tively; tarsi 1-segmented, each terminating in two claws. Abdomen yellowish to white, occasion- ally with red, red often present as dorsolat- eral stripes; tergum | with subtrapezoidal tergal plate; terga 2-6 membranous; terga 7-10 with dorsal surface sclerotized, scle- rotization on terga 9-10 occasionally ex- tending to and including ventral surface. Sternum | intimately associated with meta- sternum; sterna | and 2 not readily distin- guishable from each other; sterna 3-8 mem- branous; sterna 9-10 membranous or scler- otized. Spiracles present on segments 1-8; those on segment | located dorsolaterally, those on segments 2-8 ventrolaterally. Second instar (Fig. 2B): Length, 2.31 + 0.10; width, 0.26 + 0.01. Anteocular area widest at antennal bases, narrowest just an- terior to eyes; about 1.5 x length of post- ocular area; postocular area narrowest at posterior margin of eyes, widening to base of head; area ventrolateral of anteclypeus partially divided into maxillary plate and ventral lobe; maxillary plate adjacent to an- teclypeus, dorsal to ventral lobe. Ratio of antennal lengths about 1:1.6:5.9:5.5. Pronotum with posterior margin sub- truncate, overlapping anterior margin of mesonotum. Mesonotum at midline about 1.1 x length of pronotum; rectangular; pos- terior margin subtruncate, overlapping an- terior margin of metanotum. Metanotum at midline about 0.4 length of mesonotum; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rectangular, posterior margin more con- cave; wing pads evident at posterolateral corners. Abdominal tergum | with 2 sclerotized plates; anterior plate narrow, transverse, constricted medially; posterior plate trans- verse, elliptical. Terga 2—7 with paired small, transverse, linear plates adjacent to inter- segmental lines. Third instar (Fig. 3C): Length, 3.54 + 0.13; width, 0.28 + 0.01. Anteocular area about 1.4 length of postocular area. Area ventrolateral to anteclypeus divided into maxillary plate and ventral lobe. Area pos- teroventral to eyes with few, scattered setae. Ratio of antennal segment lengths about tleS:G:5 35516 Pronotum narrower, rectangular. Meso- notum at midline about 1.1 x length of pro- notum; rectangular; posterior margin con- cave; wing pads well developed, slightly overlapping metanotum. Metanotum at midline about 0.3 x length of mesonotum; posterior margin strongly concave; wing pads well developed. Abdominal tergum 1 with 3 sclerotized plates; first 2 as anterior pair, small, oblique; third plate posterior, transverse, elliptical. Fourth instar (Fig. 3D): Length, 5.46 + 0.15; width, 0.34 + 0.01. Ecdysial line still linear posterior to eyes, broader between and anterior to eyes to basal 4 of anteocular length before bifurcating. Setae more nu- merous posteroventral to eyes. Anteocular area about 1.8x the length of postocular area. Ratio of antennal segment lengths about 1:1.9:6.3:4.2. Pronotum with posterior margin round- ed, slightly overlapping mesonotum. Meso- notum at midline about 0.9 x length of pro- notum; wing pads well developed, extend- ing over and usually covering metanotal wing pads. Metanotum at midline about 0.6x length of mesonotum; sclerotized or membranous; wing pads often hidden by mesonotal wing pads, occasionally exposed apically. Abdominal tergum 1 with 3 sclerotized VOLUME 97, NUMBER 3 plates; first two as anterior pair, small, oblique; third plate posterior, weakly de- veloped or absent. Abdominal terga 1-6 with continuous, white, mediolongitudinal stripe bordered on either side by red stripe. Ter- gum 7 with rectangular, sclerotized plate. Tergum 8 with medial projection on pos- terior margin of sclerotized plate; projection generally larger in females. Sternum 8 oc- casionally with small, medial plate of vari- able shape. Fifth instar (Fig. 3E): Length, 7.97 + 0.10; width, 0.40 + 0.01. Body more setose. Anteocular area about 1.9 x length of post- ocular area. Ventral lobe more developed, reaching beak segment |. Ratio of antennal segment lengths about 1:2.1:6.4:3.7. Pronotum with posterior 3 narrowing posteriorly, rounded apically. Mesonotum at midline about 0.6 x length of pronotum; wing pads covering metanotal wing pads. Metanotum at midline about 1.2 length of mesonotum; membranous other than wing pads. Tergum 8, in males, with dorsomedial projection that does not exceed tip of ab- domen. Tergum 9 narrow, collarlike, ex- tending and widening laterally where it reaches abdominal sternum 9. Sternum 7 with pair of longitudinal, rectangular plates. Sternum 8 with quadrangular plate, no split present. Sternum 9 large, cuplike. Sternum 10 either lost or associated with formation of ventral side of proctiger. Tergum 8, in females, with dorsomedial projection that extends to or beyond ab- dominal tip. Sternum 7 with pair of longi- tudinal, rectangular plates. Sternum 8 with quadrangular plate that reaches lateral mar- gin; plate with mediolongitudinal split; split varying from complete to limited to distal Y), widening posteriorly. Sternum 9 with pair of small quadrangular plates; plates con- tinuing laterally and dorsally as sclerotized ring. Sternum 10 either lost or associated with formation of ventral side of proctiger. Diagnosis.— The five nymphal instars, in addition to size, can be readily separated by 727 the relative proportion of the anteocular and postocular areas; presence or absence, and degree of development, of wing pads; and length of metafemora. There is a progressive increase of the anteocular area relative to the postocular area through all the instars. Wing pad development becomes apparent in the second instar with progressive de- velopment in the subsequent instars: meso- notal wing pads begin to overlap the meta- notum in the fourth instar and completely cover the metanotal wing pads by the fifth instar. Finally, all leg segments progressive- ly increase in length through the instars, but increase is most readily seen in the meta- femur. ACKNOWLEDGMENTS We thank the following faculty and staff members of Southern Illinois University at Carbondale: J. A. Beatty and W. G. Dyer, Department of Zoology, for their critical re- views of the manuscript; and the entire staff of the Research Photography and IIlustra- tion Facility, for their technical expertise and preparation of the final drafts of the figures. We also thank R. J. Snider, De- partment of Zoology, Michigan State Uni- versity, Lansing; B. M. Burr and J. B. Stahl, Department of Zoology, SIUC; and D. W. Webb, Illinois Natural History Survey, Champaign, for identification of prey items; and R. W. Sites and Becky J. Nichols, De- partment of Entomology, University of Missouri, Columbia, for the SEM photo- graphs of the eggs. LITERATURE CITED Andersen, N. M. 1982. The semiaquatic bugs (He- miptera: Gerromorpha). Phylogeny, adaptations, biogeography and classification. Entomonograph, Vol. 3. Scandinavian Science Press, Klampenborg, Denmark. 455 pp. Bennett, D. V.and E. F. Cook. 1981. The semiaquatic Hemiptera of Minnesota (Hemiptera: Heterop- tera). Minnesota Agricultural Experiment Station Technical Bulletin 332: 1-59. Bobb, M. L. 1974. The insects of Virginia: No. 7. The aquatic and semi-aquatic Hemiptera of Vir- 728 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ginia. Virginia Polytechnic Institute and State University Research Division Bulletin 87: 1-195. China, W. E. and R. L. Usinger. 1949. A new genus of Hydrometridae from the Belgian Congo, with a new subfamily and a key to the genera. Revue de Zoologie et de Botanique Africans 41: 314-319. Gonsoulin, G. J. 1973. Seven families of aquatic and semiaquatic Hemiptera in Louisiana. Entomolog- ical News 84: 9-16. Harp, G. L. 1985. A synopsis of the Hydrometridae of Arkansas. Proceedings of the Arkansas Acad- emy of Science 39: 130-131. Herring, J. L. 1949. Taxonomic and distributional notes on the Hydrometridae of Florida. Florida Entomologist 31: 112-116. Horton, R.A. 1945. Erosional development of streams and their drainage basins: hydrophysical approach to quantitative morphology. Bulletin of the Geo- logical Society of America 56: 275-370. Hungerford, H. B. 1920. The biology and ecology of aquatic and semiaquatic Hemiptera. University of Kansas Science Bulletin 11: 3-328 (1919). Martin, J. O. 1900. A study of Hydrometra lineata. Canadian Entomologist 32: 70-76. Polhemus, J. T. and H. C. Chapman. 1979. Family Hydrometridae/marshtreaders, water measurers, pp. 43-45. In Menke, A. S., ed., The Semiaquatic and Aquatic Hemiptera of California (Heterop- tera: Hemiptera). Bulletin of the California Insect Survey 21: xi + 1-166. Smith, C. L. 1988. Family Hydrometridae, pp. 156- 158. In Henry, T. J. and R. C. Froeschner, eds., Catolog of the Heteroptera, or True Bugs, of Can- ada and the Continental United States. E. J. Brill, Pub., New York. 958 pp. Sprague, I. S. 1956. The Biology and morphology of Hydrometra martini Kirkaldy. University of Kan- sas Science Bulletin 38: 579-693. Torre-Bueno, J.R. 1905. Notes on Hydrometra mar- tini. Canadian Entomologist 37: 12-14. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 729-731 NOTE Distribution of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in North Carolina and Virginia The multicolored Asian lady beetle, Har- monia axyridis (Pallas), was first reported to be established in the United States in Louisiana and Mississippi (Chapin and Brou. 1991. Proceedings of the Entomolog- ical Society of Washington 93: 630-635). This entomophagous lady beetle feeds on a wide variety of aphids, especially arboreal aphids, as well as some coccids and other insects (Tedders and Schaefer. 1994. En- tomological News 4: 228-243). Harmonia axyridis was released in the United States as early as 1916 in California, and between 1978 and 1982, large numbers were released in the United States and Canada. The ma- jority were released along the Atlantic Sea- board, including over 87,000 released near Byron, Georgia for control of aphids on pe- can, but releases were also made in Loui- siana, Mississippi, and Washington state (L. R. Ertle, in litt.). The first collections of H. axyridis in North Carolina occurred in the fall of 1992. Reports of lady beetles aggregating on and inside houses, churches, and other struc- tures were received through the Coopera- tive Extension Service, North Carolina State University. Specimens were subsequently identified by D. L. Stephan of the Plant Dis- ease and Insect Clinic, NCSU. By Decem- ber, reports had been received from 7 coun- ties, all but one of which is in the mountain or western piedmont region of the state (Fig. 1). During the spring and summer of 1993, individual beetles were collected, coinci- dental with other field work, on a variety of host plants and at lights in 7 counties in NC. Fall aggregations had been reported from 2 of these counties, Cleveland and Clay (Fig. 1). Host plants in North Carolina in- cluded thistle (Cirsium vulgare) (Savi) Te- nore, ironweed (Vernonia sp.), Erigeron strigosus Muhl. ex Willd., dock (Rumex sp.), wheat (Triticum sp.) and cotton (Gossypium sp.). Unidentified aphid species were seen on wheat and dock, but insect hosts were not always found associated with the lady beetles. The beetles may have been feeding on nectar from floral or extrafloral nectaries or pollen; these substances are commonly fed upon by coccinellids (Hagen. 1962. An- nual Review of Entomology 7: 289-326). First reports of aggregations in Virginia came from Lee County in January 1993. Har- monia axyridis was collected 3 August 1993 in alfalfa during a coccinellid survey in Au- gusta County, VA. This is approximately 375 km (as the crow flies) from the first collection reported in Virginia. Lady beetles were also observed feeding on Myzus per- sicae (Sulzer) on tobacco (Nicotiana sp.) in Nottoway County, VA. In anticipation of fall 1993 aggregations, county offices of the North Carolina Co- operative Extension Service were contacted and asked to report activity. Initial reports were received from the western counties in mid-October and continued through the end of November, with activity shifting from west to east. On Monday 18 October, coun- ty agents from Macon County, NC to Rap- panhannock County, VA (a distance of about 630 km) were innundated with reports of aggregations that suddenly appeared the previous day. In the piedmont regions no- ticeable activity began 25 October in VA and 30 October in NC. Although to date beetles have been reported in eastern VA (to 76°30'W longitude), no reports have been received from east of about 78°W in NC. Based on reports of homeowners and oth- er observers, aggregating lady beetles came suddenly in large numbers, and landed on buildings, apparently searching for suitable 730 1992 Aggregations E —— 77/7 1993 Aggregations 1992-93 Aggregations 1992 Summer Collections Fig. 1: overwintering sites. After the initial ap- pearance, most of the beetles disappeared within a short time, but others were seen on walls or windows for several days, becoming active as the temperature warmed during the day. In Alleghany County, NC, a man was packing his car 17 October, which re- quired periodic trips into his house. In an interval of about 15 minutes, the white doors of a two-car garage became covered with **20-40,000” beetles. He continued packing and watched the beetles. Within 1.5 hours, most beetles had left, but their ultimate des- tination was unknown. A few were observed inside the garage and on the outside trim 26 October. A woman in Henderson Coun- ty, NC, described the activity as “‘like a swarm of bees” as the beetles left a stand of large white pines to the east of her house. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 7 Distribution of Harmonia axyridis Pallas in Virginia and North Carolina, 1992-1993. Beetles were most often found on light- colored walls and trim or on reflective sur- faces such as large windows. Usually they were found on the southwest, west, or south- facing sides of buildings, which were the sunniest areas during the warmer afternoon hours. A few of the houses where H. axyridis aggregated were close to others in subdivi- sions or residential areas, but often were on a rise or hill and isolated with open spaces on one or more sides. Reported overwin- tering sites included: inside houses on vault- ed or cathedral ceilings, under doorframes, in insulated attics, behind molded plastic shutters on houses, in soffit vents, and in root cellars. The beetles are able to get into small cracks and crevices, and undoubtedly Overwinter in many other places, undetect- ed. In Japan, H. axyridis reportedly over- VOLUME 97, NUMBER 3 winters in many places including the cracks and crevices of rocks on mountains, with a tendency to go toward white or light colored objects (Obata. 1986. Kontyu 54: 218-223). Tedders and Schaefer (1994) showed that white traps were more effective at attracting this species than dark, less reflective colors. We observed a variety of plants associated with the aggregation sites, with white pines, loblolly pines, Fraser firs, and apple trees found most frequently. This species is polymorphic with melanic and non-melanic forms (Komai. 1956. Ad- vances in Genetics 8: 155-188, Ayala. 1978. Scientific American 239: 56-69). All H. ax- yridis collected in North Carolina were the non-melanic form, and elytral color varied from brick-red to brownish-yellow. Elytral spots varied in number from 0 to 20 and in size from small dots to large, sometimes fused spots. This supports the findings of Chapin and Brou (1991) in Louisiana and Mississippi, and Tedders and Schaefer (1994) in Georgia, Florida, Alabama, and South Carolina populations. Since the first reports of H. axyridis in NC and VA, the lady beetle has become widespread across these states. We can only assume that it will continue to spread throughout both states within a few years. With its wide host range and preference for arboreal aphids, it has the potential to pro- vide beneficial biological control, but its 731 habit of aggregating inside houses and other buildings is viewed by many residents to be a nuisance. Acknowledgments: We thank the nu- merous county agents of the Cooperative Extension Service who assisted in the col- lection of beetles and information and the many homeowners who shared anecdotes and allowed us to collect in their homes. Kathleen A. Kidd & Christine A. Nalepa, Biological Control Laboratory, NC Depart- ment of Agriculture, P.O. Box 27647, Ra- leigh, NC 27611; Eric R. Day, Department of Entomology, Price Hall, Virgina Poly- technic Institute and State University, Blacksburg, VA 24061-0319: Michael G. Waldvogel, Department of Entomology, Box 7613, North Carolina State University, Ra- leigh, North Carolina 27695-7613. Note Added in Proof In the summer of 1994, adults of H. axy- ridis were found in cotton fields in the coast- al plain (Perquimans County in the north- east and Onslow County in the southeast) of North Carolina. In the fall of 1994, ag- gregations were reported in the western two- thirds of the state, but activity remains high- est in the far western counties. Based on the estimates of homeowners, aggregating bee- tles were more numerous than in the past. PROC. ENTOMOL. SOC. WASH. 97(3), 1995, pp. 732-734 NOTE Validation of nomina nuda of Nearctic Tethinidae, Scathophagidae, and Muscidae proposed in Manual of Nearctic Diptera In the Manual of Nearctic Diptera, Vol- ume 2 (1987), I proposed several new ge- neric and specific names that were not then validated. Three were combinations of new generic and new specific names (one in Tethinidae, two in Scathophagidae), which were not validated with the term “n.g., n.sp.” or the equivalent, as required by Article 13(c) of the International Code of Zoological No- menclature. One was a specific name in the Muscidae. These names are validated here and type material is recorded where appro- priate. Four additional new generic names were proposed for four previously described spe- cies of Scathophagidae. As distinguishing characters for each genus were given in the key to genera, and as only one species was referred to each, the requirements of Article 13(a)(i) and 68(d) of the Code were met and the generic names were validated at the time of their publication. They are listed below to draw attention to them and, in one case, to give distribution records not cited in the 1965 catalog (A catalog of the Diptera of America north of Mexico, USDA Handbook 276). Acronyms for depositions cited in this paper are BMNH (British National History Museum, London); CAS (California Acad- emy of Science, San Francisco); CNC (Ca- nadian National Collection, Ottawa); LACM (Los Angeles County Museum, Los Angeles, California; and USNM (United States Na- tional Museum of Natural History, Wash- ington, D.C.). Tethinidae Masoniella richardsi Vockeroth, New GENus, NEw SPECIES Masoniella richardsi Vockeroth, 1987: 1075 [nomina nuda, differentiating characters for the genus and species are indicated in key to genera (p. 1075) and are illustrated (Figs! 101-2,°82.95 a3; and™.14)]. Specimens examined.— Holotype 6, He- lendale [San Bernardino Co.], Cal., 18.V.1955 (W.R. Richards) (CNC). Para- types: (36: 6)..22. 2: sBMNEH,, ‘CAS SIENG LACM, USNM): same data as holotype (5 $); same locality and date as holotype but collected by W. R. M. Mason (5 @). Cali- fornia. Inyo: China Ranch, 30.V.1955 (J. Belkin et al.) (9 6, 3 2). Los Angeles: Duarte, 20.X1.1950 (N. Ehmann) (21 ¢, 13 9). San Bernardino: Victorville, 16.V.1995 (W. R. M. Mason) (1 ¢, 1 2). Distribution.-USA. California (Inyo, Los Angeles, and San Bernardino counties). Etymology.—The genus (feminine) and species are named in honor of my former colleagues W. R. M. Mason and W. R. Rich- ards both of whom, although students of Hymenoptera and Homoptera respectively, consistently collected large numbers of Dip- tera. Scathophagidae Dromogaster incompta Vockeroth, New GENus, NEw SPECIES Dromogaster incompta Vockeroth, 1987: 1096 [nomina nuda; differentiating char- acters for the genus and species are in- dicated in key to genera, p. 1096]. Specimens examined.— Holotype 4, Bil- by, Alta., June 1, 1924, D. Bryant (CAS). Paratypes (CAS, CNC, USNM): same data as the holotype (2 6, 2 2). CANADA. AI- berta: 14 mi W. Banff(4500’), 11.VHI.1955, J. R. McGillis (5 2). Saskatchewan: Attons Lake, Cut Knife, 3.VI.1940, A. R. Brooks (1 2). VOLUME 97, NUMBER 3 Distribution.— Canada. Alberta, Sas- katchewan. Etymology.—The name Dromogaster (feminine) is derived from the Greek words dromos (a type of ship) and gaster (belly) because of the large, boat-shaped sternite 7 of the female. The name incompta is de- rived from the Latin word incomptus (un- adorned) because of the lack of distinctive external characters of the species. Huckettia nearctica Vockeroth, New Genus, NEw SPECIES Huckettia nearctica Vockeroth, 1987: 1095 [nomina nuda; differentiating characters for the genus and species are indicated in key to genera, p. 1095]. Specimens examined.—Holotype 4, Beaverhill Lake, N.W.T., 66°44’N, 104°20’W, 29.VI.1966 (G. E. Shewell) (CNC). Paratypes (1694, 1589; collection dates 19.VI to 3. VIII; BMNH, CAS, CNC, Lund, USNM, St. Petersburg): CANADA. Yukon Territory: Herschel Island (J. S. Wa- terhouse). Northwest Territories: Kidluit Bay, Richards Island (J. R. Vockeroth); Fra- ser Lake, 68°45’N, 120°36'W (G. E. Shew- ell); Salmita Mines, 64°0S’N, 111°15'W (J. G. Chillcott); near Beechey Lake, 69°14'N, 106°50’W (Shewell); Ford Lake, 63°11'N, 107°19'W (Shewell); Beaverhill Lake (Shewell); Baker Lake (Chillcott); Spence Bay (Chillcott, A. E. R. Downe); Chesterfield [Inlet] (Chillcott, Vockeroth); Geillini Lake, 60°18'N., 95°35’W (C. D. Bird, Chillcott); 65°15'N, 89°30'W (Chillcott); Cambridge Bay, Victoria Island (E. H. N. Smith, G. K. Sweatman); Clyde, Baffin Island (J. E. H. Martin, Shewell); Hazen Camp, Ellesmere Island, 81°49'N, 71°18’W (R. E. Leech). Manitoba: Farnworth Lake near Churchill (Bird). Quebec: Payne Bay (E. E. Mac- Dougall, W. R. Mason); Sugluk (H. Huck- ett). USA. Alaska: Naknek (J. B. Hartley). Distribution.—Canada. Manitoba, North- west Territories, Quebec, Yukon. USA. Alaska. Etymology.—The genus (feminine) is 133 named in honor of H. C. Huckett, who pub- lished extensively on Nearctic Muscidae and Anthomyiidae from 1921 to 1977 and was one of the first to collect Diptera in northern Europe in order to compare them with Ne- arctic specimens. The species name indi- cates the wide distribution of the species in the northern part of the Nearctic Region. Brooksiella Vockeroth Brooksiella Vockeroth, 1987: 1094. Type species: Microprosopa varicornis Curran, 1927, monotypy. Name validated when published (Art. 1 3(a)(i), (b); Art 68(d). Dif- ferentiating characters for the genus are given in the key to genera (p. 1094). Specimens examined.—2 4, 15 2 (includ- ing holotype @). Etymology.—The genus (feminine) is named in honor of A. R. Brooks, who had an unrivalled knowledge of the insects of the prairies and parklands of Central Can- ada and whose preliminary manuscript re- vision of Nearctic Scathophagidae was of great value to me when I began studying this family. Neorthacheta Vockeroth Neorthacheta Vockeroth, 1987: 1096. Type species: Orthocheta dissimilis Malloch, 1924, monotypy. Name validated when published (c.f. Brooksiella). Differentiat- ing characters for the genus are given in the key to genera (p. 1096). Specimens examined.—33 6, 46 @ (in- cluding holotype @). Biology.—The larva feeds on young leaf shoots of Jris (H. J. Teskey, personal com- munication). Etymology.—The generic name (fem1- nine) is a combination of the Greek word neos (new) and the generic name Orthacheta Rondani. Peratomyia Vockeroth Peratomyia Vockeroth, 1987: 1096. Type species: Hexamitocera vittata Coquillett, 734 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1898, monotypy. Name validated when published (c.f. Brooksiella). Differentiat- ing characters for the genus are given in the key to genera (p. 1096). Specimens examined.—18 6, 15 2 (in- cluding the holotype 3). Etymology.—The generic name (femi- nine) is formed from a combination of the Greek words peratos (west) and myia (fly) because of the western distribution of the single species. Synchysa Vockeroth Synchysa Vockeroth, 1987: 1096. Type spe- cies: Coenosia tricincta Loew, 1869, monotypy. Name validated when pub- lished (c.f. Brooksiella). Differentiating characters for the genus are given in the key to genera (p. 1096). Specimens examined.—25 64, 30 @ (in- cluding a syntype 3). Distribution.— The known range has been extended from that given in the Nearctic catalog to southeast British Columbia (Fer- nie), in the West, and to Cape Breton Island, Nova Scotia, in the East. Etymology.—The generic name (femi- nine) is derived from the Greek word syn- chisis (confusion) because of the previous confusion concerning both family and ge- neric relationships. Muscidae Hydrotaea ponti Vockeroth Hydrotaea ponti Vockeroth, 1987: 1123. The specific name, ponti, is a nomen nu- dum. It was intended as a replacement name for curvipes Stein, 1920 (Trichopticus), a species referred by Huckett (1965, Memoir of the Entomological Society of Canada 42: 317) to Phaonia but which, because of the presence of a strong proclinate upper orbital bristle in the female, is referable to the genus Hydrotaea. This makes the name Hydro- taea curvipes (Stein, 1920) a secondary homonym of Hydrotaea curvipes (Fallén, 1825, Musca), which is in turn a primary homonym of Musca curvipes Gmelin, 1790. Hydrotaea curvipes (Fallén) is considered a senior unavailable synonym of Hydrotaea meridionalis Portschinsky, 1882, but is still a senior homonym of Hydrotaea curvipes (Stein). The latter is here renamed Hydro- taea ponti Vockeroth, nomen novum, in rec- ognition of the very large contribution to taxonomy of Muscidae made by A. C. Pont. The paper referred to as Vockeroth, 1984, in the footnote on page 1118 of the Manual of Nearctic Diptera (volume 2), has not been published. It was to have discussed some of the generic synonyms listed in the footnote. Acknowledgments: I thank Drs. W. N. Mathis (USNM), F. C. Thompson and C. W. Sabrosky (Systematic Entomology Lab- oratory, ARS, USDA, Washington, D.C.) for reviewing the manuscript. Dr. B. Brown (LACM) kindly loaned numerous speci- mens of Masoniella richardsi, which are designated herein as paratypes. Dr. P. H. Arnaud, Jr. (CAS) loaned the holotype and paratypes of Dromogaster incompta from Bilby, Alberta. J. R. Vockeroth, Research Associate, Bi- ological Resources Division, Centre for Land and Biological Resources Research, Agri- culture Canada, Ottawa, Ontario, Canada KIA 0C6. PROC. ENTOMOL. SOC. WASH. 976); 19955 p35 Book REVIEW The Gall Midges of the Neotropical Re- gion. R. J. Gagné. 1994. Cornell Univer- sity Press. Ithaca, New York, USA. 352 pp. $52.50 (cloth). A wide range of topics is covered on the taxonomic status and biology of more than 450 Neotropical gall midges (Diptera: Ce- cidomyiidae). The first chapter contains short bibliographies of eleven entomolo- gists who contributed to early taxonomic knowledge of gall midges and Gagné’s gentle assessment of the historic value of their work. I found this chapter a good read. It renewed my appreciation of the difficulties endured by early researchers working, sometimes, under less than ideal condi- tions. Chapters two and three discuss dis- tribution, external anatomy and biology of gall midges at each life stage. A full-page, halftone drawing of dorsal and ventral views of a third instar larva greatly aids under- standing of larval taxonomy. The entire family is evenly discussed and well sum- marized. In addition to the gall feeders, less commonly known primitive fungal feeders, inquilines, and predators are also discussed. Larvae, pupae, adults, and galls all con- tribute to the identification of gall midges and need to be properly preserved. Chapter four presents the preferred techniques for collecting, rearing and preparing specimens. Such essential information is too often omitted from taxonomic presentations. Chapter five comprises about onehalf of the book. It contains a taxonomic overview of the Cecidomyiidae and lists all genera and species known from the Neotropical Re- gion. Useful information such as distribu- tion, diagnostic characters, and references are presented here. Relationships among three subfamilies of Cecidomyiidae and the supertribes of Cecidomylinae are distin- guished by a cladogram of 18 anatomical characters. Illustrated keys, including one to genus, are easy to use although, depend- ing on the group, the three life stages larvae, pupae, and male or female adults are needed to reach some identifications. Line drawings are well done. An exception that I found was in figure 9 where the first tarsomere was so small that it was difficult to identify. Chapter six discusses the importance of plant hosts and gall types in identifying gall midges. It presents a key to cecidomyiids based on all known published records of their damage to plants even if the insects have not been described. If aware of only the plant name and gall type or shape, re- searchers and general biologists alike can easily use this key to identify an unknown gall midge. To facilitate use by a wide range of read- ers, the author has avoided technical terms whenever possible. The glossary is brief but adequate. The extensive bibliography con- tains over 400 citations and is thorough and current. This book is a companion to The Plant- Feeding Gall Midges of North America, 1989, Cornell University Press, also by Gagné. Together, these volumes provide a comprehensive and exceptionally well-writ- ten summary of present knowledge of the new world cecidomylids. Both are hard- bound and printed on acid-free paper. The two-column format affords wide margins and is easy to read. Over 300 clear, unclut- tered drawings illustrate each book and make it easier to use the keys. Gagné has suc- ceeded in presenting useful keys and de- scriptions of all the known gall midges of the New World and in reviewing gall midge biology, taxonomy, and current literature. He encourages the interested biologist and student, as well as the professional, by pro- viding a clear assessment of the current sta- tus of knowledge and indicating relevant ar- eas for further study of these insects. Paul E. Boldt, USDA, ARS, 808 Black- land Rd., Temple, Texas 76502 U.S.A. oh -¢ ay : | bigs ee pane me f meV i i inden nit Ll Walia ic} it pie jie tea Tea Win ih dA opie SAAR AE PEA a ONT RORAN MeV ce ) anes oi ¥34a 0 Muskowy 1) xeghin Linceoy paises AS nib le fa Dsdod etierete if 4; ovo Rithia Ad fa lab Way @ci | ead remnhihes, ae) et thas oye (l five ai ve Die See Tig Ci wert frat tte ry WORE Silvia Cutting holt AAAS ve SARS % h vt) tots AW Re ale Wai i, ial Gy’ Hoek ens! i j ; +9) ae Lith Rein ma nit Mi? et aga oe ; ug Wy : AH in if ; tenes » yh ! L » (mee Wy Capita hl yy | her Sigil h “et fi Hens! } : ‘ti } +t, py" met s bees ait | Tg, EL NN SRE BRR Ooi } Pima Via LA On’ » © ORC at? Sever ee ere | ’ js iil Macid 24 chasacile wt aidiaac) covatraie’ TOaeehrot of irae fred Nii : re, . : < j fart ma Vials ih ie +o inne TS cas a ve fete!) Ch ROE ea 1 | ; hin a rv ** tel yh Wal: ‘ . ; i vi | ‘ pine | og AY wt fol vai : al oni Cheg ith Ue j » res ep Tey oer fad Hind il ) ( 4 js ; ; ; om rt 7 vi) Hine Sivslie Maa: a ip 7 5 ify: “dig the aie woh | ALRASIAOG Ry be when iy i at (tA ed a Melia Ai . Pures eee RE aiitorall 7 19) Sigh, eh gaan Lif Agel CTPISPRL, teat AL i) GOR La Tee val veal he (lh Li noha ae Mio wont epee bedi ler Lethal ney 4 1) reopen newmans 2a8 ee) i Hvinaeny Se ; La Hiietys site Wp Peis 2 nels 4 el Beagles allt) gee ed 47 at TI Ae OF PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS Cynipid Galls of the Eastern United States, by Lewis H. Weld. 124 pp. 1959 Cynipid) Galls:of the Southwest, by Lewis He Weld\;35) pp; 1960 i.e ee i SLOLE DI 01243 [e gow lo N\erOa rey 3) (0 rays | KALER Gee Mia ih er Ae sO) ies Boy Magar PUL) TO ARUNR Aah A ERRURUD SM Se Bove Vleet pe hs Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. SEN WSO? GTi 121) WecBUNeh PAI CE Oo Ce ve RDN Se! BE a OHOR TERN Viars As tered | ru ODOR RUM Oe MIC coal A Short History of the Entomological Society of Washington, by Ashley B. Gurney. 15 pp. 1976 Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. RS DSS Keg tha rads Ta ce aN yr Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. SHES AER (Oo hig n) a) WIRE IT Ao) ROE BNE we nena PL SR CRM erly ecu Rhee An ai ea LL eA adie ha aa a Ore Boel A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael i Schauff. 85 pp. 1990__ No. No. MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving. O75 1530) Epa US 4PM Iu Le aor ate = eed ot Co AB NO SONIA AL aM ora Na Wen lee Bo aan TEC, OR DORE. AS BEN The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. FAD 0}) 3, 6502 PANS MiMaR Ae gMhr rasa ei BO Ut a Pl AeA Rl lee a ef ELL a eto ie Rear ct ths We rt A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi MY) cAleceA INCAS YS OTOP SA cee ek cP RA I I ia ak NN a deg Bre Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- pogonidaé), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979. 2 The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. PAULO) oy ayypn ICP sd (Ua Ne CSN A a Mrs eal ae Vale aL IM sale Subic, te Ne ees PO MU ie Ee ANT NR Ea . Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W. Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982 . A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 (2) Op SE ros See AU DAL MS OE ce DY Mesa sae Ee an NR Oa RAY a AG aU LL Ny aS Pol 4S pe AP Aa le ew . The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. OTA M0} otal HEL YAMS» LN STEN ZU eS OM ha AML A Dead I he EO APD HUY DANI Re ace PY aL Rk rc . An Identification Manual for the North American Genera of the Family Braconidae (Hyme- noptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987) $ 5.00 3.00 6.00 1.00 1.00 1.50 2.00 10.00 $15.00 (out of print) 11.00 12.00 10.00 11.00 18.00 5.00 18.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $25.00 per volume to non-members and $13.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra except on prepaid orders, Dealers are allowed a discount of 10 per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Custodian, Entomological Society of Washington, c/o Department of Entomology, MRC 168, Smithsonian Institution, Washington, D.C. 20560. CONTENTS (Continued from front cover) MALDONADO CAPRILES, J.—Notes about Vescinae, a key to the world genera, and descrip- tion of two new Chopardita (Heteroptera: Reduviidae) MORAN, MATTHEW D.—Intraguild predation between sympatric species of mantids (Man- todea: Mantidae) NEUNZIG, H. H—New genera and species of Peruvian Phycitinae (Lepidoptera: Pyralidae) ... POLHEMUS, JOHN T.—Nomenclatural and synonymical notes on the genera Diplonychus La- porte and Appasus Amyot and Serville (Heteroptera: Belostomatidae) POLHEMUS, JOHN T. and ROBERT W. SITES—The identity of Pelocoris biimpressus Mon- tandon and synonymy of Pelocoris species in the southwestern United States (Heteroptera: Naucoridae) PRICE, ROGER D. and ROBERT M. TIMM—The chewing louse genus Aoftiella (Phthiraptera: Gyropidae) from South American night monkeys, Aotus (Primates: Cebidae) RAO, NAVIN and WAYNE N. MATHIS—A revision of the shore-fly genus Rhysophora Cres- son (Diptera: Ephydridae) SCARBROUGH, AUBREY G., BETH B. NORDEN, and KARL V. KROMBEIN—A new species of Townsendia Williston (Diptera: Asilidae) from Florida with notes on its associ- ation with Perdita graenicheri Timberlake (Hymenoptera: Andrenidae) WEBB, RALPH E., GEOFFREY B. WHITE, and KEVIN W. THORPE—Response of gypsy moth (Lepidoptera: Lymantriidae) larvae to sticky barrier bands on simulated trees WHEELER, A. G., JR., and E. RICHARD HOEBEKE—Coccinella novemnotata in northeastern North America: Historical occurrence and current status (Coleoptera: Coccinellidae) WOOD, DIANE L. and J. E. McPHERSON—Life history and laboratory rearing of Hydrometra hungerfordi Torre-Bueno (Heteroptera: Hydrometridae) with descriptions of immature stages NOTES KIDD, KATHLEEN A., CHRISTINE A. NALEPA, ERIC R. DAY, and MICHAEL G. WALD- VOGEL— Distribution of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in North Carolina and Virginia VOCKEROTH, J. R.—Validation of nomina nuda of Nearctic Tethinidae, Scathophagidae, and Muscidae proposed in Manual of Nearctic Diptera BOOK REVIEW BOLDT, PAUL E.—The gall midges of the Neotropical Region VOL."97 OCTOBER 1995 NO. 4 PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED QUARTERLY OW) ees one x 4 \ ‘) GOOG NUV 27 1995 CONTENTS BARRACLOUGH, D. A. and TERRY A. WHEELER—Three new species of Cami jaae Dip- tera) from the southwestern nearctic region, the first species of the family described from PRE ING NVODLEUT LE Wik Sod Mimicaie ity Vora Rw UCR is! ae ON GACH Ma Mabey eA aos a Ser ah Hae Ane 737 CARROLL, J. F.—Laboratory evaluation of predatory capabilities of a common wolf spider (Araneae: Lycosidae) against two species of ticks (Acari: Ixodidae) .......................04. 746 EPSTEIN, MARC E.—False-parasitized cocoons and the biology of Aididae (Lepidoptera: Zy- BACMOICME ANE a ncre Ma ue etal ares iL NEL ks MAN ANY ad Us STR UM as ea ae Ya Weare Nits SAM ae ch ae 750 EVANS, HOWARD E.—A reconsideration of the cylindricus group of the genus Anoplius Du- FALE EALVIMESHOMLEr Ay LE OMIPUIdae)s Mrcaecioe see erat acieins Aa stake ake enielas MUSE Ay SN Este Salen wera & Sig si View GAGNE, RAYMOND J. and PAUL E. BOLDT—The gall midges (Diptera: Cecidomyiidae) of iBaceraris: spp-1(Asteraceae) in the United ‘Statesin i/o se Weravn sae a ye pi ae a Nae aE & 767 GOEDEN, RICHARD D., DAVID H. HEADRICK, and JEFFREY A. TEERINK—Life history and description of immature stages of Urophora timberlakei Blanc and Foote (Diptera: Tephritidae) on native Asteraceae in southern California ................. 00060. cece cece eeneeees 779 MALDONADO CAPRILES, J.—New Nearctic species of Oncocephalus Klug (Heteroptera: Re- AUIVALGACaAS LENO POG AINAG)s Sen Smee) 6a AE I ue EMM AAS CD, CaS Vaceeets CRAIN iCiare UR BRT lS ry Vay 791 McKEEVER, STURGIS, DANIEL V. HAGAN, and WILLIAM L. GROGAN, JR.—Compara- tive study of mouthparts of predaceous midges of the tribe Palpomyiini (Diptera: Cerato- poOsonidac) monte astenm United States: WN stance an Me cme Ce UNIS aN. ue dileiaahe 799 MILLER, KELLY B. and MICHAEL A. IVIE—Enallagma optimolocus, a new species of dam- Selily trom Montana (Odonata; Coenacnonidae) rir ak ys .:s seen sence waotacs bes Sesie clela pie nonitae 833 PRICE, ROGER D. and DALE H. CLAYTON—-A new genus and three new species of chewing lice (Phthiraptera: Philopteridae) from Peruvian ovenbirds (Passeriformes: Furnariidae) ... 839 RIDER, D. A. and L. H. ROLSTON—Nomenclatural changes in the Pentatomidae (Hemiptera- LETS CG PARE ea) bare Set neta Vo ALOR be Ob mM AM RC Le CAN ERI TY an ae dl rae ualcae ee Mahe athe Te 845 STAINES, C: Li—Francisco de: Asis Monr6és: A perspective’. )......00. 0 cccl cele dene ce ee eee eee eeee 856 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1995 JOHN W. NEAL, Jr., President NORMAN E. WOODLEY, Treasurer RALPH P. ECKERLIN, President-Elect NATHAN SCHIFF, Program Chairman DaRLENE D. JupD, Recording Secretary M. ALMA SOLIs, Membership Chairman Ho.uis B. WILLIAMS, Corresponding Secretary PAUL J. SPANGLER, Past President JAMES PAKALUK, Custodian THOMAS J. HENRY, Editor DAvipD R. SMITH, Associate Editor Gary L. MILLER, Book Review Editor Publications Committee DONALD R. DAVIS... TERRY L. ERWIN F. CHRISTIAN THOMPSON A. G. WHEELER, JR. Honorary President CurtTIs W. SABROSKY Honorary Members LoutsE M. RUSSELL ALAN STONE KARL V. KROMBEIN All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, MRC-168, Smithsonian Insti- tution, Washington, D.C. 20560. 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 8 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, MRC-168, Smith- sonian Institution, Washington, D.C. 20560. Members in good standing receive the Proceedings of the Ento- mological 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 P. 218 OF THE JANUARY, 1991 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. Editor: Thomas J. Henry, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- West, Beltsville, MD 20705. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 31 October 1995 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 737-745 THREE NEW SPECIES OF CAMILLIDAE (DIPTERA) FROM THE SOUTHWESTERN NEARCTIC REGION, THE FIRST SPECIES OF THE FAMILY DESCRIBED FROM THE NEW WORLD D. A. BARRACLOUGH AND TERRY A. WHEELER (DAB) Natal Museum, Private Bag 9070, Pietermaritzburg 3200, South Africa; (TAW) Dept of Natural Resource Sciences, McGill University, Macdonald Campus, 21,111 Lake- shore, Ste-Anne-de-Bellevue, Québec, H9X 3V9, Canada. Abstract.—The first endemic species of Camillidae are described from the New World and the Nearctic Region. Three new species are described: Afrocamilla arizona (south- eastern Arizona); A. bispinosa (southern California); and Camilla arnaudi (Baja Califor- nia, Mexico). All species are illustrated and a key to the endemic Nearctic species of Camillidae is presented. Camilla glabra (Fallén, 1823), previously introduced to Canada from Europe, is likely to be no longer extant in the Nearctic Region. The biology of Camillidae is briefly reviewed. Key Words: The Camillidae are a small family of dro- sophilid-like acalyptrate Diptera in the su- perfamily Ephydroidea (see McAlpine 1989). Prior to this study, the family was thought to be an exclusively Old World group, being widespread in the temperate and tropical parts of the Palearctic and Af- rotropical Regions, but not yet recorded from Madagascar. The extant Palearctic fauna comprises 11 species of Camilla Hal- iday, 1838, centred in the Mediterranean subregion and widely distributed through Europe into the northwest of the Russian Federation and Mongolia. Recent work on the Afrotropical fauna (Barraclough 1992, 1993) has shown that Camillidae are likely to be widespread in Africa. Three species of Camilla (one undescribed) are known only from South Africa. The other three Af- rotropical genera, Afrocamilla Barraclough, 1992, Katacamilla Papp, 1978, and Tera- tocamilla Barraclough, 1993, were previ- ously thought to be endemic to Africa, al- beit widespread (except Teratocamilla). Diptera, Camillidae, Afrocamilla, Camilla, Nearctic Region The only published record of Camillidae in the New World was a population of the Palearctic species Camilla glabra (Fallén, 1823), reported from Ottawa, Canada (McAlpine 1960), and apparently an acci- dental introduction from Europe (McAlpine 1987: 1023). There are 25 specimens of C. glabra in the Canadian National Collection of Insects (Ottawa), all taken at or near the Central Experimental Farm in Ottawa be- tween 1954 and 1965. The specimens may have been breeding in droppings in a turkey pound located on the Farm; when the tur- keys were removed from the farm, the C. glabra population also disappeared. It was surprising, therefore, when we dis- covered specimens of two congeneric spe- cies of Nearctic Camillidae in the Canadian National Collection of Insects. During 1993 several specialists and Diptera collections in North America were consulted about ad- ditional camillid specimens, but only one further specimen (belonging to a second ge- nus) was located. Despite the paucity of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Al ‘ esl Fig. 1. Type localities of Nearctic Camillidae. Af- rocamilla arizona (A), A. bispinosa (B), Camilla ar- naudi (C). material (each species is known from only One specimen), all three species are de- scribed, given the zoogeographic implica- tions of their discovery. They represent the first endemic species of Camillidae from the New World. Two species, from Arizona and California, are described in Afrocamilla. The third species, from Mexico, is de- scribed in Camilla. All three species should exit at Camillidae in the key to families (adults) of Nearctic Diptera by McAlpine (1981), although the preapical dorsal bristle on the mid tibia is absent in one species of Afrocamilla. MATERIALS AND METHODS Specimens examined are deposited in the following institutions (acronyms in paren- theses): California Academy of Sciences, San Francisco, USA (CASC); Canadian National Collection of Insects, Ottawa, Canada (CNCI). Holotype label data are cit- ed exactly as they appear, with supplemen- tary information in brackets; a slash denotes the end of a line of print and a semicolon separates data on different labels. Abdomens were detached at or near their base and macerated in warm to hot 80-85% lactic acid or hot 10% KOH. After exami- nation, dissected terminalia and associated abdominal parts were stored in glycerin in a microvial pinned beneath each source specimen. Morphological terminology mainly fol- lows Barraclough (1992, 1993); abbrevia- tions are not used, except for tergum (T). Bilaterally symmetrical structures are de- scribed in the singular. Head/thorax length was measured from the anterior margin of the third antennal segment to the abdominal base. Wing length was measured from the humeral crossvein to the wing-tip. KEY TO NEARCTIC SPECIES OF CAMILLIDAE Camilla glabra 1s likely to be no longer established in North America and is omitted from this key. 1. Palpus dark brown. Scutellum with apical bris- tles longer and stronger than basal pair. Ab- domen with paired median marginal T2 bristles shorter than length of T1+T2 (Fig. 5). T2 un- modified, not exserted above level of T3, and base of T3 not invaginated beneath posterior margin of T2 (Fig. 5) ....Camilla arnaudi n. — Palpus yellow or yellow-brown. Scutellum with apical bristles noticeably shorter than bas- al pair. Abdomen with paired median marginal T2 bristles much longer than length of Tl + T2 (Fig. 4). T2 modified, exserted above level of T3, and base of T3 invaginated beneath pos- terior margin of T2 (Fig. 4) . Head and thorax predominantly dark brown, mesonotum sparsely pollinose. One vibrissa. Apical scutellar bristles about one-third length of basal pair. Mid tibia without dorsal preapical bristle Afrocamilla arizona n. sp. — Head and thorax predominantly yellow-brown, mesonotum shining. Two vibrissae. Apical scu- tellar bristles about two-thirds length of basal pair. Mid tibia with dorsal preapical bristle Afrocamilla bispinosa n. sp. sp. N VOLUME 97, NUMBER 4 739 Figs. 2-5. A. arizona, abdominal terga 1—3 (lateral). 5, C. arnaudi, abdominal terga 1—3 (lateral). Afrocamilla arizona Barraclough and Wheeler, new species Figs. 3-4, 6-7 Description (based on ¢ holotype, 2 un- known).—Size: Head/thorax length 1.2 mm; wing length 2.2 mm. Color: Head mainly dark brown to black; cheek, pro- boscis, palpus, antenna, upper facial region and lower parafacial yellow or yellow- brown; sparse silver pollinosity on occiput and face. Thorax dark brown, pleuron slightly paler, humeral callus and propleu- ron mostly yellowish; yellow to pale brown pollinosity present on mesonotum, sparse on posterior section of pleuron, dense on scutellum. Both fore legs missing; mid leg yellow, apical half of femur and basal third of tibia brown; hind leg yellow, femur part- ly brown-tinged on middle and tibia brown on basal two-fifths. Wing membrane mostly slightly smoky, but with pale brown trans- verse basal fascia, which is narrow anteri- External characters of Nearctic Camillidae. 2, A. bispinosa, head (lateral). 3, A. arizona, wing. 4, orly and expanded to about twice this width at posterior margin. Veins pale yellow. Ab- domen dark brown to black with metallic reflections, pale at extreme base; Tl and T2 with sparse brown pollinosity. Head: Eye margin smoothly rounded anterodorsally in profile. Width of face and parafacials, at mid-height, subequal to length of antenna. Antenna with aristal vestiture barely dis- cernible, but apparently shorter than in A. bispinosa (see below). Upper orbital plate inconspicuous anteriorly, width here about 0.6 that of ocellar triangle. Postocellar bristles relatively well developed, length about 1.5 that of ocellar triangle. Ocellar triangle not unusually elevated above upper eye margin in profile, distance less than length of second antennal segment. Procli- nate fronto-orbital bristle present, anterior reclinate fronto-orbital about 0.25 length of proclinate bristle. One vibrissa present. Cheek height in profile about 0.25 length PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON postg TD, Gi 7 ane LE, a epiph C7 aE SE Sz er eS — __ a : Ges = Loc postg ee N 7 WS Figs. 6—7. distiphallus omitted). Scale bar = 0.1 mm. Abbreviations: epand—epandrium; hypd—hypandrium; cerc—cercus; sur—surstylus; phapod—phallapodeme; epiph—epiphallus; postg—postgonite. of third antennal segment. Thorax: Apical scutellar marginal bristles weakly devel- oped, about one-third length and strength of basal marginals. Fore femora missing (fem- oral armature unknown). Mid tibia without dorsal preapical bristle. Wing not unusually slender; without any slightly longer ventral spinules differentiated on costa between R, and R,,,. Abdomen: T2 modified, exserted above level of T3, and base of T3 invagi- nated beneath posterior margin of T2; T2 with paired median marginal bristles much longer than length of Tl + T2 (Fig. 4). Spiracles 1—5 in membrane near lateral margins of terga. Male postabdomen: T6 reduced to slender lateral vestiges, narrowly separated dorsally, spiracle 6 in tergum near ventral margin. Epandrium narrow, rectan- gular in profile, anterior margin concave (Fig. 6). Cerci well developed, closely ap- proximated, about 1.5 higher than broad, Afrocamilla arizona. 6, male terminalia (lateral). 7, hypandrium and phallic complex (ventral, with short setae. Hypandrium narrow lat- erally, broader anteromedially, convex an- teriorly, pregonites connected to anterior re- gion of hypandrium via a broad ventral strip (Figs. 6-7). Epiphallus and postgonites rel- atively closely approximated in profile, epi- phallus elongate, very narrow along much of length and tapered apically in profile (Fig. 6); postgonite unusually elongate and of uniform width along much of length, smoothly rounded apically in profile (Fig. 6). Surstylus approximately as long as height of epandrium, broad basally, nar- rowed and curved anteriorly at apex (Fig. 6). Type material.—Holotype 3d (CNCI), “Ramsey Canyon [31°27’N, 110°17'W] / Hu- achuca Mts. [Mountains] / 13.m[iles].s[outh]. Sierra Vista / Cochise Co[unty]. ARIZ[ona, U.S.A.] / Mar. 7-11.1967 / R. EF Sternitzki [sic = Sternitzky]” [all hand-written]; ““HO- VOLUME 97, NUMBER 4 LOTYPE ¢ / AFROCAMILLA / ARIZONA / Barraclough & Wheeler” [Red ink on white card, with red perimeter]. In fair condition: left antenna missing, some cephalic bristles missing, fore legs, distal part of right mid leg, and left hind leg missing. Left wing and ab- domen dissected and stored in microvial pinned below specimen. Etymology.—The species name is a noun in apposition referring to the state of Ari- zona, the source of the holotype. Remarks.—Afrocamilla arizona is distin- guished from both named Afrotropical spe- cies, as well as Afrocamilla bispinosa and Camilla arnaudi in having only one vibris- sa and the apical scutellar marginal bristles much shorter and weaker than the basal pair (typically longer and stronger in the Afro- tropical species). The presence of only one vibrissa in Afrocamilla is unusual. The male terminalia, most notably the length and shape of the postgonite, are also distinctive. All Afrotropical species dissected by the senior author have the hypandrium quite different in shape in ventral view (cf. McAlpine 1987: 1024: Fig. 2) to that of A. arizona. In A. arizona the hypandrium is much narrower and also more smoothly rounded anteriorly. Afrocamilla bispinosa Barraclough and Wheeler, new species Figs. 2, 8-9 Description (based on 2 holotype, d un- known).—Size: Head/thorax length 1.3 mm; wing length 1.8 mm. Color: Head mainly yellow-brown, face and much of oc- ciput paler; proboscis, third antennal seg- ment and arista brown to dark brown; sparse pale yellow pollinosity on face and occiput. Thorax yellow to yellow-brown, legs paler, particularly coxae and trochan- ters which are pale yellow; sparse pollinos- ity present on anterior margin of mesono- tum and posterior part of pleuron, denser, silver pollinosity on scutellum. Wing mem- brane mostly hyaline, but with pale yellow transverse basal fascia, which is uniformly narrow across base of wing. Veins pale yel- 741 low. Abdomen yellow to pale brown; T1 and T2 with sparse brown pollinosity. Head (Fig. 2): Eye margin relatively abruptly an- gled anterodorsally in profile. Width of face and parafacials together, at mid-height, about 1.2 length of antenna. Antenna with short ventral rays along apical three-quar- ters of arista, longest dorsal rays 1.1 length of third antennal segment. Upper or- bital plate inconspicuously developed an- teriorly, width about half that of ocellar tri- angle. Postocellar bristles relatively well developed, length about 1.3 that of ocellar triangle. Ocellar triangle unusually elevated above upper eye margin in profile, distance more than length of second antennal seg- ment. Proclinate fronto-orbital bristle pres- ent, anterior reclinate fronto-orbital about 0.25 length of proclinate bristle. Two vi- brissae present. Cheek height in profile about 0.4 length of third antennal seg- ment. Thorax: Apical scutéllar marginal bristles relatively well developed, about two-thirds length and strength of basal mar- ginals. Fore femur with 2 well-developed anteroventral spines on apical third, length about 0.6 maximum femoral diameter; posterodorsal and posteroventral bristles ir- regularly developed. Mid tibia with dorsal preapical bristle. Wing probably rather slen- der (folded in holotype); with 7—8 slightly longer ventral spinules differentiated on costa between R, and R,,,. Abdomen: T2 modified, exserted above level of T3, and base of T3 invaginated beneath posterior margin of T2; T2 with paired median mar- ginal bristles much longer than length of T1 + T2 (as in Fig. 4). TS about half length of T4; segments 5 and 6 with spiracles small, just discernible. Female postabdo- men: Two sclerotized spermathecae present, small, rugose, broader at base, sclerotized portion of spermathecal duct short, mem- branous portion of spermathecal duct ex- panded (Fig. 8). Ventral receptacle C- shaped, narrowed at mid-length and at each extremity, with slender, tubular, non-scler- otized appendage at one end (Fig. 9). Type material.—Holotype 2 (CNCI), PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 9 Figs. 8-9. Afrocamilla bispinosa. 8, spermathecae. 9, ventral receptacle (lateral). Scale bar = 0.05 mm. “TU.S.A.: California] San Dimas Can[yon]. [34°08’N, 117°46’W] / Los Angeles Co[unty]. / H-2-1958 / Reared III-10-58”’; “Collector / R.E. Ryckman’’; “HOLO- TYPE 2 / AFROCAMILLA / BISPINOSA / Barraclough & Wheeler” [Red ink on white card, with red perimeter]. In fairly good condition, right wing mostly missing and two fronto-orbital bristles detached. Abdomen dissected and stored in microvial pinned below specimen. Etymology.—tThe species name refers to the two anteroventral spines on the fore fe- mur (L. bi = two, spinosa = spine). Remarks.—Although this species pos- sesses the three major character states de- fining Afrocamilla (basal wing fascia, elon- gate T2 median marginal bristles, T2 apex and base of T3 modified), it differs from all known African species in two significant characters. The first of these, the presence of two anteroventral spines on the fore fe- mur, is possibly unique in Afrocamilla (the fore legs of the unique holotype of A. ari- zona are missing) and also distinguishes A. bispinosa from Camilla arnaudi. Intrage- neric variation in femoral armature does oc- cur in other camillid genera (e.g. Katacam- illa). The second character is the well-scler- otized spermathecae. In all African material examined by the senior author the sperma- thecae are less strongly sclerotized than the ventral receptacle. In addition to the above characters Af- rocamilla bispinosa is distinguished from all other species of the genus by the dorsal preapical bristle on the mid tibia. A. bispi- nosa is an unusually pale species, although the senior author has seen a comparable South African species, apparently not close- ly related. The color of A. bispinosa appears to be fully developed. The holotype was reared by R. E. Ryck- man from material collected in southern California (Fig. 1). There is no further in- formation associated with the specimen but it may have been reared from the nest of a wood rat (Neotoma sp., Cricetidae) (V. F Lee, personal communication). Camilla arnaudi Barraclough and Wheeler, new species Figs. 5, 10-11 Description (based on d holotype, 2 un- known).—Size: Head/thorax length 1.4 mm; wing length 2.5 mm. Color: Head dark brown, cheek and anterior margin of frons slightly paler and antenna (except arista) partly yellowish; pollinosity silver to pale brown, most evident on occipital re- gion. Thorax dark brown, dense, golden pollinosity present on scutellum. Legs yel- low to yellow-brown, tarsi slightly darker. Wing hyaline, veins pale yellow. Abdomen dark brown to black with metallic appear- ance; brown pollinosity present on T1 and T2. Head: First antennal segment con- cealed, not visible in profile or frontal view. Third antennal segment smoothly and even- ly rounded apically, dorso-apical section VOLUME 97, NUMBER 4 743 Figs. 10-11. without noticeably longer pubescence. Aris- ta with inconspicuous ventral rays along apical two-thirds to three-quarters, length of these at most subequal to width of arista at base, longest dorsal rays subequal in length to width of third antennal segment in pro- file. Width of face and parafacials together, at mid-height, just less than length of an- tenna. Two vibrissae (both missing in type), peristomal bristles mostly missing. Vibris- sal angle not prominent, about coincident with anterior extent of profrons in profile. Facial region somewhat excavate, epistomal margin upturned towards antennal bases. Cheek height in profile about 0.9 length of third antennal segment. Occiput with up- per half extensively excavate. Thorax: An- terior sternopleural bristle not evident (ster- nopleurals detached). Scutellar marginal bristles of similar strength, apical bristles slightly longer than basal bristles. Fore fe- Camilla arnaudi. 10, male terminalia (lateral). 11, hypandrium and phallic complex (ventral). Scale bar = 0.1 mm. Abbreviations: distph—distiphallus; epiph—epiphallus; postg—postgonite. mur with small anteroventral spine (femur crumpled, length of spine relative to fem- oral diameter not measurable). Mid tibia with dorsal preapical bristle. Wing moder- ately broad, not pointed apically. Costal vestiture between R, and R,,, relatively short and sparse, longer spinules not differ- entiated. Abdomen: T2 with paired median marginal bristles, these distinctly longer and stronger than other tergal bristles, length 0.6—0.7X length of Tl + T2 (Fig. 5). Spiracles 1—5 in membrane near lIateral margins of terga. Male postabdomen: T6 reduced to 2 slender sclerites, narrowly sep- arated dorsally, spiracle 6 in tergum near ventral margin. Epandrium narrow, width at junction with surstylus about two-thirds that at dorsal margin (Fig. 10). Cerci well de- veloped, closely approximated, about 1.5 higher than broad, with short setae. Hypan- drium narrow laterally, broader anteriorly, 1/44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘x anteriorly, pregonites connected to or region of hypandrium via a broad ventral strip (Figs. 10-11). Epiphallus in- conspicuous, downcurved from postgonite in profile. Distiphallus long, membranous (Fig. 10). Postgonite not splayed outwards, well developed and elongate (almost twice surstylus length), straight and finger-like, slightly tapered apically, length about 5.0 width (Fig. 10). Surstylus approximately half as long as height of epandrium, broad basally, narrowed and curved anteriorly at apex (Fig. 10). Type material——Holotype d (CASC): “San Bartolome / Baja Calif[ornia]. MEX[ico]. / HI-12-1953”; ‘Sefton Orca Exped[ition]. / to Gulf of Califfornia]. / P. H. Arnaud, collf[ector].’’; ““HOLOTYPE 3 / CAMILLA ARNAUDI / Barraclough & Wheeler”’ [Red ink on white card, with red perimeter]. In fair condition, left arista mostly missing, vibrissae and many frontal and mesonotal bristles detached. Abdomen dissected and stored in microvial pinned be- low specimen. Etymology.—This species is named for the collector of the holotype, Dr. P. H. Ar- naud, Jr. Remarks.—At first examination, this spe- cies appeared to be an unusually robust Af- rocamilla. However, dissection of the ab- domen showed T2 and T3 to be unmodi- fied, and the wing does not have a trans- verse basal fascia. The T2 median marginal bristles are unusually well developed for Camilla, but are substantially shorter and weaker than in Afrocamilla, in which they are usually longer than the length of Tl + T2. It is likely that there is correlation be- tween the development of these bristles and the modification of T2 and T3, as the bris- tles and associated musculature in the mod- ified T2 and T3 are probably used to move the folded wings (Barraclough 1992). We are not aware of any other named Camilla species with similarly developed bristles. However, the senior author has seen | ¢ and | 2 of a cavernicolous Camilla species from the northern Cape Province of South Africa. That species is in poor condition (in alcohol) and cannot be described, but is dis- tinguished from C. arnaudi by having a yel- low to yellow-brown head and thorax and no dorsal preapical bristle on the mid tibia. In Papp’s (1985) key to the world species of Camillidae, C. arnaudi runs closest to the Palearctic species C. glabra (Fallén, 1823), C. flavicauda Duda, 1922, C. sa- broskyi Papp, 1982 and C. mathisi Papp, 1985. The long T2 median marginal bristles are distinctive for C. arnaudi, as are the male terminalia, particularly the shape and form of the surstylus, postgonite and epi- phallus (see also figures of male terminalia in Papp (1982)). Both South African spe- cies of Camilla (see Barraclough 1993) dif- fer in lacking a dorsal preapical bristle on the mid tibia and an anteroventral spine on the fore femur. Camilla arnaudi differs from the two Nearctic species of Afrocam- illa in having the palpus dark brown, the scutellum with the apical bristles longer and stronger than the basal pair and the abdo- men with the paired median marginal T2 bristles shorter than the length of Tl + T2. The large distiphallus in this species may be apparent only in specimens cleared in lactic acid. The distiphallus is often over- cleared or lost in specimens cleared in KOH. DISCUSSION Information about the biology of Camil- lidae has been limited for many years, but has recently been supplemented by further data by Barraclough (1992, 1993). In south- ern Africa, Camillidae have been reared from the droppings of the rock hyrax, Pro- cavia capensis (Pallas) (Procaviidae), and have been shown to have a close associa- tion with the latrines of this small mammal in the subregion, and a Procavia species in Kenya (Barraclough, unpublished data). Southern African Camillidae have also been collected near droppings of Chiroptera or in areas inhabited by bats (Barraclough, un- published data). It appears that African Camillidae are coprophagous and live in as- VOLUME 97, NUMBER 4 sociation with the nests and droppings of a variety of mammals. In the Palearctic Re- gion, Camillidae have been collected near the entrances to rabbit burrows, and in as- sociation with rodent nests (Barraclough 1992). Although there are no definite biological data associated with the Nearctic species, the holotype of Afrocamilla bispinosa was reared, possibly from the nest or droppings of a wood rat (Neotoma sp.). The fact that Camillidae have not previously been col- lected from the United States may be due to a close association with the droppings, nests and burrows of small mammals. These habitats are often inaccessible or overlooked by many insect collectors. More specialized collecting techniques used in these habitats may show that Camillidae are more abundant and diverse in the south- western Nearctic Region than previously supposed. The fact that Camillidae have not been collected in greater numbers in the Nearctic Region may also be due to the phenology of the species. The types of all three species were collected in late winter and early spring (February/March), a time when there is generally much less collect- ing effort in North America. ACKNOWLEDGMENTS We are indebted to Dr P. H. Arnaud (CASC) and Dr J. M. Cumming (CNCI) for the loan of specimens in their care. Special 745 thanks go to V. F Lee (CASC), who in col- laboration with E. S. Ross and K. Cebra, provided valuable data on the collectors and type localities of Afrocamilla arizona and A. bispinosa. J. R. Vockeroth (CNCI) pro- vided information on the rise and fall of Camilla glabra in Canada. LITERATURE CITED Barraclough, D. A. 1992. Afrocamilla stuckenbergi, a new African genus and species of Camillidae (Diptera), with comments on its behaviour and bi- ology. Annals of the Natal Museum 33(1): 37-49. 1993. Teratocamilla, a new genus of Cam- illidae (Diptera) and two new species of Camilla, from southern Africa. Annals of the Natal Muse- um 34(1): 19-30. McAlpine, J. E 1960. First record of the family Cam- illidae in the New World (Diptera). Canadian En- tomologist 92: 954—956. . 1981. Key to families—Adults, pp. 89-124. In McAlpine, J. F et al., eds, Manual of Nearctic Diptera, Volume 1. Research Branch Agriculture Canada Monograph 27: 1—674. 1987. Camillidae, pp. 1023-1025. In Mc- Alpine, J. EK, ed., Manual of Nearctic Diptera, Vol- ume 2. Research Branch Agriculture Canada Monograph 28: 675-1332. 1989. Phylogeny and classification of the Muscomorpha, pp. 1397-1518. In McAlpine, J. E, and Wood, D. M., eds., Manual of Nearctic Dip- tera, Volume 3. Research Branch Agriculture Can- ada Monograph 32: 1333-1581. Papp, L. 1982. A revision of the species of Camilla Haliday described by J. E. Collin (Diptera: Cam- illidae). Memoirs of the Entomological Society of Washington 10: 125-135. . 1985. A key of the world species of Camil- lidae (Diptera). Acta Zoologica Hungarica 31: 217-227. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 746-749 ABORATORY EVALUATION OF PREDATORY CAPABILITIES OF A COMMON WOLF SPIDER (ARANEAE: LYCOSIDAE) AGAINST TWO SPECIES OF TICKS (ACARI: IXODIDAE) JME CARROEL Parasite Biology and Epidemiology Laboratory, Livestock and Poultry Sciences Insti- tute, ARS, USDA, Beltsville Agricultural Research Center, Beltsville, Maryland 20705. Abstract.—Black-legged ticks, Ixodes scapularis Say and adult American dog ticks, Dermacentor variabilis (Say) were readily attacked by the wolf spider, Schizocosa ocreata (Hentz) in petri dish bioassays. Younger (4-5 mm body length) S. ocreata were less successful in their attacks (66.7%) against J. scapularis adults than individuals with body lengths of 7-9 mm, which killed 100% of the /. scapularis adults. Against adults of D. variabilis the attacks of even the larger S. ocreata were generally ineffectual, killing only 14.3% of the larger tick. The S. ocreata were able to lift the D. variabilis from the substrate, but may have been deterred from further attack by defensive secretions known to be produced by the latter. Key Words: Lycosidae, Ixodidae Wolf spiders (Lycosidae) constitute a ma- jor group of arthropod predators on the for- est floor in northern temperate regions (Clark and Grant 1968). Like most spiders they are generalists in their choice of prey and because of their foraging activities on the leaf litter are likely to encounter host- seeking and recently fed ticks. Black-leg- ged ticks, Ixodes scapularis Say, and Amer- ican dog ticks, Dermacentor variabilis (Say) are three-host ticks; the former typi- cally a woodland species (Ginsberg and Ewing 1989) and the latter somewhat more of a woods-edge inhabitant at least in the adult stage (Sonenshine and Levy 1972). Larval and nymphal /. scapularis and D. variabilis seek hosts on the leaf litter (Gins- berg and Ewing 1989, Smith et al. 1946) and, although the adults of both species may quest for hosts as high as | m above ground level (Harlan and Foster 1990), they are exposed to wolf spider predation when Predation, Schizocosa ocreata, Ixodes scapularis, Dermacentor variabilis, moving to, or changing, questing sites. Af- ter each engorgement (three for females) both species of tick are vulnerable to attack when they leave their hosts to find molting sites Or Oviposition sites. There are few accounts of arthropod pre- dation upon ixodid ticks (e.g. Barre et al. 1991) and even fewer mention spiders. The common theridiid spider Achaearanea tep- idariorum (C. L. Koch), which often infests buildings, has been observed preying upon the lone star tick, Amblyomma americanum (Linnaeus) (Guarisco 1991). In Kenya, Mwangi et al. (1991) reported that (uniden- tified) spiders killed engorged Rhipicephal- us appendiculatus Neumann in the labora- tory. Wilkinson (1970) found that the wolf spider Lycosa godeffroyi Koch preyed upon engorged females of the cattle tick, Booph- ilus microplus (Canestrini), in Australia. According to Riechert and Lockley (1984) generalist spider predators can make VOLUME 97, NUMBER 4 a significant contribution toward biological control of insect pest species. To determine whether wolf spider predation might have any impact on the populations of two med- ically important tick species, a series of lab- oratory feeding bioassays was conducted. METHODS Wolf spiders were collected by pitfall traps and in vials by hand in mixed oak- beech woodlands and woods edges in Prince George’s County, Maryland. Spider collection sites were habitats where J. sca- pularis and D. variabilis occurred. In the laboratory spiders were maintained in petri dishes (5 cm diameter) containing moist tis- sue paper and were fed larvae of the Eu- ropean corn borer, Ostrinia nubilalis (Hub- ner) (Lepidoptera, Pyralidae). Ticks were collected with a 1-m? flannel drag, and kept in high humidity (99% R.H. for I. scapu- laris and 94% for D. variabilis) at 22—26°C and natural photoperiod. Schizocosa ocreata, the species of wolf spider collected in the greatest numbers, was used in the predation tests. Because an O. nubilalis larva constituted a large meal, spiders were tested 4 days after being fed. A single tick was placed in the petri dish with a spider and the initial encounter and activities for the next 5 min were observed. They were checked again after a second 5 min and again at 48 h after the tick was placed in the petri dish to determine tick mortality. As a reference for background tick mortality, ticks were placed in petri dishes containing moist tissue paper and checked at 48 h and | wk. Unfed male and female /. scapularis adults, unfed /. scapularis nymphs and en- gorged /. scapularis larvae were tested. The engorged nymphs dropped from their lab- oratory rat hosts <48 h before testing and were still actively crawling when confined with the spiders. Spiders of two size ranges (4-5 mm and 7—9 mm body length) were tested. Only unfed adult female D. varia- bilis (about twice the size of /. scapularis females) were tested. Spiders and_ ticks 747 Table 1. Proportion of encounters between wolf spiders, S. ocreata and ticks, /. scapularis and D. var- iabilis, confined together in petri dishes in which the tick was killed.* Body I. scapularis Length D. variabi- of S. Adults lis ocreata Unfed Fed Adult (mm) Male Female Nymph Larvae Female 7-9 1/1 18/18 1/3 2/14 4—5° 5/7 5/8 2/2 26/27 “Tick dead within 48 h after introduction in petri dish with spider. Each spider and tick only used in one test. » Spiders collected in spring contemporaneously with adult /. scapularis and D. variabilis and early I. sca- pularis nymphs. © Spiders collected in late summer and fall contem- poraneously with adult and larval /. scapularis. were tested just once. Success of S. ocreata predation against /. scapularis versus D. variabilis and between size classes of S. ocreata against adult J. scapularis were an- alyzed by Student’s ¢-test. RESULTS Almost all S. ocreata (i.e. 46 of 48) at- tacked the adult ticks (. scapularis and D. variabilis) confined with them upon their first encounter. The outcomes of these at- tacks varied. /. scapularis adults were all killed when confined with the larger (7-9 mm body length) S. ocreata, whereas 10 of 15 (66.7%) unfed /. scapularis adults were killed by the smaller (younger) S. ocreata (4-5 mm body length) (Table 1). S. ocreata belonging to the smaller size class were sig- nificantly less successful in killing female I. scapularis than were (t = 2.762, P < 0.02) the larger spiders. With one exception attacks upon the engorged /. scapularis l\ar- vae all ended with the tick eaten (Table 1). Unfed /. scapularis nymphs were also at- tacked, but because of their small size they soon found their way into cracks and crev- ices (e.g. between the petri dish and lid, or in the wad of moist tissue) inaccessible to the spiders. The larger S. ocreata had dif- ficulty in killing the unfed D. variabilis PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON » of 14, 14.3%) females. Although all ger S. ocreata but one were observed -king significantly fewer (t = 4.969, P 0.01) D. variabilis females than I. sca- pularis females were killed by S. ocreata. Attacks were occasionally triggered by the tick wandering into the spider, but in most cases the spider apparently detected the tick at distances of 5 to 10 mm, reo- riented itself, approached the tick and pounced upon it. Invariably ticks of both species responded to the attacks by drawing in their legs, appressing them close to their bodies and ceasing movement. Successful attacks usually involved a spider’s lifting the tick from the substrate. This gave the spider access to the tick’s venter, where there were more opportunities for the spi- der’s fangs to penetrate the tick’s interseg- mental membranes at the articulations of its legs. The S. ocreata were able to lift the D. variabilis females from the substrate, but the D. variabilis were usually released by the spiders within 10 sec, and the ticks re- mained motionless for several minutes. S. ocreata were observed holding /. scapular- is adults >30 min after attacking, which suggests that feeding was involved. Also exsuccous remains of fed /. scapularis nymphs were found in the petri dishes 24 h after the start of confinement of the nymphs with S. ocreata. DISCUSSION Multiple host ticks may be most vulner- able to predation by lycosids as well as many other predators just after having com- pleted engorgement and having dropped from their hosts. At this point they are still mobile and would attract the attention of a wolf spider. Furthermore, the tick’s recent blood meal would have stretched its opis- thosomal integument quite thin, presumably making it more easily bitten through by a spider than in its unfed configuration. This period of vulnerability is relatively brief, perhaps a matter of minutes or hours until the fed tick is ensconced in the leaf litter. Many fed larval /. scapularis are thought to drop off in the nests of their hosts, predom- inantly white-footed mice, Peromyscus leu- copus, and therefore safer from lycosids (Mather and Spielman 1986). The discrepancy between the two size classes of S. ocreata in their ability to kill I. scapularis adults may be due to several possible factors. One obvious explanation is size related. The larger spiders could be ex- pected to be stronger, have longer fangs and have the capacity to inject more venom into their prey. The problems the S. ocreata had with the D. variabilis females may have been due to their inability to penetrate the tick’s integument. A similar situation was reported for the jumping spider, Corythalia canosa Hentz (Salticidae), a specialist pred- ator of ants (Edwards et al. 1974). C. can- osa were unable to kill the heavily sclero- tized ants Trachymyrmex septentrionalis (McCook) and Cyphomyrmex minutus Rower, which like ticks pull in their legs and remain motionless when attacked. A different, or additional, explanation may account for the low success rate (two of 14, 14.3%) of S. ocreata against the American dog ticks they attacked. Yoder et al. (1993) found that D. variablis adults produced a waxy secretion on their dorsal surfaces when attacked by ants, or were otherwise similarly disturbed. The secre- tions caused the ants to cease their attacks, thereby protecting the ticks, a scenario rem- iniscent of the attacks by S. ocreata on D. variabilis. S. ocreata is common throughout wood- lands in the eastern United States (Dondale and Redner 1978, Stratton 1991), and was abundant in habitats where both species of tick occurred. In view of the relatively low abundance of ticks compared to other po- tential prey of S. ocreata, it is unlikely that a wolf spider such as S. ocreata will en- counter successive ticks before finding an- other meal. Therefore, tests were not con- ducted to determine the maximum number of ticks an S. ocreata is capable of eating. Further evaluation of the impact of S. ocreata and other common wolf spiders on VOLUME 97, NUMBER 4 I. scapularis populations is warranted. Also spiders, such as crab spiders (Thomisidae), that frequent adult /. scapularis questing sites on vegetation up to | m high should be considered as potentially important pred- ators of black-legged ticks. ACKNOWLEDGMENTS I thank Rod Crawford of the Burke Me- morial Museum, University of Washington, Seattle, Washington for identifying wolf spider specimens and supplying valuable information about them, and Kenneth Young of the USDA, ARS Livestock In- sects Laboratory, Beltsville, Maryland for the feeding and care of the wolf spiders. Also, I thank John Davis and J. A. Klun of the USDA, ARS Insect Chemical Ecology Laboratory, Beltsville, Maryland for pro- viding European corn borer larvae. I ex- press my appreciation to D. H. Wises, De- partment of Entomology, University of Kentucky, Lexington, KY, for his thought- full review of this paper. LITERATURE CITED Barre, N., H. Mauleon, G. I. Garris, and A. Kermarrec. 1991. Predators of the Amblyomma variegatum (Acari: Ixodidae) in Guadeloupe, French West In- dies. Experimental and Applied Acarology 12: 163-170. Clarke, R. D. and P. R. Grant. 1968. An experimental study of the role of spiders as predators of the forest litter community. Ecology 49: 1152-54. Dondale, C. D. and J. H. Redner. 1978. Revision of the nearctic wolf spider genus Schizocosa (Ara- neida: Lycosidae). Canadian Entomologist 110: 143-181. Edwards, G. B., J. E Carroll, and W. H. Whitcomb. 1974. Stoidis aurata (Araneae: Salticidae), a spi- 749 der predator of ants. Florida Entomologist. 57: 337-346. Ginsberg, H. S. and C. P. Ewing. 1989. Habitat dis- tribution of /xodes dammini (Acari: Ixodidae) and Lyme disease spirochetes on Fire Island, New York. Journal of Medical Entomology 26: 183— 189. Guarisco, H. 1991. spiders upon medically significant pests. Trans- Predation of two common house actions of the Kansas Academy of Sciences 94: 79-81. Harlan, H. J. and W. A. Foster. 1990. Micrometeoro- logic factors affecting field host-seeking activity of adult Dermacentor variabilis (Acari: Ixodidae). Journal of Medical Entomology 27: 471-479. Mather, T. N. and A. Spielman. 1986. Diurnal detach- ment of immature deer ticks (/xodes dammini) from nocturnal hosts. American Journal of Tropi- cal Medicine and Hygiene 35: 182-186. Mwangi, E. N., R. M. Newson, and G. P. Kanya. 1991. Predation of free-living engorged female Rhipi- cephalus appendiculatus. Experimental and Ap- plied Acarology 12: 153-162. Riechert, S. E. and T. Lockley. 1984. Spiders as bio- logical control agents. Annual Review of Ento- mology. 29: 299-320. Smith, C. N., M. M. Cole and H. K. Gouck. 1946. Biology and control of the American dog tick. United States Department of Agriculture Techni- cal Bulletin Number 95, 74 pp. Sonenshine, D. E. and G. EF Levy. 1972. Ecology of the American dog tick, Dermacentor variabilis, in a study area in Virginia. 2. Distribution in relation to vegetative types. Annals of the Entomological Society of America 65: 1175-1182. Stratton, G. E. 1991. A new species of wolf spider Schizocosa stridulans (Araneae, Lycosidae). Jour- nal of Arachnology 19: 29-39. Wilkinson, P. R. 1970. Factors affecting the distribu- tion of the cattle tick in Australia. Observations and hypotheses. Acarologia 12: 492—508. Yoder, J. A., R. J. Pollock, and A. Spielman. 1993. Ant-diversionary secretion of ticks: first demon- stration of an acarine allomone. Journal of Insect Physiology 39: 429-435. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 750-756 “ ALSE-PARASITIZED COCOONS AND THE BIOLOGY OF AIDIDAE (LEPIDOPTERA: ZYGAENOIDEA) MArc E. EPSTEIN Department of Entomology, NHB 105, Smithsonian Institution, Washington, D.C. 20560. Abstract.—Two or four holes in the outer mesh of cocoons in Aididae appear to mimic exit holes of parasitic Hymenoptera. Cocoon construction in Aidos amanda (Stoll) from Venezuela is described and documented with photographs, along with notes on its biology. Annona punicifolia is reported as a larval hostplant of A. amanda. Literature on the cocoons and biology of Aididae are reviewed. Citations of Guyaba as a larval hostplant of A. amanda are considered to be erroneous based on misidentified larvae. Scenarios for the evolution of cocoon construction in Aididae are discussed, as are possible functions of the false exit holes in their cocoons. Key Words: sitized cocoons The small Neotropical family Aididae (6 species), previously placed as a subfamily of Megalopygidae, is an important link in the understanding of the relationships among Limacodidae, Megalopygidae and Dalceridae (Epstein in press). Larval aidids differ from many larval megalopygids in lacking plumose setae on the dorsum, and in absences of verrucae below the spiracles and of large membranous pads on the ab- dominal prolegs (Epstein in press). Their two rows of urticating bristles on the dor- sum of the abdomen are sunken in repose and splayed out in rosettes when stimulated (Hopp 1935; Epstein in press). The earliest known description of aidid cocoons was given in a letter by Moritz (published in Klug 1836). They were de- scribed and figured by Dewitz (1878: 20- 26; pl. 1, figures 17-18), based on notes and specimens from Gollmer. Hopp’s (1930, 1935) descriptions were based on those of the earlier workers, though they were not cited in his later publication (Hopp Lepidoptera, Aididae, cocoon construction, predator avoidance, false-para- 1935). Cocoons of Aidos amanda (Stoll) were described as ‘*... spun on the under- side of leaves of different trees, look like the webs of ichneumon-flies [perhaps meaning a parasitized wasp cocoon], es- pecially since also their hiding places [emergence holes of braconids—from earlier literature; see discussion] are copied”’ (Hopp 1935:1072; english version). Forbes (1942:396) described the cocoon as “ with trap door, silken stem for suspension and one or two pairs of rounded depressions in the sides (varying in individuals of a sin- gle batch of A. amanda).” Descriptions of the cocoon and other im- mature stages of A. amanda by Hoffmann (1932) were based on misidentified Mega- lopyginae. This is deduced from features of the larvae described by Hoffmann, which occur in megalopygids and not in aidids. These include (1) matted wool covering the eggs (from the caudal end of the female ab- domen); (2) plumose setae, and a fleshy knob behind every spiracle in the larva; and (3) larval setae woven into the cocoon. VOLUME 97, NUMBER 4 Along with photographic documentation, I will describe the cocoon construction of a species believed to be A. amanda. | also report its life history, update the hostplant literature on aidids, discuss alternative functions of the false exit holed cocoons, and present evolutionary scenarios that may have brought them about. METHODS I reared a single aidid larva from Ven- ezuela (see locality data below). It was rec- ognized as an aidid from the description in Hopp (1935) and from specimens exam- ined at the Natural History Museum, Lon- don (BMNH). Although the larva perished in its cocoon as a prepupa, I tentatively identified the species as A. amanda based on adult males and females collected from the same locality in September 1990, a lo- cation where only one species of Aidos is known to occur. These specimens closely match the types of A. amanda figured in Stoll (1782, pl. 383, figs. C, D). The gen- italia of the Venezuelan and other aidid species examined appear to be rather uni- form. However, further evidence in support of the identity of this population as A. amanda is found in the labial palpus, which has only two segments; three are found in other aidid species and in muse- um specimens labelled as A. amanda from outside of Venezuela and the Guianas. The two-segmented palpus matches those of a specimen believed to be A. amanda from Guyana (Cornell University Coll.), the closest locality of any specimen examined to its Surinam type locality. Voucher spec- imens of adults, the larva and its skins are deposited in the entomological collection of the National Museum of National His- tory, Washington, D.C. (USNM). Although the cocoon was lost, another cocoon from the same locality in Venezuela is placed in the USNM collection. Additional aidid specimens examined and the institutions where they are deposited, include: adults (USNM, BMNH, and Universidad Central de Venezuela, Maracay); larvae (BMNH); 751 and cocoons (USNM, BMNH and Cornell University). LIFE HISTORY The presumed larva of A. amanda was found in the Venezuelan Ilanos at Hato Masaguaral, Edo. Guarico, 45 km S. Cal- abozo (8.57°N, 67.58°W), 75 m elevation on 9 July 1989 (Fig. 1). The larva was feeding on a leaf of Annona punicifolia Triana and Planch at ca. 1400 h in cloudy conditions during a light rain. The larva was attached to the underside of the leaf, with its prothorax visible from above at the consumed leaf apex. The Annona bush was on the edge of a savanna that is often in- undated during the rainy season (Fig. 2). In a plastic rearing container, the larva spun silk trails when moving between leaves and a silk pad on the leaf it was feeding on. When pulled from a leaf the larva would often remain motionless in the bottom of the container, with silk hooked to its crochets. Locomotion is described in Epstein (in press). The larva had seven or eight instars, based on the assumption that it was a sec- ond or third instar when first discovered (Table 1). Stadia lasted from at least six to thirteen days, though the larva was reared under a wide range of climatic conditions. It was fed the native hostplant A. punici- folia until the fifth or sixth instar when it was switched to oak (Quercus spp.). It pre- ferred oak when later given a choice of leaves of an annonaceaeous tree during its final stage. Cocoon construction.—On 9 September the prepupa began spinning the cocoon be- tween two leaves (Fig. 6) in figure-eight fashion. The final frass pellet was light brown and the prepupa became wetted down, similar to what occurs with Lima- codidae (McNaulty, 1967; Epstein in press). By 10 September the prepupa had spun a thin outer mesh with one small oval hole on each side (approx. | mm). Each hole had a more dense reinforcement of silk around its edges, roughly a third the distance from PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-5. 1, late instar larva of Aidos amanda (photo by K. V. Sandved); 2, habitat and Annona punicifolia, hostplant of A. amanda, at Hato Masaguaral, Venezuela; 3, recently completed cocoon of A. amanda (hatch end); 4, laterodorsal view of four-holed aidid cocoon with emerged pupal exuvia (photo by V. Krantz); 5, detail of aidid cocoon, hatch above (hole above 3 mm across) (photo by L. Minor-Penland). VOLUME 97, NUMBER 4 Table 1. Life history of Aidos amanda (Stoll). Dates of molting, approximate instar and host for one individual reared to prepupa. Instar/ Date Specimen Cocoon Host 09 vii 2-3 Annona 15 vii — 3-4 Annona 22 vii skin 4-5 Annona 04 viii skin 5-6 Annona/Quercus 12 vili skin 6-7 Quercus 25 vili prepupa 7-8 Quercus 9-12 ix cocoon one end of the cocoon (Fig. 7). From inside the cocoon the emerald green prepupa made the holes very apparent. On 11 September the larva constructed a u-shaped escape hatch on the end of the cocoon near the holes, reinforcing the edge of the hatch with silk in much the same manner as the holes. The “‘holes’’ were sealed on the inside of the cocoon by 700 hrs on 12 September. Other hostplant records for Aididae.— Gollmer reported A. amanda on sweet or- ange (from Dewitz 1878). Since Hoffmann (1932) misidentified a megalopygid as A. amanda, it follows that his hostplant record for Aidos on Guyaba (probably /nga sp.) is also incorrect. Hoffmann’s hostplant record was later cited in Hopp (1935:1072, 1098) and Lima (1945). An adult specimen of Ai- dos yamouna (Dognin) from Peru in the USNM collection was reared on Cinchona ledgeriana, Moens. 1876 (Rubiaciae). This is a quinine-yielding tree, with leaves ‘“‘per- fectly glabrous on both surfaces,” original- ly from Rio Mamore, Bolivia (Trimen 1881). DISCUSSION The close resemblance of their false exit holes to emergence holes of parasitic Hy- menoptera may protect aidid cocoons from vertebrate predators. If vertebrates indeed learn that parasitized cocoons have little or no food to offer, as suggested by Hinton (1955), then it is also reasonable to hypoth- esize that they would tend to avoid cocoons constructed with false holes. This view was 158 expressed by Dewitz (1878) as it related to the cocoons of A. amanda. Hinton’s (1955) review of protective de- vices in Endopterygote pupae did not in- clude A. amanda, from the literature men- tioned above, or other examples of “‘false exit holes’’ in Lepidoptera cocoons. His one example of a chrysalis considered it to mimic parasitized galls (Agriolaus maesa, Lycaenidae) (from Eltringham 1923:89-— 90). Hinton’s other examples of possible predator-avoidance mechanisms included moth cocoons that appear to have parasitoid wasp cocoons extruding from them (e.g. Deilemera antinorii Oberthur; see Eltring- ham 1923:83, figure 32). Similarly, some species of ichneumonids build a false co- coon outside of the skin of their larval host, complete with a false emergence hole, while their true cocoon is inside the dead host (e.g. Hyposoter parorgyiae and H. ru- biginosa,; Finlayson 1966 and R. Carlson pers. comm.). The characterization of aidid cocoons as having the appearance of “‘webs of ichneu- mon-files’’ in the earlier literature is con- fusing (e.g. Hopp 1935), since ichneumon- ids do not build webs other than cocoons. The “‘webs”’ may refer to large ichneumo- nid cocoons, or groups of smaller braconid cocoons with emergence holes in them, as suggested in Klug (1836). Dewitz (1878: 24) thought they resembled a parasitized cocoon of a tenthredinid. The evolutionary development of false exit holes in aidids may have been an off- shoot of hatch construction. This is plausi- ble because two holes in all examples are in close proximity to the hatch, and the pre- pupa appears to reinforce both holes and hatch in similar fashion. The cladogram of the limacodid group (Epstein in press) in- dicates that hatch construction in Aididae is plesiomorphic, and that false exit holes are uniquely derived (autapomorphic). Another autapomorphic feature in aidids, the flexible and amorphous apex of the spinneret (Figs. 10-12), may be useful in shaping and re- inforcing the silk around the holes. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cocoon construction sequence of Aidos amanda: 6, wetting down and beginning of spinning between two leaves; 7, at first visible sign of holes (note spines visible through hole); 8, outer mesh near completion, prior to filling in of holes. VOLUME 97, NUMBER 4 Fig. 9. Costa Rica. Nest of Polybia occidentalis bohemani in The false exit holes may be functionally related to hatch construction in the follow- ing way. They allow the prepupa to view the immediate environment to select opti- mal hatch placement for future adult emer- gence. This ““window”’ hypothesis is sup- ported by my observations that the prepupa constructs the holes prior to the hatch. Moreover, in cocoons examined with a sin- gle pair of holes, the hatch is constructed at the same end of the cocoon. This suggests that with proper stimuli, no further holes are made. Even if the holes were to function in selecting hatch placement, this would not preclude them from having a_predator- avoidance function as well. Forbes’s (1942) observations, quoted above, suggest a flexibility in the number of holes constructed within A. amanda. However, the hole arrangement of a four- holed cocoon (USNM, no data or adult voucher) suggests that the number of holes may be fixed in an individual. In this co- coon one of two hole sites proximate to the 10-12. from two specimens) (scale length in parentheses): 10— 11, Aidid sp. from Brazil (BMNH); 10, ventral view of spinneret, flexible apex is folded down (arrow), silk pore with silk is above (46 pm); 11, dorsolateral view (100 ym); 12, Aidid sp. from Brazil (BMNH), dorsal view of spinneret, apex is extended (20 p.m). Figs. Spinneret of larval Aididae (exuviae 756 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON vas apparently blocked by a stem at ‘ime of construction. The prepupa con- ucted a smaller hole between the two holes posterior to the hatch, perhaps to compensate for the missing hole in the usu- al location. False holes may also give aidid cocoons the appearance of being small nests of ves- pids, or of other nest-building wasps, that are built on the undersides of leaves (e.g., Polybia occidentalis bohemani;, Fig. 9). The ‘holes’? on the cocoons could appear as nest entrances, though smaller in size (1 to 3 mm compared to approx. 9 mm for nest entrance in Fig. 9). Experimental studies are needed to de- termine whether there is less predation on aidid cocoons compared to similar cocoons without false exit holes (e.g., in megalo- pygids). Other future research should be undertaken to determine whether the num- ber of holes are fixed or flexible in a given species or between species of Aididae. ACKNOWLEDGMENTS I gratefully acknowledge the assistance of the following curators or collections man- agers, and their institutions: David Carter (The Natural History Museum, London); E. Richard Hoebeke (Cornell University); and Jurg de Marmels (U. Central de Venezuela, Maracay). Of particular importance was as- sistance in German translation from Elsie Froeschner, Marion Kotrba and Fred Wal- doc. Israel Nino (University Central de Ven- ezuela) identified the Annona hostplant. Robin Foster identified and obtained foliage of a tree species of Annonaceae, and Arnold Menke (SEL) identified the Polybia in Fig. 9. Victor Kranz, Laurie Miner-Penland and Kjell V. Sandved (Smithsonian Institution) provided photographic assistance. I gra- ciously thank Tomas Blohm for permission to do research at Hato Masaguaral, and R. Rudran (National Zoological Park) for ad- ministrative assistance that made my visits to Venezuela possible. I would also like to acknowledge Robert Carlson (SEL) and Klaus Horstmann (U. Wirzburg) for stimu- lating discussion and correspondence. Fund- ing was provided by the Smithsonian Office of Fellowships and Grants and the Interna- tional Environmental Science Program (Na- tional Zoological Park, Smithsonian). The manuscript benefitted from critical com- ments by R. Carlson, Daniel H. Janzen (U. Pennsylvania), Scott E. Miller (Bishop Mu- seum) and an anonymous reviewer. LITERATURE CITED Dewitz, H. 1878. Entwickelung einiger Venezuelan- ischer Schmetterlinge nach Beobachtungen von Gollmer. Archiv fiir Naturgeschichte 44: 1-36 + plate 1. Eltringham, H. 1923. Butterfly Lore. The Clarendon Press, Oxford. 180 pp. Epstein, M. E. in press. Revision and Phylogeny of the Limacodid-group Families, with Evolution- ary Studies on “‘Slug Caterpillars’ (Lepidoptera: Zygaenoidea). Smithsonian Contributions to Zo- ology. Finlayson, T. 1966. The False Cocoon of Hyposoter parorgyiae (Vier.) (Hymenoptera: Ichneumoni- dae). The Canadian Entomologist 98: 139. Forbes, W. T. M. 1942. The Lepidoptera of Barro Col- orado Island, Panama. 2. Bulletin of the Museum of Comparative Zoology 90: 265-405. Hinton. H. E. 1955. Protective devices of Endopter- ygote pupae. Transactions of the Society for Brit- ish Entomology 12: 49-92. Hoffmann, F 1932. Beitrage zur Naturgeschichte Brasilianischer Schmetterlinge. Deutsche Ento- mologische Zeitschrift 97-148. Hopp, W. 1930. Lepidoptera: Megalopygidae in Er- gebnisse einer zoologischen Sammelreise nach Brasilien, insbesondere in das Amazonasgebeit, ausgefiihrt von Dr. H. Zerny. Annalen des Natur- historischen Museums in Wien 44: 269-277. 1935. Megalopygidae, pp. 1071-1101. Jn Seitz, A., ed., The Macrolepidoptera of the World, Vol. 6. Alfred Kernan. Stuttgart [1934-1936]. Klug, G. R. 1836. Auszug aus einem Briefe des Hrn. A. Moritz. Caracas vom 27. Febr. 1836. Archiv fiir Naturgeschichte 2: 302—306. Lima, A. M. da Costa. 1945. Insetos do Brasil, Vol. 5. 379 pages. Imprensa Nacional, Rio de Janeiro. McNaulty, B. J. 1967. Outline Life Histories of some West African Lepidoptera. Part Il Limacodidae. Proceedings of the South London Entomological and Natural History Society 1-12. Stoll, C. 1782. Pp. 165-252. In Cramer, Uitlandsche Kapellen (Papillons exotiques). Vol. 4. Trimen, H. 1881. Cinchona ledgeriana, a distinct spe- cies. Trimen’s J. Botany. pl. 222 & 223, 321-325. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 757-766 A RECONSIDERATION OF THE CYLINDRICUS GROUP OF THE GENUS ANOPLIUS DUFOUR (HYMENOPTERA: POMPILIDAE) HOWARD E. EVANS Department of Entomology, Colorado State University, Fort Collins, Colorado 80523. Abstract.—Three names are removed from the synonymy of Anoplius (Pompilinus) cylindricus (Cresson). These are Arachnophila brevihirta Banks, Pompilinus subtruncatus Dreisbach, and P. truncatus Dreisbach. Pompilinus hispidus Dreisbach is regarded as a synonym of P. subtruncatus Dreisbach, while P. clavipes Dreisbach continues to be con- sidered a synonym of A. (P.) cylindricus (Cresson). Anoplius (Pompilinus) cylindricus is restricted to the southwestern United States and is clearly distinct from other members of this complex. The remaining three species are closely related and show some evidence of intergradation; two of them, A. (P.) subtruncatus and A. (P.) truncatus, occupy more limited ranges within that of the transcontinentally distributed species A. (P.) brevihirta. Some of the variation in A. (P.) brevihirta is analyzed. This species is restricted to broad areas of open sand, and populations in different sandy areas have in some cases diverged slightly in structure and color; there are also broadly geographic trends in structure and color. Key Words: Retirement should provide time to re- think some of the problems in one’s past research in which the solutions proposed seem naggingly unsatisfactory. Many years ago (Evans 1951) I placed four names in the synonymy of Anoplius (Pompilinus) cy- lindricus (Cresson); a fifth name was later added to the synonymy (Evans 1966). Ev- idence to support this lay in the apparent continuous variation in features of the male genitalia. Further collecting in the western half of the United States in recent years has convinced me that such a simplistic solu- tion to this complex is not satisfactory. Here I propose that four species be recog- nized in this complex, with two other names being retained in synonymy. Day (1974) placed the subgeneric name Pompilinus Ashmead (generotype Pompilus cylindricus Cresson) in the synonymy of the subgenus Arachnophroctonus Howard Spider wasps, Pompilidae, Anoplius, synonymy, distribution, variation (generotype Psammochares marginalis Banks). I regard Pompilinus as a probably monophyletic New World offshoot of the widely distributed Arachnophroctonus. In 1951 I erroneously placed the European An- oplius viaticus (L.), under the name A. fus- cus (L.), in Pompilinus; more properly it belongs in Arachnophroctonus. Whether Pompilinus deserves subgeneric status can be argued, but it is a large group divisible into at least three species-groups, so it is convenient to retain it as a subgenus. The most useful distinguishing features of this group are the fact that the transverse median vein of the fore wing meets the me- dia beyond the origin of the basal vein, the third submarginal cell is petiolate, and the spines of the female tarsal comb rarely ex- ceed the width of the basitarsus. The spe- cies of this group are difficult to separate in the female sex, and the males can be sep- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with assurance only by study of the lia. In 1951 I recognized two species- sroups in Pompilinus, the subcylindricus and cylindricus groups. Here I restrict the cylindricus group to that species and three forms previously considered synonyms of it. The remaining members of the former cylindricus group of Evans (1951) may be considered to constitute the marginatus group. The following are the distinguishing fea- tures of members of the cylindricus group as here defined. Females: eyes widely spaced, middle interocular distance 0.60 to 0.65 times transfacial distance; vertex pass- ing straight across between tops of eyes; ocellar triangle broad and flat; propodeum convex, with a steep, oblique declivity, and bearing at least a few short hairs (may be quite hairy); femora usually hirsute; fore basitarsus with a comb of three or four spines, these spines one to two times as long as width of basitarsus; one or more basal tergites banded with orange. Males: subgenital plate slender, acute, hirsute (Figs. 5—8); genitalia with squamae at base of parameres prominent; digiti clothed with short setae, many of which are bent apical- ly, except latero-apically with longer, straight setae; digiti broadly truncate api- cally (one exception) (Figs. 1—4); basal ter- gites with or without orange markings. In 1951 I characterized A. cylindricus as a sand dune species, and various papers on behavior published under this name all ap- ply to sand dune populations. All of these indicate that females prey on immature Geolycosa spiders occurring in sandy situ- ations, using the spider’s burrow in which to construct their own nest cells (Krombein 1953, Kurczewski and Kurczewski 1968, 1973, Gwynne 1979, Kurczewski 1981). The name A. brevihirta (Banks) is here ap- plied to these populations. Other elements in the species-group are by no means restricted to sandy places, but are often taken on plains or prairies. Un- fortunately females cannot at present be as- sociated with certainty with any of these forms, so of course nothing can be said con- cerning their predatory or nesting behavior. Males taken away from sandy areas tend to be larger than A. brevihirta and are all black. These males can be sorted into groups equivalent to Dreisbach’s species Pompilinus subtruncatus and P. truncatus and to Cresson’s Pompilus cylindricus. On at least one occasion these three forms have been taken in one locality on the same date (Tornillo, El Paso Co., Texas). Two of them have been taken together on at least six oc- casions (localities in Texas, New Mexico, Colorado, Illinois, and Michigan). Howev- er, each of these forms (and A. brevihirta) is subject to variation in features of the male genitalia and subgenital plate, varia- tion that in some cases blurs any lines that might be drawn between species. An exception is A. cylindricus itself. Study of the genitalia of the lectotype plac- es this as a member of a population restrict- ed to parts of the Southwest. The genitalia are quite distinctive, and I have seen no specimens intermediate with other members of this complex. These other members, A. brevihirta, A. subtruncatus, and A. trunca- tus, show evidence of intergradation, the latter two forming populations (not restrict- ed to sand dunes) within the broad range of the dune-inhabiting A. brevihirta. Anoplius (Pompilinus) cylindricus (Cresson) Pompilus cylindricus Cresson, 1867, p. 92. (Lectotype: ¢, Texas, ANSP). Pompilinus clavipes Dreisbach, 1958, pp. 61-63. (Holotype: 6, Texas: Conlon [? Conlen, Dallam Co.], 8 Aug. 1952, R.R. Dreisbach, MCZ). The lectotype of Cresson’s Pompilus cy- lindricus has genitalia and a_ subgenital plate virtually identical to those figured for Pompilinus clavipes by Dreisbach, and as figured here (Figs. 1, 5). This lectotype was designated many years ago, prior to the studies of Dreisbach (1949) and Evans (1951). The other 12 specimens in Cres- VOLUME 97, NUMBER 4 T9 Ry Figs. 1-4. Male genitalia (right half, ventral aspect) of Anoplius (Pompilinus) species. 1, cylindricus (Cres- son). 2, subtruncatus (Dreisbach). 3, truncatus (Dreisbach). 4, brevihirta (Banks). 760 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5-8. Male subgenital plates (ventral aspect) of Anoplius (Pompilinus) species. 5, cylindricus (Cresson). 6, subtruncatus (Dreisbach). 7, truncatus (Dreisbach). 8, brevihirta (Banks). VOLUME 97, NUMBER 4 son’s type series are all assignable to other species. Other males before me are from Kansas: Reno Co. (MCZ, NMNH); Texas: El Paso, Jeff Davis, and Pecos counties (CU, CSU); New Mexico: Chaves Co. (CSU); and Colorado: Prowers Co. (CSU). All have essentially identical genitalia and subgenital plates. All are all black, with a fore wing length from 6.0 to 8.5 mm. The genitalia are distinctive in that the para- meres are broadly spatulate and heavily se- tose along the mesal margin; the digiti are rounded apically, rather than truncate as in other members of this complex. I have seen several females that very probably belong here. All have a small amount of short erect hair on the propo- deum, but the femora have only scattered, very short hairs. The comb spines on the fore basitarsus are short, no longer than width of the basitarsus; the basitarsus bears 3 such spines (in one specimen 4). All have the first two tergites orange; fore wing length varies from 7.7 to 10.8 mm. The strongest evidence that this association is correct is to be found in a series of 3 fe- males and 13 males, all taken by myself on 7 July 1953 at Medora, Reno Co., Kansas (MCZ, NMNH) (the males are all paratypes of Dreisbach’s Pompilinus clavipes). Other females are from Kansas: Riley Co. (CU); Oklahoma: Grant Co. (MCZ); and Texas: Bastrop, Bee, Brewster, Dimmit, Travis, Kenedy, and Kleburg counties (BFL, CSU, CUT IMCZ): Anoplius (Pompilinus) subtruncatus (Dreisbach) New Status Pompilinus subtruncatus Dreisbach, 1949, pp. 17-18, figs. 58, 59. (Holotype: 6, Nebraska, Lincoln, 14 June 1909, C.H. Gable, Univ. Nebraska). Pompilinus hispidus Dreisbach, 1949, pp. 23-24, figs. 47, 48. (Holotype: ¢, Mich- igan, Tuscola Co., 20 Aug. 1940, R.R. Dreisbach, MCZ). Anoplius hispidus, Evans, 1970, p. 482. (Teton Co., Wyoming). 761 The type specimens of Pompilinus sub- truncatus and P. hispidus do indeed appear quite different. The former has the hairs on the subgenital plate short and depressed and the inner apical margins of the digiti not angularly extended. The latter has the hairs on the subgenital plate erect and bristling and the inner apical angles of the digiti pro- jecting angularly. However, other speci- mens combine these two features different- ly. Dreisbach pointed out two small groups of setae near the midline of the subgenital plate of P. hispidus, but these are not pres- ent in specimens I have seen other than the type. Variation in the condition of the hairs on the subgenital plate is striking. In some specimens they are wholly decumbent, in others only a few are erect, and in still oth- ers most are erect. In a male from Hialeah, Florida (NMNH), these hairs are especially long and bristling. A paratype of P. hispi- dus from Baldwin, Kansas (NMNH), has a strongly hirsute subgenital plate but the dig- iti lack an angulate process on the inner, apical margin. There are other examples that, to me, suggest that this is a single spe- cies subject to discordant variation in seem- ingly important features of the male abdo- men. Some specimens have fewer setae along the mesal margin of the parameres than figured, suggesting possible intergra- dation with A. brevihirta. Smaller speci- mens fall within the size range of that spe- cies; overall size range (fore wing length) varies from 5.3 to 9.0 mm. I have been un- able to associate any females with this spe- cies with any certainty. Males are all black, but it is probable that females are marked with orange as in other members of this complex. This species is widely distributed throughout the United States from the Rockies eastward. I have seen specimens from the following localities: Florida: Hia- leah (NMNH); Maryland: Bowie (NMNH); Louisiana (without further data) (NMNH); Michigan: Tuscola Co. (MCZ); Iowa: Sioux City (NMNH); Kansas: Baldwin (NMNH), PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Co. (NMNH); Nebraska: Chadron 'v. Colorado, Boulder), Lincoln, Neligh niv. Nebraska); North Dakota: Beach (NMNH); Wyoming: Teton Co. (MCZ); Colorado: Bent, Elbert, Larimer, Otero, and Weld counties (CSU); Texas: Jeff Davis (CU), El Paso (MCZ), and Presidio (ANSP) counties. Anoplius (Pompilinus) truncatus (Dreisbach) New Status Pompilinus truncatus Dreisbach, 1949, pp. 15, 16, figs. 40, 41. (Holotype: 6, Mich- igan, Gratiot Co., 2 Aug. 1947, R.R. Dreisbach, MCZ). As the species name implies, the para- meres of the male genitalia are obliquely truncate apically (Fig. 3). An additional useful character is the absence of setae from the center of the disc of the digitus. The subgenital plate (Fig. 7) tends to be more slender and pointed than in the preceding species, but this is a subtle and not wholly dependable character; the sides of the plate are less sinuate than in Anoplius cylindricus (Fig. 5) and A. brevihirta (Fig. 8). The mes- al margin of the paramere is relatively sparsely setose, much as in A. brevihirta but in contrast to A. cylindricus and A. subtrun- catus. In a series of 6 males from Tornillo, Tex- as (CSU), taken on the same day, some agree well with A. truncatus and others with A. brevihirta, while one has the left paramere truncate (as in A. truncatus), the right paramere acute (as in A. brevihirta). In a series of 4 males from Great Sand Dunes National Monument, Colorado (CSU), taken on the same day, three have genitalia typical of A. brevihirta, while the fourth has truncate parameres (although the digiti are fully clothed with setae and the subgenital plate is sinuate laterally). These specimens, and others, suggest that there is some interbreeding between these two spe- cies. Males of A. truncatus are all black and average larger than those of A. brevihirta, fore wing length varying from 5.5 to 7.2 mm. I have been unable to associate any females with this species. Anoplius truncatus appears to have a more restricted range than A. subtruncatus, occurring from Texas and New Mexico to Illinois and Michigan. Dreisbach’s para- types are from Manistee and Midland coun- ties, Michigan, Carlinville, [linois, and Roswell, New Mexico. Other males I have seen are from Colorado: Bent, Larimer, Prowers, and Weld counties (CSU); New Mexico: Hidalgo Co. (CSU); and Texas: El Paso (CSU) and Jeff Davis (ANSP) coun- ties. Anoplius (Pompilinus) brevihirta (Banks) New Status Arachnophila brevihirta Banks, 1945, p. 105. (Holotype: 2, Chicago, Illinois, July; CG: Brues} MCZ): Anoplius (Pompilinus) cylindricus, Evans, 1951, pp. 294-297; fies) Siol232idn part). This is the most widely distributed mem- ber of this complex, ranging from coast to coast and from Canada deep into Mexico. It is the form most often identified as A. cylindricus, under which name there have been several reports on its biology (as dis- cussed above). It is characteristic of areas of open sand, and virtually every system of dunes I have visited has a population of these wasps. Males average smaller than those of the preceding three species, fore wing length varying from 4.0 to 7.0 mm. The genitalia have the parameres acute, their mesal margins sparsely setose; the dig- iti are fully clothed with setae and the dis- tal, mesal angle is prominent (Fig. 4). The subgenital plate is slender, with sinuate margins, and is clothed ventrally with sub- recumbent setae (Fig. 8). Even with the removal of the three pre- ceding species from synonymy, this re- mains a very variable species. About 75% of the males I have seen have one or more basal tergites marked with orange. All- black males cannot always be distinguished VOLUME 97, NUMBER 4 from those of other members of this com- plex without examining the genitalia. Even so, occasional specimens are puzzling. A male from Mt. Vernon, Missouri (CSU), has the parameres very slender, almost lin- ear. A few males have the parameres some- what truncate apically; two such specimens were cited under A. truncatus. Females vary in fore wing length from 4.5 to 9.5 mm. Those from any one locality often differ greatly in size, suggesting that they have developed on spiders of different sizes. Females also differ in the number of spines on the fore basitarsus and in the amount of erect hair on the femora and the propodeum. Females have at least the sec- ond tergite banded with orange, sometimes part or all of the first and third tergites and some of the basal sternites. Since sand dune systems tend to be widely spaced in nature, it is possible that local populations have di- verged slightly in structure and color. Some of the variation also has a broadly geographic component (Table 1). Males from the eastern coastal states invariably have orange coloration on the abdomen, and this is true of males from Kansas, Tex- as, and eastern Colorado. Females from these same areas usually have only 3 comb spines on the fore basitarsus, while those from Michigan and from south central Col- orado usually have 4 comb spines (in these two areas males are usually all black). Fe- males from many western sites tend to have much more hair on the propodeum and femora than those from eastern localities, which may be almost devoid of such pilos- ity. It is not clear why there are resemblances between the population at Great Sand Dunes in Colorado (represented by a series at CSU) and that at Warren Dunes, Michi- gan (represented by a series at MCZ) (Table 1). The latter site is subject to cooling winds from Lake Michigan, such that cer- tain microclimatic conditions may approx- imate those at a much higher elevation (2440 m) in Colorado; however, the Colo- rado site has a much lower annual precipi- 763 Table 1. Geographic variation in male coloration and the number of comb spines on the fore basitarsus of females of Anoplius (Pompilinus) brevihirta (Banks).! No. of Males No. of Females Marked with All 3 Comb 4 Comb Locality Orange Black Spines Spines MA and CT 7 0) Na 0) NJ and MD 3 0) 17 0) NC 7 0) 64 10) FE 20 0) 48 0) NY and PA? 0) S} 11 1 MP >) 15 3 16 KS 10 0) 19 Ti 13s 1 < ] 0) 6 0) W. TX 3} 0) l 11 EEO 8 10) 14 7 S3CO# 1 8 11 44 NM 1 10) 4 9 Totals 76 26 215 95 ' Includes only localities with a sample size of 5 or more. States are combined when records are consis- tent; states are divided when separate parts differ. > Three localities on Lakes Erie and Ontario: Erie Co., PA, Oswego and Jefferson Co., NY. 3 Warren Sand Dunes State Park, Berrien Co. 4 Great Sand Dunes National Monument, Alamosa Co. tation than the Michigan site. Specimens from Kansas, eastern Colorado, and a smat- tering of localities in the Great Plains, tend to approximate eastern specimens. A comparison of the females from two widely separated areas of extensive sand is especially interesting (Fig. 9). In coastal North Carolina there is evidently a spring generation in which females are larger and have only 1—1.5 tergites orange. This is fol- lowed by a summer generation of females that are smaller and have 2 or 3 basal ter- gites orange. (These data are based on a long series of specimens from Nags Head and Kill Devil Hills in NMNH.) Presum- ably the large females of the spring gener- ation result from the fact that females of the preceding summer took more mature Geo- lycosa spiders, while the small females of the summer generation developed on less mature spiders. At Great Sand Dunes Na- tional Monument the season is much short- 164 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4.0- 4.4- 48- 5.2- 5.6- 6.0- 6.4- 6.8- 7.2- 7.6- 8.0- 8.4- 8.8- 92- 9.6- 75 79 83 87 9.1 95 9.9 Ww WwW a a = WW LL Te Oo ) Zz 4.3 47 5.1 55 59 63 6.7 7.) LENGTH OF FORE WING (mm) Fig. 9. A size and color comparison of two generations of Anoplius (Pompilinus) brevihirta (Banks) from coastal North Carolina and the single generation from southern Colorado at 2440 m elevation. A, Kill Devil Hills, NC, 23 June to 14 Sept.: 2-3 tergites orange. B, Kill Devil Hills, NC, 24 May to 22 June: 1—1.5 tergites orange. C, Great Sand Dunes National Monument, CO, July—August: 1—1.5 tergites orange. er; here the females approximate in size and coloration members of the spring genera- tion in coastal North Carolina. However, these females have much more body hair and a stronger tarsal comb. It is possible that the more extensive orange coloration of the second generation in North Carolina is the result of summer heat on the pupal stage, as occurs in other Hymenoptera. Males from Texas, New Mexico, and east- ern Colorado are almost invariably marked with orange, and there is a corresponding tendency for females from these areas to be more extensively marked with orange. Clearly we are far from explaining the ob- served variation in this species, but it must be recognized when one attempts to cir- cumscribe this species as distinct from oth- ers: KEY TO MALES OF THE A. CYLINDRICUS GROUP . Digiti rounded apically; parameres broadly spatulate (Figs 1) 225 ee cylindricus (Cresson) Digiti broadly truncate apically; parameres less broadly expanded apically . Parameres obliquely truncate apically; center of disc of digiti devoid of setae (Fig. 3) dycgh hn sdea si ccucathecmte REL ok truncatus (Dreisbach) Parameres more or less acute apically; center of digiti covered with short setae . Mesal margin of parameres heavily setose (Fig. 2); subgenital plate somewhat broader, sides gradually tapered, often bearing erect setae (Fig. 6); all black Ld cht) Big Saad 2a ye ane elie Ba subtruncatus (Dreisbach) Mesal margin of parameres more sparsely se- tose (Fig. 4); subgenital plate more slender, with sinuate lateral margins (Fig. 8); basal ter- gites with or without orange coloration Mone thee eee ce sett Sie Meee eee brevihirta (Banks) VOLUME 97, NUMBER 4 CONCLUSIONS Members of the A. cylindricus species- group occur throughout much of North America and have commonly been consid- ered to constitute a single variable species. However, four populations can be distin- guished on the basis of the male genitalia. While the differences between them are slight, they seem sufficiently discrete to suggest that to a considerable degree they breed independently of one another. This is especially true of A. cylindricus itself, which is restricted to the southwestern states. Two others, A. subtruncatus and A. truncatus (both known from males only), occupy limited ranges within that of the very widely distributed A. brevihirta. The last is characteristic of broad areas of open sand, where females prey on Geolycosa spi- ders. While two or three of these species have sometimes been taken at the same time and place and remain recognizable, oc- casional specimens are somewhat interme- diate in characters of the genitalia. Both A. subtruncatus and A. truncatus occur pri- marily on plains and prairies and may in- tergrade with A. brevihirta on the periphery of the sand dune habitats of the latter. Fur- ther collecting and data on nesting behavior are needed to fully clarify the components of this complex. Molecular studies might be useful both for associating the sexes and for better understanding the degree to which in- terbreeding occurs. A. brevihirta, even as restricted here, is a very variable species. Populations in widely spaced sand dune systems have di- verged slightly in features of structure and color, and there are also broadly geographic tendencies relating to the length and num- ber of comb spines on the fore basitarsus of the females, the length and abundance of body pilosity, and the amount of orange coloration on the abdomen. Further collect- ing of larger series from more sand dune areas may serve to fill out details in the spatial distribution of these variable char- acters. ACKNOWLEDGMENTS For the loan of specimens, I am indebted to Arnold Menke, U.S. National Museum of Natural History (NMNH), Donald Azu- ma, Academy of Natural Sciences of Phil- adelphia (ANSP), Michael S. Kelley, Mu- seum of Comparative Zoology at Harvard (MCZ), E. Richard Hoebeke, Cornell Uni- versity (CU), and Allan Hook, Bracken- ridge Field Laboratory, University of Texas (BFL). Other specimens are located at C.P. Gillette Insect Biodiversity Museum at Col- orado State University (CSU). LITERATURE CITED Banks, N. 1945. Two new species of Psammochari- dae. Psyche 52: 105-106. Cresson, E. T. 1867. Notes on the Pompilidae of North America, with descriptions of new species. Transactions of the American Entomological So- ciety 1: 85-150. Day, M. C. 1974. A contribution to the taxonomy of the genus Anoplius Dufour (Hymenoptera: Pom- pilidae) including a revision of the Palaeotropical subgenus Orientanoplius Haupt. Bulletin of the British Museum (Natural History) 30: 373-404. Dreisbach, R. R. 1949. Psammocharini (Hymenop- tera) of North America and the Antilles: Key to genera; new species and key to males of Pompi- linus. Entomologica Americana 29: 1-58. 1958. Two new species in the genus Pom- pilinus (Hymenoptera: Psammocharidae) with photomicrographs of the genitalia and subgenital plate of the male. Entomological News 69: 61-65. Evans, H. E. 1951. A taxonomic study of the Nearctic spider wasps belonging to the tribe Pompilini (Hy- menoptera: Pompilidae). Part II: genus Anoplius Dufour. Transactions of the American Entomolog- ical Society 76: 207-361. . 1966. A revision of the Mexican and Central American spider wasps of the subfamily Pompi- linae (Hymenoptera: Pompilidae). Memoirs of the American Entomological Society 20: 1442. 1970. Ecological-behavioral studies of the wasps of Jackson Hole, Wyoming. Bulletin of the Museum of Comparative Zoology 140: 451—509. Gwynne, D. T. 1979. Nesting biology of the spider wasps (Hymenoptera: Pompilidae) which prey on burrowing wolf wolf spiders (Araneae: Lycosidae, Geolycosa). Journal of Natural History 13: 681— 692. Krombein, K. V. 1953. Kill Devil Hills Wasps, 1952. Proceedings of the Entomological Society of Washington 55: 113-135. Kurezweski, E E. 1981. Observations on the nesting 166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON behaviors of spider-wasps in southern Florida ‘Hymenoptera: Pompilidae). Florida Entomologist 64: 424-437. urczewski, E E. and E. J. Kurczewski. 1968. Host records for some North American Pompilidae (Hymenoptera) with a discussion of factors in prey selection. Journal of the Kansas Entomolog- ical Society 41: 1-33. . 1973. Host records for some North American Pompilidae (Hymenoptera). Third supplement. Tribe Pompilini. Journal of the Kansas Entomo- logical Society 46: 65-81. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 767-778 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF BACCHARIS SPP. (ASTERACEAE) IN THE UNITED STATES RAYMOND J. GAGNE AND PAUL E. BOLDT (RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, % U.S. National Museum, NHB 168, Washington, D.C. 20560; (PEB) Grassland Soil and Water Research Laboratory, Agricultural Research Service, USDA, 808 East Blackland Road, Temple, Texas 76502. Abstract.—The gall midge fauna of Baccharis in the United States now includes 12 species. Four species were previously known to attack Baccharis in the United States: three species of Neolasioptera, N. baccharicola Gagné, N. lathami Gagné, and N. rostrata Gagné, and Rhopalomyia californica Felt. Three new species and a new genus to contain one of them are described: Asphondylia bacchariola Gagné, Rhopalomyia sulcata Gagné, and Xipholasioptera Gagné and its monotypic type species Xipholasioptera ensata Gagné. Five other species, three species of Asteromyia, a Contarinia sp., and a Dasineura sp., are placed only to genus because of insufficient material for further determination. Rho- palomyia baccharis Felt is considered a junior synonym of R. californica. A key is given to the feeding niche for the 12 species. Key Words: Baccharis (Asteraceae), a genus endemic to the New World, contains 21 species that occur in the United States. These plants are woody, 0.5-4.0 m in height, and dioecious. Three species, Baccharis halimifolia L., B. neglecta Britt., and B. pteronioides DC., are weedy shrubs of economic importance. They invade pastures, rangelands, and rec- reation areas, and are unpalatable or toxic to livestock. Baccharis halimifolia, intro- duced into Australia as an ornamental but turned pest because of its invasive nature, has been controlled there to some extent by Rhopalomyia californica Felt, a gall midge introduced from California. Cultivated va- rieties of two other species, Baccharis sar- othroides Gray and B. pilularis DC., are valuable for xeroscaping and are commer- cially available in western United States (Boldt 1989). In this paper we review the gall midge Diptera, gall midges, Cecidomyiidae, fauna of Baccharis fauna of Baccharis in the United States. Twelve species are treated, including three new to science and several others that re- main undescribed. Most of the newly re- corded species and specimens in this study were collected by one of us (PEB) as part of a general survey of the phytophagous in- sects on Baccharis in southwestern United States (Boldt and Robbins 1994). The gall midges we list, except possibly the Contar- inia sp. and the Dasineura sp., appear to be specific to Baccharis and belong to species groups or genera restricted to Asteraceae. METHODS The field work for this paper was done under the direction of P. E. Boldt, the tax- onomy by R. J. Gagné. Galls were collected from Baccharis in southwestern United States from 1986 to 1993 and gall midges obtained by P. E. Boldt. Galls were re- 768 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON noved from affected plants and dissected obtain larvae, or held in small containers intil adults emerged. Specimens for study were killed and preserved in 70% ethanol. All the specimens listed are in the National Museum of Natural History (USNM), Washington, D.C. Study specimens were slide mounted for identification and scien- tific study using the method outlined in Gagné (1989, 1994). Terminology for adult morphology follows usage in McAlpine et al. (1981); that for larval morphology fol- lows Gagné (1989). KEY TO LARVAL NICHE OF NEARCTIC GALL MIDGES INFESTING BACCHARIS Details on each species in the key are under the alphabetically arranged generic and species headings that follow. [2 MInvor.onsmaturemMlOWelss coos 46 ces ete one 2 — In unopened buds, leaves or stems 2. Free in flowers Melton meperce Cre Contarinia sp. and Dasineura sp. — In swollen flower receptacles Se tet ae cht is, en NR AN Neolasioptera rostrata 3. Conical or spherical complex galls on buds or leaves — Simple swellings of leaves or stems 4. Spherical, spongy bud galls (Fig. 3) Sg ke ne ate Cplemte eee aise Rhopalomyia californica — Conical or columnar galls (Figs. 1, 4-5) .... 5 5. Conical apical gall formed by fused apical bud leaves and usually surrounded by rosette of leaves (Bice r 1) Miers. Asphondylia bacchariola and Xipholasioptera ensata — Conical or columnar galls (Figs. 4—5) Bich sEcsctrae ties ei eet eae eS Rhopalomyia sulcata 6. Swollen leaves or epidermis of green stems uh Sep ew stale sitions lev Er my - ; = LPS \ eo) ai if! og ) / 2 ‘ Sap JF] Ay \" eS 4 fT “4 ny : “is 6 if D Oo p90 v qo 9 4 < ne . "o as 2 Veron é cee ey 4 @ t ) Yay wm” . Y SE Dy vy voy yoy WY ae Y ee a = 11 SE Cie mre _—~ O10 NS See, eA ) ey ve SK a Ne Pe , N oo Sef ? ( ae ) 10 Sa eovrbomand ey \ 12 Ce is TREX. au i 5 IOs, ie AAS 14 15 Figs. 6-16. Figs. 6-11, Asphondylia bacchariola. 6, Adult mouthparts (incomplete). 7, Foretarsal claw and empodium. 8, Pupal head, ventral. 9, Same, lateral. 10, Pupal seventh and eighth abdominal segments, dorsal. 11, Larval eighth and terminal abdominal segments, dorsal. 12, Detail of larval terminal segment, dorsal. 13, Larval spatula and associated papillae. Figs. 14-16, Asteromyia spp., larval spatulae. 14, Asteromyia sp. 1 from B. angustifolia. 15, Asteromyia sp. 2 from B. pteronioides. 16, Asteromyia sp. 3 from B. salicina. Gall (Fig. 1).—Each gall is made up of several leaves forming a conical, one-celled cylinder about 4 mm long and contains a single larva or pupa. Galls appear singly at the apices of branches, are green at first, but turn brown after the single larva in each gall changes to the pupal stage. The interior gall surface is covered with a white fungal mycelium while the larva is alive. Distribution.—Arizona and western Tex- as. Hosts.—Baccharis pteronioides and pos- sibly B. bigelovii. The gall is similar on both plants but no gall midge specimens were found on the latter host to make a pos- itive identification. Remarks.—Asphondylia is a cosmopoli- tan genus with some 260 described species that mostly form galls in flowers and fruit of many families of plants. This new spe- cies falls within a group of Asphondylia that occur on Asteraceae and are characterized by having the adult labella setose and the pupal upper frontal horn bifid. A thorough comparison of this new species with other western Nearctic species from Asteraceae is VOLUME 97, NUMBER 4 not possible because most named species were described only on the basis of the adult stage. Adults of Asphondylia species have few diagnostic characters (Hawkins et al. 1986, Gagné and Waring 1990). Larvae of none of the western species from Aster- aceae have been described; pupae are known only from Asphondylia chrysotham- ni Felt and Asphondylia enceliae Felt and are similar to that of the new species. The ovipositor of A. chrysothamni is somewhat longer at 2.1 times the length of the seventh abdominal sternite, and the tarsal claws of A. enceliae are longer and thinner than on the new species. Asphondylia baccharis Kieffer and Herbst (1905) is known from Chile from a plant tentatively identified as Baccharis sp. The gall is reportedly a deformed floret 2.5—3.0 mm long, black, thin walled, and covered with hairs. The adult and pupa were only sketchily described and are pre- sumed lost (Gagné 1994), so it is impossi- ble to compare this species with A. bac- chariola. Asteromyia spp. This genus belongs to the supertribe La- siopteridi and tribe Alycaulini (Gagné 1994). Larvae of Asteromyia spp. are found in blister galls of leaves or green stems of various Asteraceae. Larvae are flattened and ovoid and have a generally reduced number of papillae with short setae. There are only three lateral papillae on each side of the spatula and only four terminal papil- lae. Gagné (1968) considered a larval speci- men from Baccharis in Texas to be Aster- omyia gutierreziae because it was similar to larvae of that species. Two other kinds of larvae based on differences in the spatulae (Figs. 14-16) have since been found. With- out pupae and adults, nothing more can be done to identify these. Because the spatulae are so different, the larvae are placed here as Asteromyia sp. 1, sp. 2, and sp. 3. Analogous blister galls on Baccharis in the Neotropics are formed by Geraldesia 771 spp. (Gagné 1994). Adults of Geraldesia are superfically similar to Asteromyia, but larvae of Geraldesia are elongate and cy- lindrical instead of ovoid and flat and pupae are pigmented and sclerotized instead of hyaline and soft as in Asteromyia. Asteromyia sp. 1, near gutiereziae Felt This segregate is known from a single larva with a three toothed spatula (Fig. 14) similar to that found in A. gutierreziae. It was placed under A. gutierreziae in Gagné (1968). The specimen was taken from a blister leaf gall on Baccharis angustifolia (probably =salicina) collected in Kenedy Co., Texas, 20-XII-1940 by R. Runyon and found in the botanical collections of the USNM. Asteromyia sp. 2 The larval spatula (Fig. 15) of this seg- regate has many fine serrations along the anterior edge. It is known from two speci- mens from Texas, each from a different spe- cies of Baccharis: B. pteronioides, Frijole Ranch, Guadalupe Mountains National Park, Culberson Co., Texas, X-16-1990, P. E. Boldt; and B. bigelovii, Point of Rocks, 12 mi. W Ft. Davis, Jeff Davis Co., Texas, VI-12-1990; PE. Boldt. Asteromyia sp. 3 The larval spatula of this segregate has a smooth anterior edge (Fig. 16). The single known larva was taken from a blister gall of B. salicina, Greenbelt Reservoir, 4 mi. N Clarendon, Donley Co., Texas, X-5-1988, P. E. Boldt and T. O. Robbins. Contarinia sp. Contarinia is a very large genus of some 250 species in the world. Larvae of most species live in leaf rolls or bud galls, but many live freely and gregariously among flower heads, as does the species on Bac- charis spp. Most are known from one host species, some feed on more than one spe- cies of a genus, and at least one is polyph- agous (Gagné 1995). Contarinia larvae PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON been taken in flower heads of several es of Baccharis spp. but have not been red to the adult stage. Many Contarinia op. that live in flower buds, including the species reared from Baccharis spp., have the hind spiracles situated on posteriorly di- rected lobes, as shown in Gagné (1989). Specimens in the USNM collected from Baccharis flower heads are as follows: Bac- charis glomeruliflora, Miami, Florida, 12- XI-1970, C. E. Stegmaier, Jr., 3 larvae; Bac- charis halimifolia, Cambridge, Dorchester Co., MD, 14-X-1984, V. Krischick; Bac- charis pteronioides, Lincoln National For- est, Carlsbad, Eddy Co., New Mexico, 21- V-1989, P. E. Boldt & R. J. Gagné, 5 larvae; B. pteronioides, Rt. 137, 1 mi. S junction to Sitting Bull Falls, Carlsbad, Eddy Co., New Mexico, 21-V-1989, P. E. Boldt & R. J. Gagné, 10 larvae; B. pteronioides, Hwy. 45, 7 mi. E Durango, Durango, Mexico, 4- VIII-1988, P. E. Boldt, 5 larvae; Baccharis salicina, Sitting Bull Falls, Lincoln Nation- al Forest, Eddy Co., New Mexico, 16-VIII- 1989, T. O. Robbins. Dasineura sp. Dasineura is a large, catchall genus with over 350 species described in the world. Many species form simple galls, including leaf rolls and swollen buds, but many live gregariously in flower heads. Larvae of an unidentified species were collected in as- sociation with Contarinia sp. larvae on flower heads of B. halimifolia, Cambridge, Dorchester Co., MD, 14-X-1984, by V. Krischick, and from B. salicina, Sitting Bull Falls, Lincoln National Forest, Eddy Co., New Mexico, 16-VIII-1989, by T. O. Rob- bins. A typical Dasineura sp. larva is fig- ured on page 67 of Gagné (1989). Neolasioptera spp. Neolasioptera is a large American genus of about 175 described species that live mainly in stem, petiole, or midrib swellings of various plants, but also in achenes or oth- er flower parts of Asteraceae. Three North American species are known from Bac- charis, as follows: Neolasioptera baccharicola Gagné This species forms a hard, woody gall at the base of the stem of B. halimifolia. It is known only from a series reared from galls taken at Virginia Beach, Virginia, V-1960, by E D. Bennett (Gagné 1971). Neolasioptera lathami Gagné Galls (Fig. 2) of this species can be found near the base of the plant as well as among apical branches of B. halimifolia, B. neg- lecta, B. salicina, and B. sarothroides. This species has two or more generations per year. It is known from New York south to Florida and west to Texas and Arizona (Gagné 1989). Palmer and Bennett (1988) recommended this species for introduction into Australia to control B. halimifolia, and Diatloff and Palmer (1987) reported on its biology and host specificity. Neolasioptera rostrata Gagné This species forms galls in receptacles of both male and female flowers (Gagné and Boldt 1989). It is widespread in the United States, from Maryland, Florida, Texas, and New Mexico where it occurs on B. glome- ruliflora, B. halimifolia, B. salicina, and B. neglecta. It also occurs in the Dominican Republic on B. myrsinities (Gagné and Boldt 1989), and P. E. Boldt has found it also in Argentina on B. pingraea and B. spartioides (Gagné 1994). Rhopalomyia spp. Rhopalomyia is a large, worldwide genus of about 150 gallforming species that most- ly occur on Asteraceae. Two species have been described from Baccharis_ pilularis from California, but one, Rhopalomyia bac- charis Felt, is considered here a junior syn- onym of R. californica. A new species is described from several species of Baccharis growing in Texas to Arizona. VOLUME 97, NUMBER 4 Rhopalomyia californica Felt This species forms lobed, succulent, usu- ally leafy bud galls (Fig. 3) on B. pilularis in California. It was imported into Australia as a potential biocontrol agent of B. hal- imifolia in 1969 and 1982. The gall midge took hold only after the 1982 introduction and since then has effectively established itself in Queensland and New South Wales (McFadyen 1985, Julien 1992, Palmer et al. 1993). It was also introduced into Texas in 1985 to control B. halimifolia and B. neg- lecta. Typical galls were observed on B. neglecta at Belton, Texas in 1986, but none was subsequently found (Boldt, unpub. data). The ecology of this species and its parasites has been investigated by Ehler (1982, 1987, 1992) and Ehler and Kinsey Ci990s T991, 1993). Thismspecies: 1s) very similar to Rhopalomyia sulcata Gagné. Dif- ferentiation between the two species is made under R. sulcata. Rhopalomyia baccharis (Felt 1908) is here considered a junior synonym of R. cal- ifornica. Rhopalomyia baccharis was. re- portedly reared from unspecified stem galls collected in December, 1885 on B. pilularis, presumably in California but unstated (Felt 1908, 1915). The two syntypes, one of each sex, that Felt studied are labelled “#3849, Deca e985 and’ “Type No. 29333 U.S.N.M.” They differ from most speci- mens of R. californica in having two in- stead of one segmented palpi. The gonosty- lus of the male syntype of R. baccharis ap- pears more bulbous than in R. californica because it is laterally compressed on the slide mount. In the USNM collection are pinned specimens with the same first label as the types and an additional one _ that reads, ““C.V. Riley Collection.’ Four males were slide-mounted to compare them prop- erly with the syntype. Their gonostyli do not appear bulbous as in the syntype but are similar to that shown for R. californica (Fig. 20). In addition, two of the specimens have two-segmented palpi like the syn- types, but two have one-segmented palpi. 773 Specimens reared from R. californica galls usually have one-segmented palpi, but oc- casional specimens are found with two-seg- mented palpi. L. E. Ehler (in litt.) writes that in outbreak conditions, galls commonly occur all along the shoots, especially when the terminals of the shoots are already galled. The unspecified “‘stem gall’? made by this R. baccharis could have come from such a gall. It appears, then, that there is no reason to maintain R. baccharis as a distinct species. Rhopalomyia sulcata Gagné, NEw SPECIES Adult.—Head: Eyes joined, about 5 fac- ets long at vertex. Male antenna with 16— 17 flagellomeres, neck of third flagellomere about 4% length of node. Female antenna with 15-16 flagellomeres without distinct necks. Frontoclypeus with 5—12 setae per side. Labellae fused, each with 1—10 setae. Palpus of one short, elliptical segment with 2—10 setae. Thorax: Wing length, d 2.9-3.3 mm; °&, 2.2—2.5 mm. Anepimeron with 12-20 setae. Claws simple. Empodia slightly longer than claws. Pulvilli about *%4 length claws. Male abdomen: Tergites 1—7 rectangular, with mostly single row setae on posterior margin intermixed with a few scales, a group of several setae laterally, and pair of anterior trichoid sensilla. Tergite 8 narrower and with sparser to no setae posteriorly and laterally but with pair of anterior trichoid sensilla. Pleura with scales. Sternites 2—8 covered with setae except for a small area about *% distance from anterior edge, all with anterior pair of trichoid sensilla. Gen- italia (Figs. 17—18): Gonocoxite cylindrical. Gonostylus elongate-cylindrical, tapering to narrow, toothed apex. Cerci rounded api- cally with several setae on disk and along apex. Hypoproct convex apically with 1—2 pairs of apical setae. Claspettes and aede- agus blunt tipped, aedeagus about as long as hypoproct. Female abdomen: Tergites 1—6 rectan- gular, with mostly single row of setae on 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 17-30. Figs. 17-18, Rhopalomyia sulcata. 17, Genitalia, dorsal. 18, Claspettes and aedeagus, dorsal. Figs. 19-20, Rhopalomyia californica. 19, Claspettes and aedeagus, dorsal. 20, Gonostylus, dorsal. Figs. 21-22, Rhopalomyia sulcata. 21, Pupal head, ventral. 22, Same, lateral. Figs. 23-24, Rhopalomyia californica. 23, Pupal antennal horns and prothorax. 24, Pupal head, lateral. Figs. 25-30, Xipholasioptera ensata, female. 25, Antenna. 26, Third flagellomere, ventral. 27, Same, dorsal. 28, Wing. 29, Tip of ovipositor, including hypoproct and fused cerci. 30, Abdomen, sixth segment to end, dorsoventral. VOLUME 97, NUMBER 4 posterior margin, rarely with scales inter- mixed, | to several setae laterally, and a pair of anterior trichoid sensilla; tergite 7 narrower than preceding tergite, with dou- ble row of posterior setae and pair of an- terior trichoid sensilla; tergite 8 wedge shaped, narrower but longer than preceding tergite and narrowest posteriorly, with sev- eral short setae on posterior third. Pleura with scales. Sternites 2—7 with sparser setae than in male, setae present only posteriorly and anteromesally, each with pair of trich- oid sensilla anteriorly. Distal half of ovi- positor about 3 times length of seventh ter- gite, fused cerci bulbous, covered with se- tae and setulae. Pupa.—Head as in Figs. 21—22. Antennal horns moderately elongate, bifid in frontal view. Face without horns. Prothoracic spi- racle short. Larva.—Unknown. Holotype-—From columnar gall on B. sal- icina, Pecos R., Loving, Eddy Co., New Mexico, 12-I'V-1990, T. O. Robbins, emerged 22-IV-1990, deposited in USNM. Paratypes.—AlIl from columnar galls: same data as holotype, pupa, 9 6, 5 2; same data as holotype except 19-VII-1990, d, pupal exuviae; same data as holotype except 20-V-1989, P. E. Boldt; B. salicina, Pecos R., Iraan, Pecos Co., Texas, 11-IV- 1990, T. O. Robbins, pupa; B. neglecta, Pe- cos R., Iraan, Pecos Co., Texas, 2-II-1990, emerged 7-II-1990, T. O. Robbins, 3 ¢ on slides; B. pteronioides, Appleton-Whittel Research Ranch, 4 mi. SE Elgin, Santa Cruz Co., Arizona, 8-III-1991, emerged 18 to 20-III-1991, P. E. Boldt and T. O. Rob- bins, 2 pupae, 4 3d, 1 2; B. salicina, Sitting Bull Falls, Lincoln National Forest, Eddy Co., New Mexico, 22-VIII-1988, PE. Boldt, 2 2; B. halimifolia, greenhouse, Temple, Texas, 21-III-1988, T. O. Robbins, 4 3,2 9%, 3 pupal exuviae. Other material examined.—Same data as holotype, 3 galls; B. neglecta, Pecos R.., Iraan, Pecos Co., Texas, 2-II-1990, emerged 7-1-1990 iO Robbins; 2°); 3°2, S’galls on pins; B. pteronioides, Appleton-Whittel folia, B. 775 Research Ranch, 4 mi. SE Elgin, Santa Cruz Co., Arizona, 8-II-1991, gall; B. pter- onioides, Hwy 61, 8 km N La Ascension, Nuevo Leon, Mexico, 28-IV-1990, P. E. Boldt, gall; B. heterophylla, Hwy 23, 7 mi S Durango, Durango, Mexico 5-VIII-1988, P. E. Boldt, galls; B. salicina, Rt. 31, under Pecos R. bridge, near Carlsbad, New Mex- ico, 20-V-1989, P. E. Boldt, @, galls; B. neglecta, Hwy 190, 1 mi. E Iraan, Pecos Co., Texas, 11-I-1990, T. O. Robbins, 10 galls; B. neglecta, Eagle pass, Maverick Co;,.) Texas;, 7-V=19945 150 Robbins; galls; B. sarothroides in greenhouse, Tem- ple Texas, T. O. Robbins and P. E. Boldt, pupae and galls; B. sarothroides, Sierra Vista, Cochise Co., Arizona, galls. Etymology.—The name sulcata is an ad- jective meaning furrowed, with reference to the furrowed surface of the gall. Distribution.—United States: Arizona, New Mexico, western and southern Texas; Mexico: Durango, Nuevo Leon. Hosts: This species is reported from B. halimifolia, B. heterophylla, B. neglecta, B. pteronioides, B. salicina, and B. sarothro- ides. The record on B. heterophylla is based only on similarity of the galls, but the other host records are based on adults and/or pu- pae. A generation of R. sulcata was reared from B. halimifolia and B. sarothroides in a greenhouse from a generation that came from field-collected galls on B. salicina. Remarks.—Males from B. pteronioides differ slightly from males from B. halimi- neglecta, and B. salicina in the shape of the hypoproct. The apex in dor- soventral view is convex in specimens from B. pteronioides, concave in the other hosts, but for now all specimens are considered to be the same species. Rhopalomyia sulcata differs from R. cal- ifornica in the male genitalia and the pupa. Rhopalomyia californica has a shorter gon- ostylus, apically oblique instead of blunt claspettes (compare Figs. 17-18 with 19— 20), and longer pupal antennal horns (com- pare Figs. 21—22 with 23-24). 776 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON \ipholasioptera Gagné, NEW GENUS ,dult (female only).—Head: antenna Figs. 25-27) with 11 flagellomeres; first and second flagellomeres connate, as wide as pedicel, the remaining flagellomeres nar- rower, cylindrical. Palpus four segmented. Labella short, hemispherical. Thorax: Wing (Fig. 28): R5 joining C near midlength of wing; M 3 + 4 and Cu weak. Scutum with four longitudinal rows of setae and scales, the rows of scales al- most contiguous. Anepisternum with scales on dorsal third; mesepimeron covered with setae and scales; katepisternum bare. Tarsal claws with large tooth; empodia as long as claws. Abdomen (Figs. 29-30): First through sixth tergites rectangular, more than twice as wide as long, with anterior pair of trich- oid sensilla, posterior margin with single, sparse row of setae, and otherwise covered with scales; seventh tergite rectangular, about twice as long as wide, with anterior pair of trichoid sensilla, posterior margin with single row of setae, and covered with scales on posterior two-fifths; eighth tergite elongate, pointed posteriorly, about twice as long as wide, not including the narrow, an- terior extensions, with anterior pair of trich- oid sensilla as only vestiture; second to sixth sternites square, without anterior pair of trichoid sensilla but with setae laterally and posteriorly and scales elsewhere; sev- enth sternite triangular, narrowest anterior- ly, somewhat longer than wide, posterior margin with single row of setae, and cov- ered with scales on posterior two-fifths; eighth sternite about twice as long as wide, narrowing posteriorly, without vestiture; Ovipositor elongate, posterior half bilater- ally flattened, strongly sclerotized, pointed apically. Pupa and larva.—Unknown. Type species.—Xipholasioptera ensata Gagné. Etymology.—The name Xipholasioptera combines the Greek “‘xiphos”’ for sword, with reference to the shape of the oviposi- tor, and “‘lasioptera”’ for the scale-covered wing. The latter segment is a common suf- fix in the tribes Alycaulini and Lasiopterini. The name is feminine. Remarks.—This genus belongs to the tribe Alycaulini of the supertribe Lasiopter- idi, as defined in Gagné (1994), because of the thick covering of scales, the very short R5 wing vein, the short antenna, and the lack of two lateral groups of large setae on the eighth abdominal segment. This genus differs from all known Alycaulini in the shape of the postabdomen and ovipositor (Fig. 30) and in the lack of the anterior pair of trichoid sensillae on each of the abdom- inal sternites. The strongly modified abdo- men so obscures relationships that the clos- est relative of Xipholasioptera is not appar- ent. All other genera of Alycaulini except Protaplonyx and Edestosperma have soft, cylindrical ovipositors. Protaplonyx, known form one species on a Sarcobatus (Cheno- podiaceae), has only the eighth abdominal tergite of the female modified and has a very different, short, strongly sclerotized ovipositor (Gagné 1969); Edestosperma, known from one species reared from seeds of a grass, has the seventh and eighth ter- gites and eighth sternite modified and a short, bilaterally flattened, but soft ovipos- itor (Gagné 1969). Xipholasioptera ensata Gagné, NEW SPECIES Adult (female only).—Antenna as in Figs. 25-27. Wing length, 1.7 mm, venation as in Fig. 28. Abdomen as in Figs. 29-30; dorsal area of fused cerci weakly striate, de- void of setae; a longitudinal row of setae present on each side, below which cerci pit- ted and not so strongly pigmented; hypo- proct short, narrow, with apical pair of short setae. Pupa and larva.—Unknown. Etymology.—The name ensata is a Latin adjective meaning swordlike, after the shape of the ovipositor. Holotype.—Female, ex branch apex of Baccharis pteronioides, Pine Springs, Gua- VOLUME 97, NUMBER 4 dalupe Mountains National Park, Texas, V- 2-1988, P. E. Boldt, deposited in U.S. Na- tional Museum. Remarks.—The single known specimen was taken in association with leaf fascicle galls made by Asphondylia_ bacchariola. Although this species is known from a sin- gle female, it is certainly distinctive and likely to be found again on Baccharis. The Ovipositor is shaped for piercing and, there- fore, the eggs are presumably laid in plant tissue. ACKNOWLEDGMENTS We thank T. O. Robbins of the Grassland, Soil and Water Laboratory, USDA, ARS, Temple, Texas, for technical assistance, L. Heath Lawrence of the Systematic Ento- mology Laboratory for Figs. 1—5, officials and personnel of Guadalupe Mountains Na- tional Park, National Park Service, Salt Flat, Texas and Appleton-Whittell Research Ranch, National Audubon Society, Elgin, Arizona, for assistance and permission to collect specimens; P. Malikul for preparing the slide mounts; and L. E. Ehler, of the University of California, Davis, G. C. Steyskal, Gainesville, Florida, S. Nakahara and A. L. Norrbom of the Systematic En- tomology Laboratory, and an anonymous reviewer for their comments on drafts of the manuscript. LITERATURE CITED Boldt, P E. 1989. Baccharis (Asteraceae), a review of its taxonomy, phytochemistry, ecology, eco- nomic status, natural enemies and the potential for its biological control in the United States. Texas Agricultural Experiment Station Miscellaneous Publication 1674: 1—32. Boldt, P. E., and T. O. Robbins. 1994. Phytophagous insect fauna of Baccharis salicina, B. pteronioides and B. bigelovii (Asteraceae) in the southwestern United States and northern Mexico. Environmen- tal Entomology 23: 47-57. Diatloff, G. and W. A. Palmer. 1987. The host spec- ificity of Neolasioptera lathami Gagné (Diptera: Cecidomyiidae) with notes on its biology and phe- nology. Proceedings of the Entomological Society of Washington 89: 122-125. Ehler, L. E. 1982. Ecology of Rhopalomyia califor- NSE nica Felt and its parasites in an urban environ- ment. Hilgardia 50: 1—32. 1987. Ecology of Rhopalomyia californica Felt at Jasper Ridge (Diptera: Cecidomyiidae). Pan-Pacific Entomologist 63: 237-241. 1992. Guild analysis in biological control. Environmental Entomology 21: 26—40. Ehler, L. E. and M. G. Kinsey. 1990. Influence of gall size on survival and fecundity of Rhopalomyia californica (Diptera: Cecidomytidae), a biologi- cal-control agent for Baccharis halimifolia (As- teraceae). Environmental Entomology 19: 1558-— 1565. 1991. Ecological recovery of a gall midge and its parasite guild following disturbance. En- vironmental Entomology 22: 1295-1300. 1993. Spatial distribution of progeny in a parasitoid of a gall midge and its relevance to ap- plied biological control. Biological Control 3: 116-126. Felt, E. P. 1908. Appendix D. New York State Mu- seum Bulletin 124: 286-422, pls. 33-44. Gagné, R. J. 1968. A taxonomic revision of the genus Asteromyia (Diptera: Cecidomyiidae). Miscella- neous Publications of the Entomological Society of America 6: 1-40. 1969. A tribal and generic revision of the Nearctic Lasiopteridi (Diptera: Cecidomyiidae). Annals of the Entomological Society of America 62: 1348-1364. 1971. Two new species of North American Neolasioptera from Baccharis (Diptera: Cecido- mylidae—Compositae). Proceedings of the Ento- mological Society of Washington 73: 153-157. 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ithaca, New York. xi and 356 pp. and 4 pls. 1994. The Gall Midges of the Neotropical Region. Cornell University Press, Ithaca, New York. xv and 352 pp. . 1995. Contarinia maculipennis (Diptera: Ce- cidomyiidae), a new pest of Dendrobium orchids in continental USA. Bulletin of Entomological Research 85: 209-214. Gagné, R. J. and P. E. Boldt. 1989. A new species of Neolasioptera (Diptera: Cecidomyiidae) from Baccharis (Asteraceae) in southern United States and the Dominican Republic. Proceedings of the Entomological Society of Washington 91: 169— 174. Gagné, R. J. and G. L. Waring. 1990. The Asphon- dylia (Cecidomyiidae: Diptera) of creosote bush (Larrea tridentata: Zygophyllaceae) in North America. Proceedings of the Entomological So- ciety of Washington 92: 649-671. Hawkins, B. A., R. D. Goeden, and R. J. Gagné. 1986. Ecology and taxonomy of the Asphondylia spp. (Diptera: Cecidomyiidae) forming galls on Atri- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON , spp. (Chenopodiaceae) in southern Califor- iia. Entomography 4: 55-107. Julien, M. H. 1992. Biological control of weeds. A world catalogue of agents and their target weeds. Third ed. CAB Intern. Kieffer, J.-J. and P. Herbst. 1906. Description de gal- les et d’insectes gallicoles du Chili. Annales de la Société Scientifique de Bruxelles. 30: 223-236, 1 pl. McAlpine, J. E, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds. 1981. Manual of Nearctic Diptera, vol. 1. Re- search Branch, Agriculture Canada, Monograph 27. vi + 674 pp. McFadyen, P. J. 1973. Introduction of the gall fly, Rhopalomyia californica, from the USA into Aus- tralia for the control of the weed Baccharis hal- imifolia, pp. 779-787. In Delfosse, E., ed., Pro- ceedings VI International Symposium of Biolog- ical Control of Weeds. Vancouver, Canada. Palmer, W. A. and ED. Bennett. 1988. The phytoph- agous insect fauna associated with Baccharis hal- imifolia L. in the eastern United States. Proceed- ings of the Entomological Society of Washington. 90: 216-228. Palmer, W. A., G. Diatloff, and J. Melksham. 1993. The host specificity of Rhopalomyia californica Felt (Diptera: Cecidomyiidae) and its importation into Australia as a biological control agent for Baccharis halimifolia L. Proceedings of the En- tomological Society of Washington. 95: 1-6. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 779-790 LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF UROPHORA TIMBERLAKE! BLANC AND FOOTE (DIPTERA: TEPHRITIDAE) ON NATIVE ASTERACEAE IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN, DAvip H. HEADRICK, AND JEFFREY A. TEERINK Department of Entomology, University of California, Riverside, California 92521. Abstract.—Urophora timberlakei Blanc and Foote is a bivoltine, probably trivoltine, tephritid which develops in the flower heads of Acamptopappus, Amphipappus, Chryso- thamnus, and Haplopappus spp. in southern California. For the first time, the egg, second and third instar larvae, and puparium are described, and the larvae and puparium are figured. Distinctive morphological features noted for the immature stages include eggs that are smaller and of a shape different from Palearctic Urophora. The third instars resemble certain described Palearctic Urophora, but they differ from larvae of other Ne- arctic tephritids in having a bluntly truncated, dark brown to black caudal segment covered by minute dome-shaped papillae and bearing several deep pits mediad and ventrad of the posterior spiracular plates. The verruciform sensilla circumscribing the prothorax and the wedge-shaped acanthae that circumscribe the meso-, metathoracic, and abdominal seg- ments also are distinctive features. The larvae feed mainly on the ovules and soft achenes. Pupariation occurs in the larval feeding chamber among fragments of achenes. Premating and mating behaviors are described for the first ttme for any Nearctic Urophora, distin- guished only by the males displaying abdominal pleural distensions throughout courtship and copulation. Eupelmus sp. (Hymenoptera: Eupelmidae) is reported as a solitary, en- doparasitic, larval-pupal or pupal parasitoid. Key Words: Insecta, Urophora timberlakei, nonfrugivorous Tephritidae, mating behav- ior, immature stages, Asteraceae, flower-head feeding eight introduced Old World species, were reviewed and distinguished by Foote et al. The genus Urophora (Diptera: Tephriti- dae) is a polyphyletic assemblage of flori- vorous and/or gallicolous species of fruit flies best known from the Palearctic Re- gion, where several species infesting knap- weeds and thistles [Asteraceae: Cynareae (Centarinae and Carduinae, respectively)] were extensively studied and subsequently exported to North America for biological control of weeds (Varley 1937, Zwéolfer 1988, Harris 1989, Julien 1992). Known species of New World Urophora were de- picted in a pictorial key by Steyskal (1979), and those found North of Mexico, including (1993). Little is known about the biologies of the eight indigenous North American species of Urophora (Foote et al. 1993); however, all seven California species are non-gallicolous, seed-feeders in flower heads of hosts in the subtribe Solidagininae of the tribe Astereae (Asteraceae) (Goeden 1987, unpublished data). This paper describes the life history and immature stages of Urophora timberlakei Blanc and Foote, the first North American species of Urophora to be studied in any detail. RO PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MATERIALS AND METHODS his study began in 1989 and was based part on dissections of selected subsam- ples of mature flower heads of Asteraceae infested by U. timberlakei from among many samples collected annually through- out California in the manner described by Goeden (1985, 1987, 1992). Adults studied were reared in wooden, 35 X 32 X 35-cm, muslin cloth-backed, glass-topped sleeve cages in the insectary of the Department of Entomology, University of California, Riv- erside, from 1|-liter samples of mature flow- er heads at 14-h photophase and 27 + 1°C. Additional adults and parasitoids were reared from larvae and puparia of U. tim- berlakei dissected from flower heads and held separately in cotton-stoppered glass shell vials within humidty chambers in the laboratory at 23 + 2°C and 76% R.H. All larvae and five puparia dissected from flow- er head samples were fixed in 70% EtOH for scanning electron microscopy (SEM). Specimens for SEM later were rehydrated to distilled water in a decreasing series of acidulated EtOH, rinsed in hexanes, and post-fixed in 2% aqueous osmium tetroxide for 24 h. They were then dehydrated through an increasing series of acidulated EtOH, critical-point dried, mounted on stubs, sputter-coated with a gold-palladium alloy, and studied with a JEOL JSM C-35 scanning electron microscope in the De- partment of Nematology, University of Cal- ifornia, Riverside. SEM micrographs were prepared on Polaroid® P/N 55 film at 15 kV accelerating voltage. Most adults reared from isolated puparia, as well as overwintered adults swept from preblossom and early-blossom Acampto- pappus sphaerocephalus (Harvey and Gray) Gray | km N of Snow Creek Village, San Gorgonio River flood plain, 350m el- evation, Riverside Co., in April, 1992-94, were individually caged in 850-ml, clear- plastic, screened-top cages each fitted with a cotton wick and basal water reservoir and provisioned with a strip of paper toweling impregnated with yeast hydrolyzate and su- crose. These cagings were used for longev- ity studies and oviposition tests. Virgin male and female flies of known ages ob- tained from emergence vials were paired in clear-plastic, disposable petri dishes provi- sioned with a flattened, water-moistened pad of absorbant cotton spotted with honey for direct observations, videorecording, and still-photography of their general behavior, courtship, and copulation (Headrick and Goeden 1991). Three trials with U. timber- lakei reared from the flower heads of Chrysothamnus teretifolius (Durand and Hilgard) Hall were held together for a max- imum of 14 days, and observations were made as opportunity allowed throughout each day. Plant names used in this paper follow Munz and Keck (1959) and Munz (1968, 1974); names for flower head parts follow Hickman (1993). Tephritid names and ana- tomical terms follow Foote et al. (1993); no- menclature used to describe the immature stages follows Headrick and Goeden (1990, 1991), Goeden and Headrick (1990, 1991a, b; 1992), and the telegraphic format of Goe- den et al. (1993, 1994). Means + SE are used throughout this paper. Voucher speci- mens of reared adults of U. timberlakei and its parasitoids reside in the research collec- tions of RDG; preserved specimens of lar- vae and puparia are stored in collections of immature Tephritidae maintained by JAT. RESULTS AND DISCUSSION Taxonomy Urophora timberlakei was described by Blanc and Foote (1961). Photographs of the characteristically hyaline wing occurring in both sexes of this species were provided by Foote and Blanc (1963) and Foote et al. (1993). The aculeus of the female was il- lustrated by Steyskal (1979). The adults of U. timberlakei heretofore have not other- wise been illustrated nor the immature stages described or illustrated. Egg.—Egg body smooth, shiny, white, elongate-ellipsoidal; anterior end tapered, VOLUME 97, NUMBER 4 bearing peg-like, 0.02 mm-long pedicel; posterior end tapered, smoothly rounded. Twenty-one ova dissected from two field- collected females were 0.55 + 0.004 (range, 0.52—0.58) mm long and 0.15 + 0.004 (range, 0.12—0.20) mm wide. Wadsworth (1914) described the ova of Urophora solstitialis (L.) as being white, smooth, cresent-shaped, 0.70 mm long and 0.20 mm wide. Persson (1963) described U. solstitialis eggs as shiny white, 0.80 mm long, 0.20 mm wide, and having the same shape as those of U. stylata. The eggs of U. timberlakei are smaller in size and differ in shape from the Palearctic species described above, being more elongate-ellipsoidal than cresent-shaped. Third instar—Third instar elongate-cy- lindrical, tapering slightly anteriorly, blunt- ly truncated posteriorly (Fig. 1A); integu- ment white, venter of meso-, metathorax with brown infuscation; caudal segment dark brown or black (Fig. 4B, C); minute acanthae circumscribe meso-, metathorax and abdominal segments (Fig. 1A); gnath- ocephalon conical, smooth, lacking rugose pads (Fig. 1B); paired dorsal sensory or- gans, dorsomediad of anterior sensory lobe, consist of a single dome-shaped papilla, (Fig. 1C-1, 1D-1); anterior sensory lobes separated by medial depression (Fig. 1C-2), each bearing terminal sensory organ (Fig. 1D-2), pit sensory organ (Fig. 1D-3), lateral sensory organ (Fig. 1D-4), and supralateral sensory organ (Fig. 1D-5); stomal sense or- gan ventrolaterad of anterior sensory lobe near mouth lumen (Fig. 1C-3, 1D-6); lateral and ventrolateral sensilla verruciform with a central pore (Fig. 1B-1); mouth hooks bi- dentate, teeth stout, conical (Fig. 1C-4); median oral lobe tapered anteriorly, later- ally flattened, attached to labial lobe (Fig. 1C-5); prothorax smooth, minute acanthae anteroventrally (Fig. 1B), verruciform sen- silla circumscribe segment, arranged 3-ven- tral, 3-medial, 2-dorsomedial, 3-dorsolater- al (Fig. 1B-2); anterior thoracic spiracles dorsolateral on posterior margin of protho- rax each bearing two papillae (Fig. 1B-3, 781 E); lateral spiracular complex with spiracle (Fig. 1F-1) and two verruciform sensilla on meso- and metathorax, a single verruciform sensillum on abdominal segments (Fig. 1 F- 2); minute acanthae which circumscribe TII-AVII wedge-shaped with ovoid bases (Fig. 1F-3); caudal segment covered by minute, dome-shaped acanthae (Fig. 1G-1), and numerous deep pits located dorsome- diad, mediad and ventrad of posterior spi- racular plates (Fig. 1G-2); posterior spirac- ular plates (Fig. 1G-3, H) distinct, raised, spheroid, bear three oval rimae ca. 0.02 mm in length (Fig. 1H-1), and four, spine-like interspiracular processes ca. 0.006 mm in length (Fig. 1H-2); stelex sensilla surround margin of caudal segment. Urophora timberlakei larvae fit the gen- eral description of other Urophora species (Wadsworth 1914, Varley 1937, Persson 1963), and appear closely allied with U. quadrifasciata. Both species have anterior spiracles with two papillae, the integument roughened with minute tubercles and the posterior spiracular plates without a pattern of fine grooves or pale border (Varley 1937). Urophora timberlakei is different from other Nearctic, non-frugivorous, tephritid species previously examined (Goeden and Headrick 1990; 19914, b, 1992, ‘Goeden’ et al. 1993, 1994, Green et al. 1993, Headrick and Goe- den 1991, 1993, Novak and Foote 1968, Steck and Wharton 1986) in having a blunt- ly truncated, dark brown to black caudal segment covered by minute dome-shaped papillae and bearing numerous deep pits dorsomediad, mediad and ventrad of the posterior spiracular plates. The caudal seg- ment of Euaresta stigmatica is brown, but smooth and rounded posteriorly (Headrick et al., in press). The posterior spiracular plates appear to be similar to other described spe- cies of Urophora (Wadsworth 1914, Varley 1937, Persson 1963), but differ from other Tephritidae in being distinctly raised and spheroid in shape (Goeden and Headrick 1990, Goeden et al. 1994, Green et al. 1993, Steck and Wharton 1986). The verruciform sensilla circumscribing the prothorax and the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON A , 4 a. | , . p ee ) pa W Oe , "BSRU xi300' o911 16.00 F @8kU X1500. aaa9 ° ioaU Fig. 1. Third instar larva of U. timberlakei: (A) habitus, anterior to left; (B) gnathocephalon, prothorax, anterior view, | lateral and ventrolateral sensilla, 2—verruciform sensillum, 3—anterior thoracic spiracle; (C) gnathocephalon, anterior view, 1—dorsal sensory organ, 2—anterior sensory lobe, 3—stomal sense organ, 4— mouth hooks, 5—median oral lobe; (D) anterior sensory lobe, |—dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4 lateral sensory organ, 5—supralateral sensory organ, 6—stomal sense organ; (E) anterior thoracic spiracle; (F) lateral spiracular complex, first abdominal segment, 1—spiracle, 2—verruci- VOLUME 97, NUMBER 4 wedge-shaped acanthae which circumscribe the meso-, metathoracic, and abdominal seg- ments are morphological features which may be distinctive of the North American Uro- phora. Second instar.—Second instar cylindri- cal, tapered anteriorly, bluntly truncated posteriorly (Fig. 2A); integument white throughout (Fig. 4A); gnathocephalon con- ical, laterally flattened, smooth, lacking ru- gose pads (Fig. 2C); paired dorsal sensory organs consist of single round papilla, dor- somediad of anterior sensory lobe (Fig. 2B- 1, 2C-1); anterior sensory lobes separated by medial depression (Fig. 2C-2), and bear terminal sensory organ (Fig. 2B-2), pit sen- sory organ (Fig. 2B-3), lateral sensory or- gan (Fig. 2B-4) and supralateral sensory or- gan (Fig. 2B-5); stomal sense organs indis- tinct, ventrolaterad of anterior sensory lobes, near mouth lumen (Fig. 2B-6, 2C-3); mouth hooks bidentate, teeth sharply curved (Fig. 2C-4); median oral lobe ta- pered apically, laterally flattened (Fig. 2C- 5); small integumental petals dorsad of mouth lumen (Fig. 2C-6); prothorax smooth with minute acanthae ventrad of gnatho- cephalon; anterior thoracic spiracles consist of two papillae (Fig. 2D); lateral spiracular complex not seen; caudal segment wrinkled with several large depressions mediad of posterior spiracular plates (Fig. 2E); poste- rior spiracular plates bear three oval rimae ca. 0.007 mm in length (Fig. 2F-1), and four, spine-like interspiracular processes ca. 0.004 mm in length (Fig. 2F-2); stelex sen- silla surround margin of caudal segment. The second instar larva is more cylindri- cal than the third instar in general body shape. Most morphological features of the second instar are similar in shape and place- ment to those of the third instar, but not as structurally distinct as on the mature larva. The caudal segment of the second instar lar- e 783 va differs from that of the third instar larva. In the second instar the caudal segment is truncated, white in color, wrinkled and with several large circular depressions. The cau- dal segment of the second instar lacks the minute dome-shaped acanthae and deep pits mediad and ventrad of the posterior spirac- ular plates which are present on the third instar. Because of the wrinkled nature of the second instar larvae prepared for SEM, the lateral spiracular complex and the area where prothoracic verruciform sensilla would occur were not observed. Puparium.—Puparium light brown, an- terior end dark brown, caudal segment dark brown to black; elongate cylindrical, taper- ing anteriorly, truncated posteriorly (Fig. 3A), minute acanthae circumscribe seg- ments (Fig. 3A); anterior end bears the in- vagination scar (Fig. 3B-1), raised bilobed anterior thoracic spiracles (Fig. 3B-2), and round verruciform prothoracic sensilla (Fig. 3B-3); posterior end comprising of minute dome-shaped acanthae (Fig. 3C-1) and nu- merous deep pits dorsomediad, mediad and ventrad to the posterior spiracular plates (Fig. 3C-2); posterior spiracular plates bear three oval rimae ca. 0.02 mm in length (Fig. 3D-1), four spine-like interspiracular pro- cesses ca. 0.004 mm in length (Fig. 3D-2) and the ecdysial scar (Fig. 3D-3); stelex sensilla surround margin of caudal segment (Fig. 3C-3). Three puparia averaged 2.42 + 0.62 (range, 2.33—2.54) mm in length and 0.97 + 0.01 (range, 0.95—0.98) mm in width. DISTRIBUTION AND HOSTS The North American distribution of U. timberlakei North of Mexico was mapped by Foote et al. (1993) to include California, Colorado, Idaho, Oregon, and Utah, adding Colorado and Utah to the distribution given by Steyskal (1979). form sensillum, 3—wedge-shaped acanthae; (G) caudal segment, 1—dome-shaped acanthae, 2—deep pits, 3— posterior spiracular plate; (H) posterior spiracular plate, 1—rima, 2—interspiracular process. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SKY A126 15KVU Keeoe 68006 16. Al ye iSKV X3200 goa3 i8.6U F Fig. 2. Second instar larva of U. timberlakei: (A) habitus, anterior to left; (B) anterior sensory lobe, 1— dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4 lateral sensory organ, 5—supralateral sensory organ, 6—stomal sense organ; (C) gnathocephalon, anterolateral view, 1—dorsal sensory organ, 2— anterior sensory lobe, 3—stomal sense organ, 4—mouth hooks, 5—median oral lobe, 6—integumental petals; (D) anterior thoracic spiracle; (E) caudal segment; (F) posterior spiracular plate, 1—rima, 2—interspiracular process. VOLUME 97, NUMBER 4 «<3 SKU X22" Bigs 3: SKU K1866 16. 6U D 682 Puparium of U. timberlakei: (A) habitus, anterior to right; (B) anterior end, 1—invagination scar, 2— anterior thoracic spiracle, 3—verruciform sensillum; (C) caudal segment, 1—dome-shaped acanthae, 2—deep pits, 3—stelex sensillum; (D) posterior spiracular plate, |—rima, 2—interspiracular process, 3—ecdysial scar. Goeden (1987) reported this tephritid as reared from flower heads of Acamptopap- pus shockleyi Gray, A. sphaerocephalus, Chrysothamnus nauseosus (Pallas) Britton, C. teretifolius, C. viscidiflorus (Hooker) Nuttall, Haplopappus cuneatus Gray, and H. laricifolius Gray. To these confirmed re- cords we add the following new rearing re- cords from ungalled flower heads: Amphi- pappus freemontii Torrey and Gray, 9 6 and 7 2, W. of Homer Mountain at 884-m elevation, NE San _ Bernardino Co., 5-V219883/G: .albidus. Jones) Greene;2. 2; Fish Slough, 9 km N of Bishop at 1,225 m, Inyo Co., 9.x.1990; C. paniculatus (Gray) Hall, 1 d and 1 2, N of Pioneertown on N rim of Chaparrosa Wash at 1,356 m, SW San Bernardino Co., 31.x.1990; H. cooperi formosa (Coquillett), U. (Gray) Hall, | 5, along Black Canyon Rd., E of Colton Hills, E. Mojave Desert at 1,200 m, NE San Bernardino Co., 30v1991: As first noted by Goeden (1987), and as these new rearing records confirm, the host- plants of U. timberlakei, apparently like those of its native congeners in California, all belong to the Subtribe Solidagininae of the Tribe Astereae (Munz and Keck 1959). Most of these new records, like those re- ported by Goeden (1987), involve only a few flies reared from bulk samples each containing hundreds of flower heads of its mostly small-headed hosts. Like other oli- gophagous Urophora in California, e.g. U. timberlakei has been reared alone, but more commonly 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON along with other congeneric or nonconge- ¢ species from single samples of host flower heads (synphagy), sometimes as the dominant phytophage, but more commonly subordinate to one or more species of Neas- pilota, Tephritis, or Trupanea (Goeden 1987, unpublished data). Thus, U. timber- lakei was only occasionally swept, reared, or otherwise encountered in nature, usually in synphagy with other species, and there- fore was not easily studied, even in Cali- fornia, where it first was described and since has been reported most frequently (Foote and Blanc 1963, Foote et al. 1993). BIOLOGY Egg.—One of eight (12.5%) infested heads of A. shockleyi, the host with the largest-sized head in California, and only one of 61 (2%) infested smaller heads of C. teretifolius and C. viscidiflorus examined during the study period contained two lar- vae of U. timberlakei. Therefore, single eggs usually are oviposited by individual females in preblossom heads, but none was recovered during the present study. Larva.—All second and third instar lar- vae observed in dissected heads had con- fined their feeding to ovules and soft achenes of individual flower heads (Fig. 4A, B). Larger larvae were found in feeding cavities located between adjacent achenes or were confined to individual achenes, and only partially damaged other ovules as ear- ly instars. The feeding cavities of U. tim- berlakei always appeared dry, which sug- gested that sap was not an important sup- plemental food source for this tephritid, un- like Neaspilota viscidiflorus Quisenberry (Goeden and Headrick 1992) and several other, ovule and soft achene-feeding tephri- tids studied in California to date (Headrick and Goeden 1990, 1991; Goeden and Head- rick 1992). In eight (4%) infested heads of A. shockleyi of 200 examined, third instars attacked an average of only 1.3 + 0.2 (range, 1—2) achenes of an average total of 38 + 1.7 (range, 32-46) achenes, or about 3% of the achenes therein. The receptacle was slightly scored in only one of these eight flower heads, which, again, are the largest among U. timberlakei’s known hosts in California. On the other hand, an average of 3.5 + 0.4 achenes were damaged and the receptacles scored in eight, small, infested heads of C. viscidiflorus. Four other sub- samples of 200 heads each of C. viscidiflo- rus from different locations contained only 0, 2, 3 and 4 (0%, 1%, 1.5%, and 2%, re- spectively) immature U. timberlakei. In a subsample of 400 heads of another small- headed host, C. teretifolius, 28 (7%) were infested by U. timberlakei larvae that whol- ly or partly damaged an average of 2.2 + 0.12 (range, 1—4) ovules and soft achenes or about half of an average total of 5.0 + 0.13 (range, 3—6) achenes in each head. The receptacles were slightly scored in only two (7%) of these 28 infested heads; whereas, all four achenes in each of four infested heads were completely destroyed, and three of the receptacles scored in a different sub- sample of 200 heads of C. teretifolius from a different location. The fully grown third instar (Fig. 4C) oOverwinters inside the feeding cavity within the excavated head which usually remains attached to the dormant host plant, as with N. viscidiflorus (Goeden and Headrick 1992), but without forming a hard, protec- tive cell like the latter species. Among flo- rivorous, southern California Tephritidae, only Urophora, Neaspilota, and possibly Xenochaeta spp. mainly overwinter within dead flower heads as nonfeeding, diapaus- ing, fully grown larvae (Goeden and Head- rick 1992, Goeden and Teerink, unpub- lished data); whereas, most other florivo- rous species emerge in the spring, summer, or fall and overwinter as adults, e.g., Neo- tephritis, Paracantha, Stenopa, Trupanea, and Tephritis spp., or emerge from heads as larvae, drop to the ground, and overwinter as buried puparia, e.g., Orellia and many Paroxyna spp. (Goeden, Teerink, and Head- rick, unpublished data). Puparium.—Pupariation occurs in spring after overwintering inside the dead flower VOLUME 97, NUMBER 4 787 Fig. 4. viscidiflorus; (B) third instar larva feeding in flower head of C. teretifolius; (C) nonfeeding, overwintering, prepupal, third instar dissected from flower head; (D) empty puparium in flower head of C. viscidiflorus; (E) female; (F) mating pair. Bars = | mm. head. The larva reverses its orientation 180° and the puparium is formed with its anterior end facing outward, away from the recep- tacle (Fig. 4D). The adult emerges through the pappus bristles when exiting the pupar- ium and flower head (Fig. 4D). Adult.—The superficial resemblance of U. timberlakei adults to Agromyzidae in their lack of wing pattern and in their body size, pigmentation and pattern is noteworthy as possibly representing convergence or mimicry, but otherwise remains unexplained and unaddressed by the present study (Fig Life stages of U. timberlakei: (A) Second instar larva in attacked immature achenes of Chrysothamnus 4E, F). Males and females emerged together from mid-November to mid-January in the insectary from bulk samples of flower heads collected in mid-October, 1990 and 1991, that initially contained overwintering larvae. This emergence probably was abnormally early, as the spring hosts of U. timberlakei first bloom in April and May. All told, 23 3 and 22 2 and 20 ¢ and 10 2 were reared from these two samples, respectively. The most adults ever recovered by RDG from a single sample was 42 ¢d and 31 @ reared from a liter sample of heads of C. viscidiflo- 188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON s collected in Landers Meadow, Sequoia Forest, Kern Co., on 3.ix.1981. These 4 other published (Goeden 1987) and un- published rearing records of RDG suggested a slightly male-biased sex ratio for U. tim- berlakei. Four of five females newly emerged from heads of C. viscidiflorus in the insectary contained two to <60, fully formed ova; the fifth female contained im- mature ovaries and considerable fat stored as 0.04-0.28-mm, irregular globules in the he- molymph. This suggested that the former fe- males may have remained inside the flower heads, perhaps still inside their puparia, for undefined periods after they had molted and sexually matured and before they emerged. Like the early insectary emergence noted above, this suspected delay may have rep- resented laboratory behavioral artifacts. Wing displays.—Adults exhibited hama- tion as described for N. viridescens (Goe- den and Headrick 1992). The wings in both sexes were held away from the body at ca. 45° without supination when at rest, then extended back and forth through 25—30° in a plane parallel to the substrate. Both sexes exhibited this display throughout the day concurrent with other behaviors, i.e. groom- ing, resting and feeding. Males also main- tained hamation displays while facing a fe- male for courtship. Swaying.—Both sexes of U. timberlakei occasionally displayed swaying during ha- mation while facing another individual. In- dividuals swayed less during hamation when no other fly was nearby. Swaying during hamation sometimes preceded or followed side-stepping. Both sexes side- stepped during hamation while facing other individuals as described for N. viridescens (Goeden and Headrick 1992). The display- ing adult moves sideways, up to one body length, in a semicircle and then back while facing the other individual. This movement continued for variable times and ended with one or the other fly decamping. Courtship.—Male aggregative displays occurred throughout the day and lasted for several hours. Males typically held their black abdomens slightly raised with the pleura fully distended. The abdominal pleu- ra in U. timberlakei males undulated in waves of contraction that moved along the pleura from anterior to posterior. U. timber- lakei males continued to hold their pleura distended throughout courtship and copu- lation, unlike other tephritid males that dis- play abdominal pleural distension (cf., Headrick and Goeden 1990, 1991, 1993, Goeden and Headrick 1992, Headrick et al. 1993). Males occasionally interrupted their courtship displays to groom or feed. Males also commonly hung upside down in arenas visually following females while displaying hamation and abdominal pleural distension. Males displayed hamation and abdominal pleural distension during all hours of the day. Courtship was brief. Males approached females and abruptly jumped onto their dor- sa with or without having exhibiting any prior wing displays toward them (n = 6). Females resisted males and jumped away from their mounting attempts. Males that were able to maintain their grip began cop- ulatory induction behavior (n = 4). Copulatory induction behavior (CIB).— Mounted males grasped females with the front legs gripping the dorsum of the ab- domen near the thorax with the foreclaws hooked into the abdominal pleura. The mid- dle legs wrapped around the base of the oviscape and the hind legs bent underneath the oviscape. Males used their hind legs to first raise the oviscape and place its apex against the epandrium. Generally, females strongly resisted males by using their hind legs to push at them. If males remained, the hind legs were rubbed vigorously against the sides and venter of the oviscape. When viewed from above, males shook rapidly from side-to-side through 30° over the fe- male dorsum, as described for N. virides- cens (Goeden and Headrick 1992). Copulation.—The oviscape of Urophora females is relatively longer than that of most other species studied by us, except for Procecidochares stonei Blanc and Foote (Green et al. 1993). This caused some VOLUME 97, NUMBER 4 unique problems for U. timberlakei males, as also observed with P. stonei. When re- ceptive females exserted the aculeus and the male epandrium was not situated to receive its apex, the aculeus usually slid beneath the epandrium and posteriad of the male terminalia. This typically invoked vigorous CIB by males and repositioning of the ter- minalia. Males rocked back and forth over the female dorsum while the females con- tinued to struggle. Males successful in en- gaging the aculeus tip were pushed farther backward as females exserted the ovipositor to its full length. The front legs of the male then grasped the base of the oviscape, the middle legs grasped the apex of the ovis- cape, and the hind tarsi were pressed against the fully exserted aculeus. This po- sition was difficult for males to maintain and they struggled to move forward on the female by pressing the terminalia against the aculeus. Rhythmic retraction and exer- tion of the aculeus then began and contin- ued until the final copulatory position was attained (Fig. 4F) (cf., Headrick and Goe- den 1995). Two copulations were timed for their full duration and lasted | and 1.5 h. Seasonal history.—Urophora timberlakei is at least bivoltine, probably trivoltine (see below), and synovigenic in southern Cali- fornia. Eleven males lived an average of 39 + 5 days (range, 18—66 days); 14 females, 31 + 6 days (range, 8—88 days) under in- sectary conditions. These mean longevities are longer than those of proovigenic fe- males, e.g., P. stonei (Greene et al. 1993), but shorter than other synovigenic, univol- tine females, e.g. Paracantha gentilis Her- ing (Headrick and Goeden 1990), or syno- vigenic, bivoltine females, e.g. Tephritis ar- izonaensis Quisenberry (Goeden et al. 1993). In southern California, the F, gen- eration is produced in flower heads of spring-blooming hosts (Acamptopappus and Amphipappus spp., present study and Goeden 1987); the F, generation utilizes fall-blooming hosts (Chrysothamnus and Haplopappus spp., present study and Goe- den 1987) and overwinters, as noted above, 789 as non-feeding third instars in dried, senes- cent flower heads. The summer is passed as F, adults, which we have observed to ovi- posit first on low-altitude populations of al- ternate hosts such as C. viscidiflorus, a widespread species which successively blooms from June through September at ever higher altitudes (RDG, unpub. data; Munz 1974). Thus, the F, and F, genera- tions of U. timberlakei probably is pro- duced on these long-blooming hosts at dif- ferent elevations. Natural enemies.—Only one parasitoid was reared specifically from U. timberlakei during this study, i.e. a single, solitary, en- doparasitic female Eupelmus sp. (Hyme- noptera: Eupelmidae) from one of four pu- paria dissected from a subsample of 400 flower heads of C. viscidiflorus. This sub- sample also yielded Procecidochares min- uta (Snow). ACKNOWLEDGMENTS Once again, we thank A. C. Saunders, Curator of the Herbarium, Department of Botany and Plant Science, University of California, Riverside, for identification of plants: J. LaSalle, for identification of the parasitoid while he was with the Depart- ment of Entomology, University of Califor- nia, Riverside; and FE L. Blanc and A. L. Norrbom for reviewing a late draft of our manuscript. LITERATURE CITED Blanc, E L. and R. H. Foote. 1961. A new genus and five new species of California Tephritidae. Pan- Pacific Entomologist 37: 73-83. Foote, E L. and FE L. Blanc. 1963. The fruit flies or Tephritidae of California. Bulletin of the Califor- nia Insect Survey 7: 1-117. Foote, R. H., EF L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca. Goeden, R. D. 1985. Host-plant relations of native Trupanea spp. (Diptera: Tephritidae) in southern California. Proceedings of the Entomological So- ciety of Washington 87: 564—571. . 1987. Host-plant relations of native Uropho- ra spp. (Diptera: Tephritidae) in southern Califor- 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON via. Proceedings of the Entomological Society of Washington 89: 269-274. 1992. Analysis of known and new host re- cords for Trupanea from Californica (Diptera: Te- phritidae). Proceedings of the Entomological So- ciety of Washington 94: 107-118. Goeden, R. D. and D. H. Headrick. 1990. Notes on the biological and immature stages of Stenopa af- finis Quisenberry (Diptera: Tephritidae). Proceed- ings of the Entomological Society of Washington 92: 641-648. 199la. Life history and descriptions of im- mature stages of Tephritis baccharis (Coquillett) on Baccharis salicifolia (Ruiz and Pavon) Persoon in southern California (Diptera: Tephritidae). Pan- Pacific Entomologist 67: 86—98. 1991b. Notes on the biology, hosts, and im- mature stages of Tomoplagia cressoni Aczél in southern California (Diptera: Tephritidae). Pro- ceedings of the Entomological Society of Wash- ington 93: 549-558. 1992. Life history and descriptions of im- mature stages of Neaspilota viridescens Quisen- berry (Diptera: Tephritidae) on native Asteraceae in southern California. Proceedings of the Ento- mological Society of Washington 94: 59-77. Goeden, R. D., D. H. Headrick, and J. A. Teerink. 1993. Life history and descriptions of immature stages of Tephritis arizonaensis Quisenberry (Diptera: Tephritidae) on Baccharis sarothroides Gray in southern California. Proceedings of the Entomological Society of Washington 95: 210— 99 1994. Life history and description of imma- ture stages of Procecidochares flavipes Aldrich (Diptera: Tephritidae) on Brickellia spp. in south- ern California. Proceedings of the Entomological Society of Washington 96: 288-300. Green, J. E, D. H. Headrick, and R. D. Goeden. 1993. Life history and description of immature stages of Procecidochares stonei Blanc & Foote on Vigui- era spp. in southern California (Diptera: Tephrit- idae). Pan-Pacific Entomologist 69: 18-32. Harris, P. 1989. The use of Tephritidae for the biolog- ical control of weeds. Biocontrol News and Intor- mation 10: 9-16. Headrick, D. H. and R. D. Goeden. 1990. Description of the immature stages of Paracantha gentilis (Diptera: Tephritidae). Annals of the Entomolog- ical Society of America 83: 220-229. 1991. Life history of Trupanea californica Malloch (Diptera: Tephritidae) on Gnaphalium spp. (Asteraceae) in southern California. Proceed- ings of the Entomological Society of Washington 93: 559-570. 1993. Life history and description of imma- ture stages of Acuirina thoracica (Diptera: Te- phritidae) on Baccharis sarothroides in southern California. Annals of the Entomological Society of America 86(1): 68-79. 1995. Reproductive behavior of California fruit flies and the classification and evolution of Tephritidae (Diptera) mating systems. Studia Dip- terologia 2 (In press.) Headrick, D. H., R. D. Goeden, and J. A. Teerink. 1994. Life history and description of immature stages of Euaresta stigmatica Coquillett (Diptera: Tephritidae) on Ambrosia spp. (Asteraceae) in southern California. Annals of the Entomological Society of America. 87 (In press.) Hickman, J. C. (ed.). 1993. The Jepson Manual. Uni- versity of California Press, Berkeley and Los An- geles. Julien, M. H. 1992. Biological Control of Weeds: A World Catalogue of Agents and their Target Weeds. Third Edition. C.A.B. Internation, Wal- lingford, Oxon, U.K. Munz, P. A. 1968. Supplement to A California Flora. University of California Press, Berkeley. 1974. A Flora of Southern California. Uni- versity of California Press, Berkeley. Munz, P. A. and D. D. Keck. 1959. A California Flora. University of California Press, Berkeley. Novak, J. A. and B. A. Foote. 1968. Biology and immature stages of fruit flies: Paroxyna albiceps (Diptera: Tephritidae). Journal of the Kansas En- tomological Society 41: 108-119. Persson, P. I. 1963. Studies on the biology and larval morphology of some Trypetidae (Diptera). Opus- cula Entomologica 28(1—2): 33-69. Redfern, M. 1968. The natural history of spear thistle- heads. Field Studies 2: 669-717. Steck, G. J. and R. A. Wharton. 1986. Descriptions of immature stages of Eutreta (Diptera: Tephriti- dae). Journal of the Kansas Entomological Society 59(2): 296-302. Steyskal, G. C. 1979. Taxonomic studies on fruit flies of the genus Urophora (Diptera: Tephritidae). Special Publication, Entomological Society of Washington. 61 pp. Varley, G. C. 1937. The life history of some trypetid flies, with descriptions of the early stages. Pro- ceedings of the Royal Entomological Society of London (Ser. A) 12: 109-122. Wadsworth, J. T. 1914. Some observations on the life- history and bionomics of the knapweed gall-fly Urophora solstitialis Linn. Annals of Applied Bi- ology 1(2): 142-169. Zwolfer, H. 1988. Evolutionary and ecological rela- tionships of the insect fauna of thistles. Annual Review of Entomology 33: 103-122. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 791-798 NEW NEARCTIC SPECIES OF ONCOCEPHALUS KLUG (HETEROPTERA: REDUVIIDAE: STENOPODAINAE) J. MALDONADO CAPRILES Department of Crop Protection, University of Puerto Rico, Mayaguez, Puerto Rico 00681; home address: Urb. Aponte 6 I 1, Cayey, Puerto Rico 00736. Abstract.—The three new species Oncocephalus mochis from Sinaloa, Mexico, O. pen- insularis from Lower California, Mexico, and O. variabilis from Arizona, California, and New Mexico are described; and O. nubilus Van Duzee is redescribed. A key to the species occurring in Mexico and United States of America is given. Key Words: tion, United States, key Thanks to Norman D. Penny, California Academy of Science (CAS), the Golden Gate Park, San Francisco, California, I was able to study the specimens of Oncocepha- lus in the collections under his care. Types of the new species herein described are de- posited in CAS. Giacchi (1984) keyed all nine known American and Giacchi and Maldonado (1983) added the new species O. quadrivittatus from Nicaragua. Many re- cords of O. nubilus probably are O. varia- bilis new species. There are about 195 Old World species (Maldonado 1990). At least among the specimens of Onco- cephalus studied the species were found to be very variable. For example, markings on the connexival margins and segments can be absent, or they may be one or two; the postocular margins can be _ parallel-sided and then angularly bent or rounded to col- lum; the relative lengths of the lobes of the pronotum may differ, the annuli on the legs can be dark, pale or absent, the humeral width of the pronotum and the size of the lateral tubercle before the transverse prono- tal constriction may vary. To aid in describ- ing these variations, illustrations of pronota are divided in half, with each half illustrat- Reduviidae, Oncocephalus new species, O. nubilus Van Duzee, redescrip- ing a different color pattern. Extreme mea- surements and color variation for O. iabilis n. sp. are given in parenthesis. The discal cell in Oncocephalus is open caudally, the arrow in Fig. 14 pointing to area where closing vein is lacking. This is a generic character overlooked until now. Ten Old World species examined exhibit this character. The hypopygial caudal pro- cess is located before the caudal margin and on each side, a setose process is found (Fig. 2). The caudal margin of the hypopygium exhibits a different, simple pattern of fine Setae imjeach species (Figs. 35, 11): As in most stenopodaine genera the fore tibia has an apical tuft of sensory hairs and a calamistrumlike organ. The eyes are large, slightly surpassing the lower surface of the head and narrowly apart underneath, more so in females. Depth of femur is its thickness in lateral aspect. S-tubercle stands for setigerous tu- bercle throughout the text. The anteocular space is the distance from the anterior mar- gin of eye to the apex of the antennophore and the postocular space the distance from the posterior margin of eye to the insertion of the collum. All measurements are in mm. var- 192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON KI O THE SPECIES OF ONCOCEPHALUS IN MEXICO AND NORTH AMERICA Anterior lobe of pronotum with a lateral tuber- cle before transverse constriction; males with pilosity of first antennite shorter than diameter of segment; discal cell of corium and outer cell of membrane with black patches .......... 2 Anterior lobe of pronotum without such tuber- cle; males with pilosity of first antennite 2X or 3 diameter of segment; discal and outer cell of membrane without patches . Second antennite about 2.5 times longer than first, the latter twice as long as anteocular mar- gin; posterior tibia with dense long setae, setae 1.5—2 times as long as diameter of tibia. Color gray cinereus O. geniculatus Stal Second antennite nearly 3 times as long as first, the latter 2/5 longer than anteocular margin; posterior tibia with dense short setae, setae not much longer than diameter of tibia. Color more Strain OUStesiesek a ene O. apiculatus Reuter . Head, pronotum, and hemelytra except later- ally, black, connexivum reddish dorsally; an- tennite II about 2.5 times as long as first. Baja California O. erectus Van Duzee Head, pronotum and hemelytra brownish, stra- mineous or yellowish; connexivum not reddish, more or less stramineous with or without blackish ornamentation; antennite II variable Anterior lobe of pronotum with median stripe 3 as wide as median sulcus (Fig. 4); posterior lobe mostly brown, with paramedial carinae, | + 1 small spots lateral to carinae, and humeral angles stramineous; hemelytra uniformly pale brown. Sonora, Mexico O. mochis n. sp. Anterior lobe of pronotum with median stripe narrow, not much broader than median sulcus; posterior lobe of pronotum less extensively brown, lateral margins stramineous; hemelytra differently colored, paler or clearly bicolored . Hemelytra bicolored (Fig. 7), laterally to pter- ostigma and outer longitudinal half of corium yellowish-stramineous, contrasting with the darker scutellum, inner longitudinal half of cla- vus and corium, and membrane; the yellowish area of the hemelytra continuous with the sim- ilarly colored lateral areas of pronotum Se OAs Se eon O. nubilus Van Duzee Hemelytra without such contrasting areas, al- most uniformly colored; the yellow areas of the pronotal margins not extending into hemelytra . Posterior lobe of pronotum with narrow para- medial longitudinal fasciae connected basally by a narrow transverse band, thus forming a thin U; corial veins usually set on stramineous stripes resulting in darker discal areas mar- gined with stramineous; short section of vein R + M at level of discal corial cell brown, of same color as connexival marginal markings. Baja California O. peninsularis n. sp. — Posterior lobe of pronotum with brown fasciae and basal band broad, not forming a thin U; corial cells uniformly colored; hemelytra and vein R + M concolorous. Southwestern United States O. variabilis n. sp. DESCRIPTIONS Oncocephalus mochis Maldonado, NEW SPECIES Figs. 1-4 Male.—Head: anterior lobe stramineous above, with a brownish longitudinal stripe each side of median line, laterally with a longitudinal brown stripe from anterior margin of eye to apex of head; posterior lobe dark brown above, brown below; ocel- lar callus black. I antennite brownish, base stramineous, pilosity golden, internal setae about twice diameter of segment; II seg- ment brownish, slightly paler than I, a long preapical stramineous band, long fine pi- lose, pilosity 3 times diameter of segment, gradually decreasing in length toward apex, more abundant on inner half of perimeter; III and IV segments dark gray, pilosity scarce, about twice as long as diameter of segment. Rostrum stramineous, last seg- ment darker toward apex. Pronotum (Fig. 4): anterior lobe mostly stramineous dor- sally, anterolateral angle blackish brown laterally, a relatively broad median longi- tudinal stripe, | + 1 paramedial brownish stripes; posterior lobe dorsally mostly brown, humeral angles, | + 1 small anterior spots, and carinae stramineous. Pronotum laterally, pleurae, coxae, trochanters, and thoracic sterna stramineous. Femora stra- mineous, profemur and metafemur with in- complete annulus at midlength, preapically brown; mesofemur with a preapical brown band; metatibia with short basal and middle annuli. Scutellum blackish brown, apical spine stramineous. Hemelytra pale brown, VOLUME 97, NUMBER 4 793 -———3 —} ———4 ————} —— I | 10> == / y (an / pS i ee 11 Figs. 1-12. Oncocephalus mochis Maldonado n. sp., male holotype. 1, clasper, dorsal view. 2, caudal spine and dorsolateral process (dlp) of hypopgyium, caudal view. 3, dorsocaudal margin of hypopygium, caudal view. 4, pronotum (two color variations), dorsal. Oncocephalus nubilus Van Duzee, male holotype. 5, head, dorsal view and detail of surface. 6, head, lateral view. 7, coloration of body, dorsal view. 8, clasper, dorsal view. 9, caudal spine and dlp, caudal view and lateral detail of dlp. 10, profemur, lateral view. 11, dorsocaudal margin of hypopygium. 12, female, VIII tergum. membranal cells with scattered stramineous _ spiracle, all three arranged in a zig-zag pat- spots. Connexival margin with or without a tern. White scale-like setae each side of ty- long and a short brown spot; abdominal lus, above metapleura, and suture before sterna yellowish stramineous, each with first abdominal sternum. Head: length to two brownish spots caudad to the brownish apex of collum 2.3, width across eyes 1.85, 794 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘reatest width of anteocular lobe 1.0, width of postocular lobe behind eyes 1.3, width f eye 0.5, height of eye 0.97, length of eye 0.71, interocular space 0.8, anteocular mar- gin 0.65; distance between eyes underneath 0.2, width of ocellar callus 0.75, length of collum 0.3. Length of antennal segments: I, 2.25% Il, -a alee 226: SNe 2 SA sethkee (S= tubercles lateroventrally behind eyes, the rest of head with minute globose s-tuber- cles. Pronotum: width across anterolateral angles 1.44, width across humeral angles 3.25, width at transverse constriction 2.05, length of anterior lobe 1.54, length of pos- terior lobe 1.40, anterior lobe with very small s-tubercles in rows and along lateral margins; median sulcus on apical third only; surface of head as in O. nubilus (Fig. 5); posterior margin slightly concave above scutellum. Scutellum: basal width 1.2, length to base of apical spine 1.0, length of spine 0.5, with scarce granulations and small s-tubercles. Hemelytra: M + Cu vein with or without small s-tubercles; not or slightly surpassing apex of abdomen; length 9.75, greatest width 3.38. Fore leg, lengths: coxa 1.0, trochanter 0.9, femur 4.4; depth of femur 1.16; armature consisting of 10 conical teeth alternating with 3 to 5 small s-tubercles; tibia length 3.9, setae decum- bent and slightly shorter than diameter of segment; tarsi: first two together slightly longer than third, 6.5:5, claws expanded ba- sally, curved. Middle leg, lengths: coxa 0.75, trochanter 1.0, femur slightly thicker apically, 4.25, tibia 4.62; first two tarsal segments combined as long as third, 0.65; claws and pilosity as in fore legs. Hind legs: coxa 1.24, trochanter 0.95; femur 6.90, slightly expanded apically; tibia 7.8, thinner toward apex, consequently pilosity after midlength longer than diameter of segment; first two tarsal segments together longer than third, 0.8:4.5; claws as in fore legs, less curved. External genitalia as in Figs. 1—3. Total length of body 14-15 mm. Holotype male.—Sinaloa, Los Mochis, MEXICO, vii-20-1922, C. T. Dodds collec- tor; 5 paratypes; all from the type locality, July 1922, C. T. Dodds collector. One in JMC, others in CAS. The trivial name refers to the type local- ity. The almost uniform pale brown color of the hemelytra makes this species stand out from nearly all of its congeners because of their mostly stramineous or clearly bi- colored fore wings as in O. nubilus. Only O. erectus Van Duzee, from Baja Califor- nia, has darker, but almost black hemelytra. Oncocephalus nubilus Van Duzee Figs. 5-12 Redescription of male holotype.—Head (Figs. 5, 6): Anterior lobe above strami- neous, laterally brownish or dark brown, ventrally dark brown with a transverse fig- ure-8 yellow spot midway from bucculae to eyes; posterior lobe, neck, and underneath between eyes brown. Ocellar elevation brown, black between ocelli. Eyes black. White scale-like setae around tylus and an- tennophore, in front and around eyes. An- tenna: antennite I brown, paler below; II slightly paler brown than I laterally, stra- mineous internally, apex grayish brown; III grayish brown; IV missing. Rostrum stra- mineous, apex brown. Pronotum—anterior lobe: median longitudinal stripe brown; area on either side of median stripe stra- mineous on basal half, posterior half almost orange; disc of posterior lobe blackish brown, marginal areas stramineous (Fig. 7); mesopleura pale brown; metapleura brown. Mesosternum medianly with abundant, thick, decumbent whitish setae, narrowly margined with brown. Metasternum brown, margined laterally and anteriorly with a band of whitish setae. Scutellum blackish brown, from disc to apex of spine yellow. Legs—anterior: coxa and trochanter yel- lowish; femur with basal third stramineous, middle third pale brown with subcircular yellowish areas ventrally and with more ex- tensive yellow areas dorsally, apical third brown with yellow apical margin; tibia yel- lowish, postbasal and about midlength brown annuli; tarsi yellowish; middle: coxa and trochanter yellowish, apical third VOLUME 97, NUMBER 4 brown; hind: coxa brownish, trochanter stramineous, femur with a median and api- cal long brown annulus; tibia with basal and postbasal brown annuli. Hemelytra: base of clavus and corium yellowish, rest of wing brownish (Fig. 7). Abdomen brownish each side of ventral keel, reddish brown below line of spiracles, both bands of about same width; connexivum variable, usually yel- low, margin with a central black, narrow marking. Head: Length from interocular sulcus to apex 1.9, width across eyes 1.90, width of eye 0.55, length of eye 0.7, height of eye 1.10, interocular space 0.8, anteocular space 0.7, postocular space 0.6, width behind eyes 1.25; minutely granulose (Fig. 5); lateral margins of postocular lobe rounded to col- lum. Antennal segments: I, 2.2; II, 4.51; III and IV missing; I with abundant long, fine setae on ventral surface, short decumbent setose dorsally; II with similar pilosity, se- tae gradually decreasing in length toward apex; III long pilose, hairs twice as long as thickness of segment. Rostral segments: I and I, 0.9; III, 0.6. Pronotum with mod- erately abundant small granules; length 3.5, width at anterior angles 1.6, width at hu- meral angles 3.94, length of anterior lobe 1.62; two inconspicuous carinae extending from anterior lobe to basal third of posterior lobe; posterior margin shallowly concave above scutellum. Scutellum: width at base 1.08, length to base of apical spine 1.4, length of spine 0.5, spine slight upcurved. Fore leg: coxa 0.5, trochanter 0.9, two small spines on ventrolateral edge; femur (Fig. 10): length 4.7, greatest depth at about mid- length, 1.12, armature usually consisting of 10 short teeth alternating with 3 to 5 small s-tubercles; tibia length 4.70, slightly thin- ner apically; tarsi 0.4; 0.4; 0.9; pilosity de- cumbent and shorter than corresponding segment, except on tarsi where it is slightly longer than their diameter. Middle leg: coxa 0.70; trochanter 0.90; femur 5.00, slightly thicker at midlength, 0.4; tibia 4.81 long; tarsi 0.25; 0.30; 0.60; pilosity as in fore leg. Hind leg: coxa 1.10; trochanter 1.10; femur 795 7.82, straight; tibia 9.2; tarsi 0.4; 0.4; 0.8; claws of middle and hind legs less thick- ened basally than on front leg; pilosity of femur and tarsi as in first two legs, setae on tibia longer than thickness of segment. He- melytra almost reaching apex of abdomen. Abdomen keeled to base of VI sternum. Ex- ternal genitalia as in Figs. 8, 9, 11. Total length of body 17.5 mm. Females are very similar in morphometry and_ coloration; VIII tergum as in Fig. 12; overall length of body from 14 to 18 mm. Holotype.—San Bernardino Ranch, Co- chise Cty, ARIZONA, FE. H. Snow collector, in Van Duzee collection at the CAS, San Francisco, California. Reported by Henry and Froeschner (1988) from Texas, Arizo- na, and California. Also known from north- ern Mexico. Oncocephalus peninsularis Maldonado, NEW SPECIES Figs. 13-17 Male.—Mostly stramineous. Head: ver- tex with short brown dash each side of me- dian line; laterally with poorly defined brown line not quite reaching apex of head, beneath blackish brown; posterior lobe dark brown above and laterally, below strami- neous; ocellar callus black. Antenna: I an- tennite brown, basally stramineous, II brownish basally gradually deepening to brown apically, II] and [V brown. Prono- tum (Fig. 17): disc of posterior lobe with a median, relatively broad brownish stripe and a narrower stripe on each side, the me- dian extending into anterior lobe as a nar- row stripe. Scutellum dark brown, medianly from base or postbasally to apex of spine with a tapering yellowish-white stripe. Ros- trum: first segment mostly yellowish, sec- ond stramineous or brownish above, third brown. Coxae and trochanters yellowish. Femora yellowish: anterior—with long api- cal brown annulus, annulus with irregular small yellowish areas; middle—with apical brown annulus; hind—with apical and mid- length brownish annuli. Tibiae yellowish; anterior—three-annulate, the last two fused, 796 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON exw ee See es a» Figs. 13-23. Oncocephalus peninsularis Maldonado n. sp., male holotype. 13, clasper, dorsal view. 14, hemelytron. 15, caudal spine and dlp. 16, dorsocaudal margin of hypopygium. 17, pronotum and scutellum, dorsal view. Oncocephalus variabilis Maldonado n. sp., male holotype. 18, clasper, dorsal view. 19, apex of clasper, internal view. 20, caudal spine and dlp, caudal view. 21, pronotum, dorsal. 22, profemur, lateral. 23, VIII tergum of female, dorsal. faint or incomplete; middle—similar to an- terior; posterior—with apical and preapical annuli. Tarsi yellowish. Hemelytra (Fig. 14): corial veins set on stramineous stripes, thus cells with pale brown discal areas; R + M yellowish, section laterad of discal cell dark brown. Connexival segments un- marked, marked basally or with dark brown margin. Head.—Length from interocular sulcus to apex of head 1.35, width across eyes 1.35, width of eye 0.54, length of eye 0.65, anteocular space 0.32, postocular space 0.51, width behind eyes 1.1, anterior mar- VOLUME 97, NUMBER 4 gin of ocellar callus at about midlength of eye, ocellar callus as wide as interocular space. Antenna: I, 2.0; II, 4.4; III, 1.13; IV, 1.21; I abundantly long pilose on outer half of perimeter, II long pilose, setae gradually shorter towards apex; III and IV scarce pi- lose, setae about twice as long as segment. Head—surface as in nubilus (Fig. 5); con- spicuous s-tubercle lateral on antenniferous tubercle, 2—3 s-tubercles ventrolaterally be- hind eyes. Pronotum (Fig. 17): length 2.75, anterior width 1.24, humeral width 3.42, width at constriction 1.45, length of anterior lobe 1.34, carinae across constriction ob- solete, anterior lobe with scarce small gran- ulations; posterior lobe with abundant small granulations and very small globose s-tu- bercles; posterior margin shallowly concave above scutellum; anterolateral and humeral angles sharp but not produced. Scutellum: width 1.4, length to base of spine 1.2, length of spine 0.4. Hemelytra: length 10.5, greatest width 3.36. Abdominal sterna keeled to midlength of sixth. Fore leg: coxa length 0.6, trochanter length 0.9; femur— with scarce and short s-tubercles; length 4.2, depth 1.0; armature: 9 teeth, each one alternating with one small s-tubercle; tibia slightly thinner preapically, setae on side facing femur shorter than diameter of seg- ment, length 4.1; tarsi: first two segments together shorter than third (0.51:0.65); claws as in nubilus. Middle leg—coxa length 0.3, trochanter length 1.0; femur thickness at midlength 0.32, slightly nar- rower preapically; tibia length 3.4. Hind leg: coxa length 0.6, trochanter length 0.75; femur decumbent setose; tibia length 7.2, long setose on apical 4/5. Genitalia as in Figs 13, 15, 16. Total length of body 15.0 mm. Holotype male.—15 mi. N, El Refugio, LOWER CALIFORNIA, vii-1938, Mich- elbacher and Ross colls., in CAS. Para- types—4, same data as holotype; | in JMC others in CAS. The trivial name makes reference to the peninsula that characterizes Lower Califor- nia. The key characters identify this spe- TST. cies; the blackish marking along R + M vein or inner vein of embolium is visible to the naked eye. Oncocephalus variabilis Maldonado, NEW SPECIES Figs. 18—23 Male.—Overall color stramineous; head, pronotum, first pair of legs darker than he- melytra and last two pairs of legs; orna- mented moderately with shades of brown. Head: postocular space brownish, before and after eyes brownish laterally; gular re- gion brown; I and II antennite pale brown, same as anteocular region, the latter with short apical annulus, III and IV gray. Ros- trum: segments I and II slightly paler than I antennite, III brown. Eyes and ocellar cal- lus blackish brown. Pronotum (Fig. 21) slightly paler than head; anterior lobe—lat- eral margin dark brown, median longitudi- nal stripe and short extensions of fasciae of posterior lobe brown; posterior lobe with median subtriangular area, paramedial fas- ciae brown and common base brown (para- types of both sexes show same pattern but with much wider paramedial fasciae, no two exactly alike). Meso- and metapleura with slightly darker anterior areas (these darker areas present in two paratypes only). Coxae and trochanters stramineous; femora brown apically, on anterior femora not forming an annulus; tibiae—anterior with basal, postbasal, and apical ill-defined an- nuli (traces of these annuli in most para- types); last two femora with well defined apical annulus only; tarsi quite dark, an ev- ident abnormality as these are stramineous in all other specimens. Head: Length 2.25 (2.19-2.44), width across eyes 1.69 (1.50), interocular space 0.81 (0.68), anteocular space 1.00, postoc- ular space 0.37; pronotum—anterior lobe length 1.56 (1.32—1.69), posterior lobe length 1.31 (1.25-1.50), humeral width 3.18 (3.00—3.25). Antennal segments—I, 137°C. 3) ty 3.87612); MR1V. thinner and shorter than first, too twisted for accu- rate measurements. Rostrum I, 1.00; II, 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON [IIl, 0.75; first not reaching anterior rzin of eye. Scutellum as wide as long 8 (1.12—1.25). Legs—length of femora 1.12 (3.74—3.87), 4.37, 6.75; depth of pro- femur 1.12 (Fig. 22), armature consisting 8 or 9 small tooth-like spines each alternating with 3—4 small hemispherical s-tubercles; tibiae -3175; "4:00" 72003 ‘tarsi 1-757) 100; 1.37. Fore wing length 9.38, not quite reaching apex of abdomen, width 3.50. Genitalia as in Figs. 18—20. Overall length 14.7 (13-16 mm). Female.—Overall coloration as in male. Dimensions fall inside the extremes given for the males. Two specimens are 17.0 long. VIII tergum as in Fig. 23. Holotype.—Male, S. Luis Yuma, ARI- ZONA, viii.11.40, presented by E. C. Van Dyke, in CAS. Paratypes 27, both sexes; ARIZONA—4, same data as holotype; 4, Ehrenberg, vii.20.1946, E. C. Van Dyke collector; CALIFORNIA—1, Ft. Yuma, Aug.21.1924, E. PR Van Duzee collector; 8, Coachella, yol7.1928, B.C. Van Dyke col- lector; 2;. Blythe, .vii1.3.40, J. W.. Tilden coll-:32;, Blythe: v.12.37, Ee. PVansDuzee coll cone. Pine Inyo.€o., Diaz, Lake, vi. 16:37, EsC) Van Dykeicoll:ai Browleys v.1940, R. P. Allen coll.; NEW MEXICO— Carlsbad, Carera, 111.38, R. P. Allen coll; 2 from S. L. Yuma in JMC, others in CAS. Discussion.—The trivial name variabilis refers to the very variable morphometry of the species. The overall coloration and the first rostral segment not reaching the ante- rior margin of eye are constant characters. LITERATURE CITED Barber, H. G. 1930. Essay on the Subfamily Steno- podinae of the New World. Entomologica Amer- icana, New Series 10: 149-238. Giacchi, J. C. 1984. Revision de los stenopodainos americanos. VI. Las especies americanas del ge- nero Oncocephalus Klug, 1830 (Heteroptera-Re- duviidae). Physis Seccion C 42(103): 39-42. Giacchi, J. C. and J. Maldonado Capriles. 1983. A new species of Oncocephalus Klug, 1830 (Redu- viidae: Stenopodainae) from Nicaragua. Caribbe- an Journal of Science 19(3-4): 61-63. Henry, T. J. and R. C. Froeschner. 1988. Catalog of the Heteroptera, or true bugs, of Canada and Con- tinental United States. J. E Brill, New York. 958 Pp. Maldonado Capriles, J. 1990. Systematic Catalogue of the Reduviidae of the World (Insecta: Heter- optera). Special Publication of the Caribbean Jour- nal of Science, University of Puerto Rico, College of Arts and Sciences, Mayagiiez, Puerto Rico 00681. x + 694 pp. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 799-832 COMPARATIVE STUDY OF MOUTHPARTS OF PREDACEOUS MIDGES OF THE TRIBE PALPOMYIINI (DIPTERA: CERATOPOGONIDAE) FROM EASTERN UNITED STATES STURGIS MCKEEVER, DANIEL V. HAGAN, AND WILLIAM L. GROGAN, JR. (SM) Professor Emeritus and (DVH) Professor, Department of Biology, Institute of Arthropodology and Parasitology, Georgia Southern University, Statesboro, Georgia 30460-8042; (WLG) Professor, Department of Biological Sciences, Salisbury State Uni- versity, Salisbury, Maryland 21801-6837. Abstract.—Mouthparts of females of 16 species and males of 13 species representing Bezzia, Phaenobezzia, and Palpomyia of the insectivorous tribe Palpomyiini were ex- amined by light and scanning electron microscopy. Greatest differences among species occurred in the sclerotized mandibles of females where the mean number of coarse medial teeth ranged from 6.00 to 10.31, small basal teeth from 0.37 to 3.47, length of the tooth row in relation to length of the mandible from 14.9 to 29.8% and mean number of lateral teeth from 1.50 to 6.61. An interlocking mechanism midlength of the blade functions as a fulcrum; abduction of the base of the mandibles results in divergence of their tips and enables their strong teeth to cut the cuticle of the prey. The non-insectivorous males have mandibles with large medial non-functional teeth and lack an interlocking mechanism. Lateral teeth are present on the mandibles of some males of eight species. The unarmored labrum, hypopharynx and lacinia are inserted into the wound made by the mandibles. The mandibles remain between the labrum and hypopharynx during feed- ing and serve as the floor of the food canal and roof of the salivary canal. The laciniae, which are smooth in both sexes, form the lateral proximal boundaries of the food canal. Cylindrical structures at the tip of the labrum of females are thought to be mechanore- ceptors, chemoreceptors or both; they are reduced in size or absent in males and, in most species, are replaced by spicules. Labra of males and females have species specific char- acters, but differences are less pronounced in males. The hypopharynx of females is generically specific in its shape and species specific in distribution of its spicules. The hypopharynx of males is species specific in size and shape of its spicules which are longer than in those of conspecific females. Sensilla on the maxillary palp sensory organ are species specific in both sexes with respect to number, points of origin on the palp of individual sensilla, shape of the head of the sensilla and total length. The number of sensilla per sensory organ is approximately equal in both sexes of a given species and they probably do not function in prey detection and/or location, which is thought to be more related to visual stimuli in predaceous midges. Key Words: Insecta, Ceratopogonini, Palpomyiini, mouthparts The Palpomyiini Enderlein, one of the Clastrieromyia Spinelli and Grogan, Pach- insectivorous tribes of Ceratopogonidae yhelea (Wirth), Palpomyia Meigen, and Newman is comprised of the genera Ame- Phaenobezzia Haeselbarth (Spinelli and rohelea Grogan and Wirth, Bezzia Kieffer, Grogan 1985). Females of this tribe, as well 800 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON as those of Heteromyiini Wirth, and Sphae- siini Newman, and most Ceratopogonini ewman feed on Chironomidae, Chaobor- idae and Ephemeroptera (mayflies), usually males, that are captured from mating swarms. In addition, in Palpomyiini, Het- eromyiini and Sphaeromiini, when the fe- male enters mating swarms of her own spe- cies, she captures a male that she feeds upon while he is in copula with her (Downes 1977). Females of the tribe Sten- oxenini Coquillett also have well developed mandibles with large teeth, thus indicating that they too are insectivorous; however, observations on their swarming, mating or feeding habits are lacking (Wirth and Ra- tanaworabhan 1972). Mouthparts of both males and females of an insectivorous species of Ceratopogoni- dae, Probezzia (as Dicrobezzia) venusta (Meigen), a member of the Sphaeromiini were described in detail by Gad (1951). All structures were present in both sexes, with male structures being smaller and less de- veloped. Gad also presented data and draw- ings of some structures of other insectivo- rous genera, including Bezzia and Palpo- myia, and compared mouthparts of insectiv- orous forms with hematophagous and flower-feeding representatives of the fami- ly. Downes illustrated and briefly described mouthparts of Probezzia flavonigra Coquil- lett (1971) and Palpomyia quadrispinosa Goetgebuer (1978). Mouthparts of insectiv- orous Ceratopogoninae are characterized by large mandibular teeth and reduced, tooth- less laciniae, an exception being members of the insectivorous anophelis group of Culicoides Latreille subgenus Trithecoides Wirth and Hubert that have large mandib- ular teeth and toothed laciniae (Wirth and Hubert 1989). Hematophagous forms, e.g. Culicoides have small, closely set mandib- ular teeth and toothed laciniae. Gad found the tips of the mandibles, labrum, laciniae, and hypopharynx to be weakly sclerotized and without functional teeth in Atrichopo- gon pavidus (Winnertz), subfamily Forci- pomylinae; therefore, he considered the species to be a nectar feeder. Many studies have used mouthpart struc- ture to distinguish taxa. For example, Wirth and Hubert (1959) used the structures of el- ements of the proboscis of the subgenus Trithecoides (tribe Culicoidini), for identi- fication of species and species groups. Glu- khova (1982) found that each element of the proboscis of 13 species of Trithecoides exhibited considerable species diversity, in contrast to a high degree of similarity of other structures (wing, maxillary palpus and spermathecae), and used diversity of these elements as diagnostic characters. She found no species-specific differences in the maxillary palpus of species of Trithecoides, but Chaika (1978) found the palpal sensory organ to be of taxonomic significance in some other species of Culicoides. Endopar- asitic species of Atrichopogon Kieff. (sub- family Forcipomyiinae), were divided into four groups by Glukhova (1981) based upon structure of the female mouthparts. She related this diversity to differences in the various insect hosts (beetles, lacewings, dragonflies, etc.) and the parts of their body upon which these ectoparasitic species feed. She found even greater diversity in mouth- parts of species of the related genus For- cipomyia which she attributed to diversity of their hosts. Wirth and Grogan (1988) in- cluded mandibles and palpi to partially characterize genera of Ceratopogonini. In a recent comparative study of ten species rep- resenting seven genera of predaceous midges of the tribe Ceratopogonini, we found the mandibles, labrum and _palpal sensory organs of females to be species specific and the hypopharynx generically specific (McKeever et al. 1991). Descriptions of mouthparts of the pre- daceous midge tribe Palpomyiini (subfam- ily Ceratopogoninae) are few and generally confined to the mandibles. Dow and Turner (1976) presented data on the number of mandibular teeth of female Bezzia along with drawings of female mandibles of five of their 40 Nearctic species. Wirth (1983a) VOLUME 97, NUMBER 4 presented data on the number of mandibular teeth of two species of Bezzia with a draw- ing of the mandible of one. In a later paper, Wirth (1983b) presented drawings, accom- panied by data on the number of teeth, for mandibles of seven species of Bezzia fe- males, as well as a drawing of the male mandible of one species. Wirth and Grogan (1983) gave data on the number of mandib- ular teeth of all species of Bezzia. Grogan and Wirth (1981) presented data on the number of mandibular teeth of 10 species of Amerohelea accompanied by drawings of the mandibles of two of the species; all of these species occur in Central and South America, and one is found in southwestern United States as well. Grogan and Wirth (1979) gave the number of mandibular teeth of female Palpomyia of North America and presented drawings of the mandibles of fe- males of nine of thirty-one species and males of three species. Downes (1978) pre- sented photomicrographs of mouthparts of female Palpomyia quadrispinosa Goetghe- buer and discussed in detail their functions. Wirth and Grogan (1982) presented a draw- ing of the female mandible of Phaenobezzia opaca (Loew), and Gad (1951) illustrated the female mandible of Palpomyia luteife- morata Edwards. To date, no studies of mouthparts of predaceous Palpomyiini have employed electron microscopy. This is a re- port of a detailed comparative study, by light and scanning electron microscopy (SEM), of the morphology of mouthparts of females of 16 species and males of 13 spe- cies in three genera of Palpomyiini to de- termine whether generic differences or spe- cies-specific differences, or both, exist in any or all structures of the mouthparts. MATERIALS AND METHODS Females of the following species were examined: Bezzia bivittata (Coquillett), B. imbifida Dow and Turner, B. nobilis (Win- nertz), B. glabra (Coquillett), B. dorsase- tula Dow and Turner, Phaenobezzia opaca, Palpomyia subaspera (Coquillett), P. cres- soni (Malloch), P. lineata (Meigen), P. ple- 801 beia (Loew), P. rufa (Loew), P. pseudorufa Grogan and Wirth, P. basalis (Walker), P. flaviceps (Johannsen), P. scalpellifera Gro- gan and Wirth, and P. hastata Grogan and Wirth. Males of all the foregoing species except P. cressoni, P. flaviceps and P. has- tata were examined. All specimens except 13 P. nobilis that were collected at Hatchet Creek, Alachua County, Florida, were collected on Patuxent Wildlife Research Center and Beltsville Ag- ricultural Research Station, both near Lau- rel, Prince Georges County, Maryland or at Salisbury, Wicomico County, Maryland. Specimens were collected with Malaise traps or by sweeping vegetation and stored in 70% ethanol. Later, they were dehydrated in a graded ethanol series, dried in a critical point drier and dissected with tungsten nee- dles sharpened in boiling potassium nitrate. Dissected structures were suspended hori- zontally from fresh micro-drops of CMC mounting medium (Turtox, Chicago) on 15 mm glass coverslips that had previously been coated with a thin film of CMC. Spec- imens were examined with a light micro- scope, measured, then gold-coated and later photographed with an ISI Super II scanning electron microscope. Measurements of 13 characters of the various mouthparts of 127 females and 77 males were made with an ocular microme- ter at 400 and values expressed in mi- crometers (jm). In addition, the number of mandibular teeth was determined at 400. The number of spicules on the labrum and hypopharynx and the number of palpal sen- sillae were counted directly on the SEM screen. Length of the sensilla and camber and pitch of the mandibular teeth were de- termined from SEM photomicrographs. Statistical differences were determined by t-tests. Measurements were determined as in our previous study (McKeever et al. 1991) as follows. Mandibles: (total length)—from attach- ment of adductor muscle at outer angle of base of mandible to tip of distal tooth; (length of tooth row)—from proximal base Table 1. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Measurements (um) of mandibles of female Palpomyiini. Total Length Maximum Width Width at Basal Tooth Length of Tooth Row Species Mean SE n Mean SE n Mean SE n Mean SE n B. bivittata 142.10 13.46 13 SYD ARS 118} L821 4°33) 13331045 OSes B. imbifida 170.00 6.56 11 S70 Ae6s: 12 yin A/S NW EOI BBO" 1D B. nobilis LG837i/ ay S32 B84 BXOO)) ANBAR S37 1944) 135) 39) 50129) 3259s B. glabra AD SOD Oe SAS). » P2eThs3s PX) 1958) 54 25 4926 F596 B. dorsasetula 126.34 6.36 33 226, OT AAS5 12-94 OF49) 55335) 29388 2S Ph. opaca 147.06 5.48 37 34:20) 1:45 35 abs) OTs) SI) 27/808) 1505 37 P. subaspera 232.08 — 2 43.22 — 2 25.93 — 2 64.50 — 2 P. cressont 186258" 3:07 fl 45.88 2.62 10 5:56) 127 10 3 8:047° 3:62 5 a0 P. lineata 206:835 15:67 ~ 38 43.74 3.56 s) 18.62 3.54 2 Spi) 8.52 y P. plebeia 15820) 5 3.015 9 30.47 0.81 22 [ICO NOS C225 33:67 OSS P. rufa 205.66 6.99 34 39'045 ll-55\5" 34 LSU 2OTAS5 9437 0245S P. pseudorufa 159.41 3.48 28 28:60); 20:94 29 10:92, Ae 285) 37200 1.90 8 P. basalis 152295) S110 4 Sp) PUA = Sia 4 14.30 3.16 4 28.26 20.30 4 P. flaviceps AD OO SZ ane2 32222, WEAN 24 [4226)) hI 25,5 30:08 1-835 26 P. scalpellifera WIS 230 22:05 8 36.41 4.48 8 13296) 059 8 25577 1.45 8 P. hastata 146.96 — 2 3321 — 2 12.46 — 2 24.60 — Dy of proximal tooth to tip of distal tooth; (maximum width)—from medial to lateral edges, near midpoint, measured at 90° to long axis of mandible; (width of basal tooth)—from proximal base of proximal tooth to lateral edges, measured at 90° to long axis of mandible; (length of longest tooth)—from tip to base; (camber of the teeth)—proximal angle between distal edge of the fifth tooth (line from tip to distal base) and a line parallel to base of the tooth row; (pitch of teeth)—proximal angle be- tween proximal edge of the fifth tooth (line from tip to proximal base) and a line par- allel to base of the tooth row. Labrum: (length)—from junction with hypopharynx to tip; (width)—maximum immediately proximal to teeth. Hypopharynx: (length)—from junction with labrum to tip; (width)—maximum im- mediately proximal to teeth. Laciniae: (length)—from junction with maxillary palpus to tip; (greatest width)— distance between lateral edges at junction with maxillary palpus, measured at 90° to long axis of the laciniae. Sensilla: (length)—from origin in socket to apex. There are no voucher specimens, because the specimens were destroyed in preparing the mouthparts for examination. Specimens of all species described herein are in the Florida State Collection of Arthropods, Gainesville, the U.S. National Museum of Natural History, Washington, D.C., and the Canadian National Collection of Insects, Ottawa. The mandible, labrum, hypophar- ynx, lacinia, and palpal sensillae are illus- trated by SEM photomicrographs (Figs. 1— 17). Size relationships of the structures are presented tabularly, as the mean and two standard errors. (Tables 1—8). RESULTS General Descriptions of Structures Mandible.—The strongly sclerotized mandibles of females have an interlocking mechanism approximately midlength of the blade that is similar in structure and func- tion to that described for Culicoides (Gad 1951) and the Ceratopogonini (McKeever et al. 1991). On the medial edge the distal first and usually second antrorse teeth are followed by a series of longer, coarse, re- trorse teeth that may be uniform throughout or may decrease in length and size proxi- mally (Figs. 1-5). There were 6—17 such VOLUME 97, NUMBER 4 80 WwW Table 2. Measurements (1m) and number of teeth of mandibles of female Palpomyiini. Length of Longest Tooth No. of Coarse Medial Teeth No. of Small Basal Teeth No. of Lateral Teeth Species weMicane PaSEL o ova Mean SE n Mean SE n TiMeanl SE nn B. bivittata Salve Or4 13 10.31 1252 19 3.47 1.40 19 3.20 1.00 20 B. imbifida QT 0.12 12 9.42 0.99 1 2.17 0.59 12 35) O61 12 B. nobilis 4.4] 0.27 39 8.96 0.25 45 335) 3059" ~45 444 0.47 45 B. glabra SOE OMA] 26 10.11 es 7/ less OSS) 227/ S00, 0:89) 27 B. dorsasetula 22009 33 7.71 Oy Sil 2.58 054 31 3.64 055 33 Ph. opaca 6:27, 10338 39 OT ONO 45 0.89 0.30 45 6.61 3.36 44 P. subaspera 7.98 — 2 9.50 — 2, 2.00 _ 2 3.50 = 2 P. cressoni 5-59) 0l60" 39 TAO! O63 10 2.60 0.69 10 4.09 1.04 10 P. lineata 5.91 0.90 9 8229 0M,9 14 4.71 0.83 i4 6.57 0.43 14 P. plebeia »DOn OSs 22 M05—) O37 2 DegXsy — (OS10) PI 4.54 045 22 P. rufa 5.69 O28) 34 7:10 O15 41 3227 OT al 39/8) 1058) 741 P. pseudorufa 454 0.32 28 697 (0NBy) 35 2208 0338" 35 3:09) 305455535 P. basalis 3.99 0.00 4 6.00 oo 8 0.37 =—0..89 8 3H) 1059 8 P. flaviceps 3109) 033") 26 (06 O34 33 OG 0435 3s 2227, 1 0'O9 33 P. scalpellifera 4.40 0.66 8 6.25 0.43 8 1.87 0.94 8 5.29 1.38 7 P. hastata 2.66 — 2, 6.00 = 2 1.50 - 2 1:50 = 2 teeth on the distal 14.9 to 29.8% of the blade in all 16 species examined. Proxi- mally, up to eight small basal teeth may be present. Up to 11 small, antrorse projec- tions, considered to be accessory teeth, may occur on the lateral edge in all species, but these may be absent in some specimens. Greatest width of the mandible ranges from 18-24% of the total length among the 16 species and is attained at the point of the interlocking mechanism; little or no de- crease occurs proximally. The base curves approximately 45—90° to the long axis of the blade. Muscle attachment and function are as described for the Ceratopogonini (McKeever et al. 1991). Mandibles of males of the 13 species ex- amined are poorly sclerotized, broad blades Table 3. Measurements (4m) and number of spicules of labra and hypopharynx of female Palpomyltini. Labra Hypopharynx Total Length No. of Spicules Total Length No. of Spicules Species Mean SE n Mean n Mean SE n Mean SE n B. bivittata 170.24 22.40 5 19:50) 16102 4 170.05 19.02 v] 20s) WEDS a B. imbifida 7223 5.38 3 DIDO VAS 5) 178.78 1.91 3 L938 1.84 6 B. nobilis 189.60 10.99 18 25.14 4.42 Wf 199:20 10:47 16 £29.27 Gis 1 B. glabra 209.70 16.20 12 191645 3°91 DIO G8) 12 23°36 Sory | Il B. dorsasetula 158.36 B54.) 15 1-925 R68) 2 155.14 4:33) 17 20.50 1.46 14 Ph. opaca 188.73 16.66 14 26H OOS aly: 185.40 15.66 15 15.94 See) we P. subaspera 266.00 _- 1 21.00 — | DSNESi --- | 36.00 _— l P. cressoni 223.08 35.64 3} 18.00 7.45 3 217.68 39.98 3 37.91 10.24 5 P. lineata 249.25 30.24 3 20.50 — 2 268.55 — ] 24.00 — 1 P. plebeia 184.37 8.59 8 23 Si ol V 179.81 8.95 9 — — 0) P. rufa 239109 N29 12 15283)" 354 6 24440 15.62 13 11.75 6.67 + P. pseudorufa 187.29 8.06 11 L722) 22877 9 187.33 TeO a2 19.62 4.66 8 P. basalis 183.54 —- 2 — — 0 186.20 — 2 7.00 _ 2 P. flaviceps 169.29 11.98 7 18.57 7.04 of 167.36 8.46 12 5.60 5.38 5 P. scalpellifera 198583") 39:42 4 — os 0 LOTS ile Si 3 10.00 — 2 P. hastata — —- O 18.00 — I —- a 10) 6.00 — | PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. SEM photomicrographs of mandibles of female. A, Bezzia bivittata. B, B. imbifida. C, B. nobilis. D, B. glabra. E, B. dorsasetula. F. Phaenobezzia opaca. G, Palpomyia subaspera. H, P. cressoni. White bars equal 8 wm. VOLUME 97, NUMBER 4 805 Fig. 2. SEM photomicrographs of mandibles of female. A, Palpomyia lineata. B, P. plebeia. C, P. rufa. D, P. pseudorufa. E, P. basalis. F, P. flaviceps. G, P. scalpellifera. H, P. hastata. White bars equal 8 jm. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. SEM photomicrographs of mandibles of female. A, Bezzia bivittata. B, B. imbifida. C, B. nobilis. D, B. glabra. E, B. dorsasetula. F Phaenobezzia opaca. White bars equal 8 wm. VOLUME 97, NUMBER 4 807 Fig. 4. SEM photomicrographs of mandibles of female. A, Palpomyia subaspera. B, P. cressoni. C, P. lineata. D, P. plebeia. E, P. rufa. F, P. pseudorufa. White bars equal 8 1m. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs: pellifera. D, P. hastata. White bars equal 8 wm. with up to 13 long, coarse, antrorse, some- times branched, teeth on the medial edge and, in nine species, up to 10 short antrorse teeth on the lateral edge (Figs. 6—9). Max- imum width is attained at approximately mid-length of the blade and is 14.1—-19.3% of their total length. There is little reduction in width proximal to mid-length. The artic- ulating mechanism of female mandibles is absent in males. The basal portion curves approximately 45° to the long axis of the blade. Labrum.—The labrum of both sexes is moderately sclerotized, ventrally concave in cross section and proximally joined to the hypopharynx on both sides. Two short sen- silla basiconica occur in pits located asym- metrically on the ventral surface near the proximal end of the row of lateral spicules (Figs. 10, 11). In females the tip is rounded and terminates in two median short, peg- like structures with rounded points; lateral to these are three, sometimes two, rounded structures. Antrorse spicules on the distal ventrolateral surface are short triangular or stout spinelike structures that occur in a sin- SEM photomicrographs of mandibles of female. A, Palpomyia basalis. B, P. flaviceps. C, P. scal- gle series on each side. The labrum of males is more pointed than in conspecific females, in most species the terminal me- dian peg-like structures are replaced by an- trorse spicules, and the lateral rounded structures are reduced in size or absent. The ventrolateral spicules are longer and more slender than those of females and occur in one to three rows. Hypopharynx.—In both sexes the mod- erately sclerotized hypopharynx has a deep dorsal salivary groove located above a ven- tral keel; both structures become less pro- nounced distally and terminate proximal to the tip. Antrorse spicules of varying lengths occur on the rounded or semipointed tip of the hypopharynx of females and, in most species, extend for varying distances prox- imally along its lateral edges. The spicules vary in shape from tooth-like to spike-like to filiform (Fig. 12). The hypopharynx of males is less heavily sclerotized, shorter and has longer spicules than those of con- specific females (Fig. 13). Lacinia.—In both sexes of all species ex- amined, the laciniae are acute triangular in VOLUME 97, NUMBER 4 809 imbifida. C, B. nobilis. D, Fig. 6. SEM photomicrographs of mandibles of male. A, Bezzia bivittata. B, B. al B. glabra. E, B. dorsasetula. F, Phaenobezzia opaca. G, Palpomyia subaspera. H, P. lineata. White bars equ 8 pm. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 7. shape, have no teeth or spicules on either the tip or edges and are one-half to three- fourths the length of the mandibles (Figs. 14-15). Their structure indicates that they serve as the lateral boundary of the pro- boscis food canal. The base is thick and, on broken specimens, appears to be hollow (Fig. 14C). Laciniae of both sexes of all species are similar and do not differ dis- tinctively in appearance from each other, but females have longer laciniae than con- specific males. Maxillary palpal sensory organ.—In both sexes of all species the filiform sensilla, known as Newstead’s sensilla or bulb or- gans (Lewis, 1973), that constitute the sen- sory organ, arise from an unindented sur- face on the distal end of the third palpal segment as in Corethrellidae (McKeever 1986), Ceratopogonidae (McKeever et al. 1991) and Psychodidae (Brinson et al. SEM photomicrographs of mandibles of male. A, Palpomyia plebeia. B, P. rufa. C, P. pseudorufa. D, P. basalis. E, P. scalpellifera. White bars equal 8 pm. 1993). In contrast, the sensilla arise from a pit in Culicoides (Chu-Wang et al. 1975) and some Ceratopogonini where the pit ranges from shallow, as in Alluaudomyia bella (Coquillett), to deep, with heads of the sensilla below its rim, as in Downeshelea stonei (Wirth) (McKeever et al. 1991). In Palpomyiini the long stalk of each sensil- lium originates from a recessed socket as in Culicoides (Rowley and Cornford 1972) and Ceratopogonini (McKeever et al. 1991) and the tip is expanded to form a head, the shape of which varies among the species (Figs. 16, 17). In some species, the sensilla are closely grouped and their heads are in approximately the same plane, but in others they originate along over half the length of the palpal segment and the stalk of one sen- sillium may not extend to the point of ori- gin of a more distal one. The number of VOLUME 97, NUMBER 4 811 B, B. imbifida. C, B. nobilis. D, Fig. 8. SEM photomicrographs of mandibles of male. A, Bezzia bivittata. B. glabra. E, B. dorsasetula. F, Phaenobezzia opaca. White bars equal 8 pm. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 9. SEM photomicrographs of mandibles of male. A, Palpomyia subaspera. B, P. lineata. C, P. plebeia. D, P. rufa. E, P. pseudorufa. F, P. scalpellifera. White bars equal 8 pm. VOLUME 97, NUMBER 4 813 Fig. 10. SEM photomicrographs of labra of female. A, Bezzia bivittata. B, B. imbifida. C, B. nobilis. D, B. glabra. E, B. dorsasetula. F, Phaenobezzia opaca. G, Palpomyia cressoni. H, P. lineata. 1, P. plebeia. J, P. rufa. K, P. pseudorufa. L, P. basalis. M, P. flaviceps. N, P. scalpellifera. O, P. hastata. White bars equal 8 pm. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 11. SEM photomicrographs of labra of male. A, Bezzia bivittata. B, B. imbifida. C, B. nobilis. D, B. glabra. E, B. dorsasetula. F Phaenobezzia opaca. G, Palpomyia subaspera. H, P. lineata. I, P. plebeia. J, P. rufa. K, P. pseudorufa. L, P. basalis. M, P. scalpellifera. White bars equal 8 jm. VOLUME 97, NUMBER 4 815 Fig. 12. SEM photomicrographs of the hypopharynx of female. A, Bezzia bivittata. B, B. imbifida. C, B. nobilis. D, B. glabra. E, B. dorsasetula. F Phaenobezzia opaca. G, Palpomyia cressoni. H, P. lineata. 1, P. plebeia. J, P. rufa. K, P. pseudorufa. L, P. basalis. M, P. flaviceps. N, P. scalpellifera. O, P. hastata. White bars equal 8 p.m. sensilla is approximately equal in both sex- es of the same species (Tables 4, 8). Description of Mouthparts by Species Bezzia_ bivittata. Mandible (female) (Figs. 1A, 3A): 114-175 pm (n = 13) long, 23-37 wm (n = 13) wide; tooth row 23% of total length; 7-15 (n = 19) small coarse medial teeth, uniform throughout, their camber 35°, pitch 71°; 0-8 (n = 19) small basal teeth, sometimes bifurcate; 0-6 (n = 20) small lateral teeth. Mandible (male) (Figs. 6A, 8A): 121- 140 pm (n = 10) long, 19-24 wm (n = 10) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 13. SEM photomicrographs of the hypopharynx of male. A, Bezzia bivittata. B, B. imbifida. C, B. nobilis. D, B. glabra. E, B. dorsasetula. F, Phaenobezzia opaca. G, Palpomyia subaspera. H, P. lineata. I, P. plebeia. J, P. rufa. K, P. pseudorufa. L, P. basalis. M, P. scalpellifera. White bars equal 8 ym. VOLUME 97, NUMBER 4 817 Fig. 14. opaca. D, Palpomyia cressoni. E, P. lineata. F, P. plebeia. White bars equal 8 pm. wide; 4—13 (n = 10) large medial teeth, api- cal ones slender, some proximal ones may be bifurcate; O—2 (n = 10) small basal teeth; O—1 (n = 10) small lateral projections. Labrum (female) (Fig. 10A): 146-189 wm (n = 5) long, 21-29 wm (n = 7) wide, 14—22 (n = 4) short, coarse, widely-spaced spicules. Labrum (male) (Fig. 11A): 157-160 pm (n = 2) long, 16-21 wm (n = 2) wide; 18- 20 (n = 2) short, coarse, widely-spaced spicules in a single row on the ventrolateral surfaces; additional spicules on the subapi- cal ventral surface; spicules replace the ter- minal median projections of females. Hypopharynx (female) (Fig. 12A): 142- 200 pm (n = 7) long, 14-40 pm (n = 7) wide; 17—32 (n = 4) short, semi-blunt spic- ules on margins and apex. Hypopharynx (male) (Fig. 13A): 142— 186 wm (n = 4) long, 23-32 pm (n = 4) wide; 7—21 (n = 3) long, slender spicules on distal margins and apex, minute spicules on margins proximally. Palpal sensory organ (female) (Fig. 16A): 3-4 (n = 3) closely grouped sensilla 14-15 pm (n = 2) long, heads spatulate. Palpal sensory organ (male) (Fig. 17A): 2-6 (n = 5) closely grouped sensilla 16—19 wm (n = 3) long, heads spatulate. SEM photomicrographs of laciniae of female. A, Bezzia nobilis. B, B. dorsasetula. C, Phaenobezzia Bezzia imbifida. Mandible (female) (Figs. 1B, 3B): 158-182 wm (nm = 11) long, 27— 35 pm (n = 12) wide; tooth row 24% of total length; 7-12 (mn = 12) small, coarse medial teeth, diminishing in size proximal- ly, their camber 25°, pitch 75°; 14 (n = 12) small, basal teeth; 3-6 (n = 12) minute lateral teeth. Mandible (male) (Figs. 6B, 8B): 134— 165 pm (n = 10) long, 20—23 pm (n = 10) wide; 6—11 (m = 11) large medial teeth, api- cal ones slender, proximal ones coarse; 0— 2 (n = 11) small basal teeth; 0-3 (n = 11) minute lateral teeth. Labrum (female) (Fig. 10B): 176—180 wm (n = 2) long, 28-29 wm (n = 5) wide; 20-30 wm (n = 5) coarse, closely spaced spicules. Labrum (male) (Fig. 11B): 152—169 pm (n = 4) long, 19-21 wm (n = 6) wide; 17 (n = 1) sharp, widely-spaced spicules on the ventrolateral surfaces and as replace- ments for the median projections of fe- males. Hypopharynx (female) (Fig. 12B): 177— 178 wm (n = 3) long, 19-25 wm (n = 6) wide; 17-22 (n = 6) short, sharp, irregu- larly spaced spicules on lateral margins, and sparse, irregularly spaced projections on apex. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 15. SEM photomicrographs of laciniae of male. A, Bezzia bivittata. B, B. imbifida. C, B. glabra. D, B. dorsasetula. E, Phaenobezzia opaca. F, Palpomyia subaspera. G, P. lineata. H, P. plebeia. 1, P. rufa. J, P. pseudorufa. K, P. scalpellifera. White bars equal 8 wm. VOLUME 97, NUMBER 4 aE Fig. 16. 819 SEM photomicrographs of sensilla of female. A, Bezzia bivittata. B, B. imbifida. C, B. nobilis. D, B. glabra. E, B. dorsasetula. F. Phaenobezzia opaca. G, Palpomyia subaspera. H, P. cressoni. 1, P. lineata. J, P. plebeia. K, P. rufa. L, P. pseudorufa. M, P. flaviceps. N, P. scalpellifera. O, P. hastata. White bars equal 8 pm. 4-7 (n = 5) closely grouped sensilla 16—19 Hypopharynx (male) (Fig. 13B): 153 168 pm (n = 3) long, 19-20 pm (n = 4) wide; 11—21 (n = 4) long, slender, closely spaced spicules on distal margins and apex. Palpal sensory organ (female) (Fig. 16B): 5—8 (n = 7) closely grouped sensilla 37.5—37.5 wm (n = 3) long, heads spatulate. Palpal sensory organ (male) (Fig. 17B): wm (n = 4) long, heads spatulate. Bezzia nobilis. Mandible (female) (Figs. IC, 3C): 142-196 wm (n = 34) long, 31- 96 pm (n = 37) wide; tooth row 30% of total length; 7—10 wm (n = 45) large coarse medial teeth, diminishing in size proximal- ly, their camber 32°, pitch 71°; 0-8 (7 = PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 17. SEM photomicrographs of sensilla of male. A, Bezzia bivittata. B, B. imbifida. C, B. nobilis. D, B. glabra. E, B. dorsasetula. F, Phaenobezzia opaca. G, Palpomyia subaspera. H, P. lineata. 1, P. plebeia. J, P. rufa. K, P. pseudorufa. L, P. scalpellifera. White bars equal 8 wm. VOLUME 97, NUMBER 4 821 Table 4. Measurements (1m) of laciniae and measurements and number of sensilla of maxillary palpal sensory organ of female Palpomyliini. Laciniae Sensilla Total Length Maximum Width No. per Organ Total Length Species Mean SE n Mean SE n Mean SE n Mean SE n B. bivittata 105.51 10.94 6 26.38 37 6 S167 E43 3 14.75 —- 2 B. imbifida 106.90 7.47 8 26.27 1.95 8 6.14 1.41 i 37.50 — 3} B. nobilis 115.42 5.02 14 29719. 229" is 6.87 1.06 16 L940" e555 B. glabra 91.77 20.08 4 26.27 952 -+ 529i 120) 16 EZ OR OVA SG B. dorsasetula 83.09 3163) 7 19 25.62 0.70 16 5.42 0.43 24 19580) 2103, 5 Ph. opaca 81.36 28s 29.65 IRSORD 27 6160) 0:72 5 18.47 1.85 6 P. subaspera 127.68 = 2 43.22 -- 2 10.00 oo | 2650) 3:68) 6 P. cressoni 142.58 21.57 5) 3957 3.80 5 WAS) als, 7 DIESST 2245 6 P. lineata 110.83 76.30 3 BONS) 24735 3 12.00 — ] 28:93) 228i) 16 P. plebeia 115.49 59 als 28.21 2.13 14 Tess» 11X0)i1 13 1650) 1/26 P. rufa 154.92 8.67 19 36.48 Possisy INE) S105 eS 4 Sas WE83) 23, 16 P. pseudorufa 121.47 12.74 9 Deli 1.96 9 Ss) (053! 17 20°80) 1055) P. basalis = an O _ _- 0) _- = O — — 0) P. flaviceps 97.62 24.42 5 27.04 16.63 3 Syoe ORWY 17/ 28:00) 2a > P. scalpellifera 130.78 50.04 3 37.68 15.61 3 9.80 3.56 5 30.85 1.42 5 P. hastata _ = 0) — _ 0) 12.00 a= ] 201345 3:07 45) small, basal teeth; 2—8 (n = 45) prom- inent lateral teeth. Mandible (male) (Figs. 6C, 8C): 120— 169 wm (n 12) long, 20-32 wm (n = 15) wide; 2-13 (n = 15) small short medial teeth, apical ones long and slender; 0-4 (n 15) minute basal teeth; 0-3 (n 15) minute lateral teeth, proximal ones short, apical ones long and slender. Labrum (female) (Fig. 10C): 146-225 pm (n = 18) long, 23-40 wm (n = 20) wide; 17—30 (n = 7) coarse, closely spaced spicules in single row distally and double row proximally on ventrolateral surfaces. Table 5. Measurements (4m) of mandibles of male Palpomyiini. Total Length Maximum Width Width at Basal Tooth Length of Tooth Row Species Mean SE n Mean SE n Mean SE n Mean SE n B. bivittata 133.93 GullSa 10 22S 1k42) 10 1220552299 WO 25.80 6.95 10 B. imbifida 147.92 8.06 9 20:80), (072, wil ES Se O WS ane 33.12, 6:03 11 B. nobilis 147.63 8.89 12 25.36) 92530 25 9.84 2.44 15 26.60 8.79 15 B. glabra 138.54 9.77 6 25.67 W442 “10 TOOe $2306, 10 14:89" 5:55 lO B. dorsasetula 114.61 2.14 17 21.87 [E355 as PESO 0295 18 222A ae aS Ph. opaca 118.51 6.14 18 24.80 1.54 20 10.83 129) 20 7-861" 2:03 20 P. subaspera 130.34 =10.38 5 WS\0B) P57) 6 13.60 0.93 u 30.02 5.40 7 P. cressoni —- = 0 — _— 0) a — 0 — a 10) P. lineata 146.30 11.17 14 26118) 822525 SiS 11.53 1925 Vall 23.16) D:469 5 P. plebeia 118.59 2.31 18 20:02, RSI 18 7.54 1.04 18 15.81 1.60 18 P. rufa 134.42 8.02 15 234 KGS 15 10.02 O07 als PHlese) a Spy} II) P. pseudorufa L337 3:89 22 2031 10:80 22 TEDO OMS" = 22 PANTANS Il 7272 P. basalis 105.07 — 2 19.95 — 2 — — 0) — — 0) P. flaviceps -— — O — -—— 0) — — O — — O P. scalpellifera LOS) 25289 4 22.61 175 4 TESS OaLS 4 [330930 4 P. hastata — = O —_ 0 — — 0 — — 0 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ble 6. Measurements (wm) and number of spicules of mandibles of male Palpomyiini. Length of No. of Coarse No. of Small Longest Tooth Medial Teeth Basal Teeth No. of Lateral Teeth Species Mean SE n Mean SE n Mean SE nD Mean SE n B. bivittata 1.92 0.53 9 7.40 2.06 10 0.20 0.45 10 0.70 0.23 10 B. imbifida Ds 0.12 12 8.36 1.24 11 0.36 0.54 11 0.45 0.70 11 B. nobilis 4.41 0.27 39 5.87 1.61 15 0.67 0.77 15 0.40 0.50 1S B. glabra 5.19 0.44 26 4.30 1.58 10 0.50 0.61 10 0.20 0.30 10 B. dorsasetula 1.92 0.41 18 5.94 0.47 18 0.00 a 18 0.00 — 18 Ph. opaca 2.24 0.47 19 4.10 0.60 20 0.00 — 20 055 0.72 20 P. subaspera 1.90 0.66 7 4.14 NES) 7 —- os 10) 0.71 0.66 Tl P. cressoni a — 10) os —- 10) a — 10) — — 10) P. lineata 2S 0.59 IIS) 5.94 1.00 16 0.31 0.42 16 0.19 0.29 16 P. plebeia 1.81 0.37 18 5.00 el 18 0.00 = 18 OP, 0.26 18 P. rufa 2.08 0.57 15 4.80 0.70 15 0.27 0.57 15 0.00 a 14 P. pseudorufa 1.99 0.27 22 4.86 0.31 22 0.00 os 22 0.00 as 22 P. basalis —_— — 10) _— —- 0 —_ _- (0) oe a 10) P. flaviceps — — 10) —_ -—— 10) = — 10) _- os 10) P. scalpellifera 1233 10) 4 4.25 2.00 4 0.00 — 4 0.00 — 4 P. hastata — — 10) a — 0 — -— 0) i —- 10) Labrum (male) (Fig. 11C): 160—186 wm (n = 7) long, 20-27 wm (n = 8) wide, 16— 29 (n = 8) spicules, the proximal ones coarse, distal ones long and slender. Hypopharynx (female) (Fig. 12C): 165— 229 wm (n = 16) long, 22—37 pm (n = 17) wide; 15-46 (n = 11) short, sharp, irregu- larly spaced spicules on lateral margins and apex. Hypopharynx (male) (Fig. 13C): 172-— 184 pm (n = 7) long, 17-33 pm (n = 8) wide; 11—22 (n = 6) long, slender, closely spaced spicules on distal margins and apex. Palpal sensory organ (female) (Fig. 16C): 2-10 (n = 16) closely grouped sen- silla 17-21 wm (n = 5) long, heads spatu- late. Palpal sensory organ (male) (Fig. 17C): Table 7. Measurements (ym) and number of spicules of labra and hypopharynx of male Palpomyiini. Labra Hypopharynx Total Length No. of Spicules Total Length No. of Spicules Species Mean SE n Mean SE n Mean SE n Mean SE n B. bivittata 158.27 _- A) 19.00 —_ 2 159.27 29.99 4 133337) SI7Eo2 3) B. imbifida [58:60 Mile7i7 4 17.00 — 1 159.15 18.79 3 16.50 6.69 4 B. nobilis 173.09 8.33 a Dili Seasstole oS 176.89 4.49 W/ 16.32 S19 il 6 B. glabra 154.72 8.31 3 14.00 3.04 5 159.27 21.04 4 13333 2.87 6 B. dorsasetula 142.16 4.75 9 18.67 4.28 6 139.92 4.59 9 15.87 1.92 8 Ph. opaca 141.74 11.09 7 22S OF 4S 137.39 9.76 10 122 2.66 8 P. subaspera 147.30 S257, 4 S67 S745 3 147.63 11.45 3 13.50 —- 2 P. cressoni — — 0) —- —- 0) — — 10) — a 10) P. lineata LPS 7 Fe22418 5 15.50 4.78 6 L/S2 lS 6 13.30 3.36 6 P. plebeia 144.78 6.68 7 USTs 2655 7 139.65 HAS: 6 13.30 1.47 10 P. rufa 160.40 10.64 10 14.25 2.08 8 162.09 15.44 8 11.50 5.40 8 P. pseudorufa 129.01 6.88 8 Li —2:29 69 129.01 5:07 14 14.82 1.46 11 P. basalis 143.64 a l 16.00 — i 140.98 oo 1 11.00 a 1 P. flaviceps — = 10) = — oO _- 0 a — 0) P. scalpellifera 133.00 — ] 11.00 —- ] IS 1267 — 1 4.00 a 1 P. hastata — — 0) _ 0) — _- (0) — a 0) VOLUME 97, NUMBER 4 823 Table 8. Measurements (um) of laciniae and measurements and number of sensilla of maxillary palpal sensory organ of male Palpomyiini. Laciniae Sensilla Total Length Maximum Width No. per Organ Total Length Species Mean SE n "Mean SE. Mean SE n Mean SE n B. bivittata 88.78 16.51 4 23.61 7.60 4 3.40 2.08 5 17.48 2.79 3 B. imbifida 95.91 3.45 9 ZO AT a 2285 6 5:50 2:05 5 16.93 1.75 4 B. nobilis L00:5077710:30; 10 ZItS2.) 3:06) iil 429) = O}86r 12 17.86 0.88 4 B. glabra 35:33 13.84 5 Lisi Sige 9'54' DS 1.00 9 ssi Koy PATAW) 5 B. dorsasetula 72.61 Syel les} = AIO) 20.21 236-510 A 25a OS, 8 L9F73i5 2228 3 Ph. opaca M35 9.94 if pe Syl B00) Vz 4:22 O71 18 L29) O87 4 P. subaspera 82.46 5.65 6 2305) 5241 5 4.17 1.68 6 21.06 2.13 4 P. cressoni —— -= (0) — — 0 — — (0) — — 0) P. lineata 85.56 5.09 9 28.50 3.61 V/ 3.86 0.64 7 LA O00 3 P. plebeia 86.24 21 Os: 22300 1e28en3 4.50 0.63 14 19.33 2.67 14 P. rufa 91.27 8.86 8 19.42 2.78 > 3285.) 0169) 7 13 17.60 2.12 5) P. pseudorufa 79.92 3.44 11 15.23 Neti 3.005 VO595> 18 19.39 = 1.96 5 P. basalis — — 0) — — (0) — — 0 — — O P. flaviceps = —— 0) — — 0) -- a 0) — — 0) P. scalpellifera lI a l 35.90 — 1 8.00 -_— 2 18.40 0.74 5 P. hastata — — 0 —_ ~~ 0 — — (0) — — 10) 2-6 (n = 12) closely grouped sensilla 17— 274 pm (n = 12) long, 20-39 um (n = 12) 18 wm (n = 4) long, heads spatulate. Bezzia glabra. Mandible (female) (Figs. 1D, 3D): 148-226 pm (n = 24) long, 28- 50 wm (7 = 26) wide; tooth row 28% of total length; 7-17 wm (nm = 27) medium size medial teeth, diminishing in size prox- imally, distal ones widely spaced, their camber 43°, pitch 89°; 0-4 (n = 27) small, basal teeth; O-10 (n = 27) small to promi- nent lateral teeth. Mandible (male) (Figs. 6D, 8D): 128— 149 pm (n = 6) long, 23—29 pm (n = 10) wide; 2—10 (n = 10) small to medium me- dial teeth, proximal ones slender; 0—2 (n = 10) minute basal teeth; O—1 (n = 10) minute lateral teeth. Labrum (female) (Fig. 10D): 169-269 pm (n = 11) long, 29-43 pm (n = 11) wide; 12-29 (n = 11) coarse, closely spaced spicules. Labrum (male) (Fig. 11D): 152-158 pm (n = 3) long, 21—28 ym (n = 6) wide, 11— 17 (n = 5) coarse spicules on the antero- ventral surfaces, apex without spicules, similar to that of females, but structures less developed. Hypopharynx (female) (Fig. 12D): 186— wide; 6—37 (n = 11) short, sharp, widely and regularly spaced spicules on_ lateral margins and apex. Hypopharynx (male) (Fig. 13D): 149— 178 pm (n = 4) long, 13-16 pm (n = 5) wide; 9-16 (n = 6) long, slender closely spaced spicules on most distal margins and apex, short spicules on margins proximally. Palpal sensory organ (female) (Fig. 16D): 1—9 (n = 15) closely grouped sensilla 10-12 pm (n = 6) long, heads spatulate. Palpal sensory organ (male) (Fig. 17D): 1-5 (n = 9) closely grouped sensilla 13—18 wm (n = 5) long, heads spatulate. Bezzia dorsasetula. Mandible (female) (Figs. 1E, 3D): 116-144 pm (n = 33) long, 24-32 pm (n = 35) wide; tooth row 23% of total length; 6—9 (n = 31) large coarse medial teeth, distal ones uniform, proximal ones progressively smaller, their camber 35°, pitch 85°; O-S (n = 31) small basal teeth; 2—7 (n = 33) prominent lateral teeth. Mandible (male) (Figs. 6E, 8E): 106—120 pm (x = 17) long, 20-31 pm (7 = 18) wide; 4-7 (n = 8) medium medial teeth, lateral teeth absent. Labrum (female) (Fig. 1OE): 149-172 324 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ="15) lone, 21-27 ner =a) ide: -12=23° @r = 12) coarse; ~closely: spaced spicules. Labrum (male) (Fig. 11E): 134-153 wm (n = 9) long, 17—24 pm (n = 9) wide, 15— 26 (n = 6) long, slender spicules in 2-3 rows on the ventrolateral surfaces and apex, apical structures of females absent. Hypopharynx (female) (Fig. 12E): 133— 169 wm (n = 17) long, 20-24 pm (n = 18) wide; 18—24 (n = 14) very short, regularly and widely spaced spicules on lateral mar- gins, those on the rounded apex closely spaced. Hypopharynx (male) (Fig. 13E): 133- 150 wm (n = Y) long, 15-20 pm (n = 10) wide; 12-19 (n = 8) short, widely and evenly spaced spicules on margins and apex. Palpal sensory organ (female) (Fig. 16E): 3-8 (n = 24) closely grouped sensilla 18— 22 wm (n = 5) long, heads filiform. Palpal sensory organ (male) (Fig. 17E): 3-5 (n = 8) closely grouped sensilla 19-20 wm (n = 3) long, heads fusiform. Phaenobezzia opaca. Mandible (female) (Figs. 1F 3F): 120-180 wm (n = 37) long, 29-43 wm (n = 35) wide; tooth row 24% of total length; 7-8 (n = 8) large coarse medial teeth, the row terminates abruptly, proximal tooth largest, others uniform, their camber 45°, pitch 71°; 0-2 (n = 8) small basal teeth; 3—6 (n = 8) small lateral teeth. Mandible (male) (Figs. 6E 8F): 104—142 um (n = 19) long, 20-32 pm (n = 21) wide; 2—6 (n = 21) medium medial teeth, O—1 (n = 21) small lateral teeth. Labrum (female) (Fig. 1OF): 153-245 wm (n = 14) long, 19-39 pm (n = 19) wide; 21—32 (n = 17) long, slender, closely spaced spicules in single row on both ven- trolateral and subapical surfaces. Spicules most abundant in this species of any ex- amined. Labrum (male) (Fig. 11F): 121-146 wm (n = 8) long, 19-44 um (n = 10) wide; 14—36 (n = 8) long, slender, closely spaced spicules in 2—3 rows on the ventrolateral and subapical ventral surfaces. Spicules IL (7 most abundant in this species of any ex- amined. Hypopharynx (female) (Fig. 12F): 152- 245 pm (n = 15) long, 4-37 pm (n = 17) wide; 0-9 (n = 17) long, slender spicules on the apex, absent from lateral margins. Hypopharynx (male) (Fig. 13F): 120- 156 pm (n = 11) long, 13—20 pm (n = 10) wide; 10-19 (n = 9) long, very slender spicules on pointed apex, absent from mar- gins. Palpal sensory organ (female) (Fig. 16F): 4-9 (n = 15) closely grouped sensilla 17— 21 wm (n = 6) long, heads spatulate. Palpal sensory organ (male) (Fig. 17F): 2-7 (n = 6) closely grouped sensilla 11—12 wm (n = 4) long, heads spatulate. Palpomyia subaspera. Mandible (female) (Figs. 1G, 4A): 226-238 wm (n = 2) long, 40-47 wm (n = 2) wide; tooth row 28% of total length; 9-10 (n = 2) large coarse me- dial teeth, largest midlength of tooth row, their camber 58°, pitch 85°; 2—2 (n = 2) small basal teeth; 3—4 (n = 2) minute to small lateral teeth. Mandible (male) (Figs. 6G, 9A): 118— 140 pm (n = 5) long, 20-27 pm (n = 6) wide; 3—7 (n = 7) medium medial teeth, O— 2 (n = 7) small lateral teeth. Labrum (female) (no Fig.): 266 wm (n = 1) long, 52 wm (n = 1) wide; 21 (n = 1) spicules. Labrum (male) (Fig. 11G): 144-150 wm (n = 4) long, 25-27 pm (n = 3) wide; 16— 20 wm (n = 3) spicules, long and slender and replace midapical structures of females, minute to short on ventrolateral surfaces. Hypopharynx (female) (no Fig.): 251 wm (n = 1) long, 49 wm (n = 1) wide; 36 (n = 1) spicules. Hypopharynx (male) (Fig. 13G): 145— 156 pm (n = 3) long, 20-21 pm (n = 3) wide; 7-20 (n = 2) medium length slender spicules on rounded apex and distal mar- gins. Palpal sensory organ (female) (Fig. 16G): 10 (n = 1) closely grouped sensilla 22-30 um (n = 6) long, heads spatulate. Palpal sensory organ (male) (Fig. 17G): VOLUME 97, NUMBER 4 2-7 (n = 6) closely grouped sensilla 20—23 wm (n = 4) long, heads same diameter as stalk. Palpomyia cressoni. Mandible (female) (Figs. 1H, 4B): 181-192 wm (n = 7) long, 40-50 pm (n = 10) wide; tooth row 20% of total length; 6—9 (n = 10) medium coarse medial teeth, largest midlength of tooth row then rapidly diminishing in size proximally, their camber 46°, pitch 86°; 14 (n = 10) small, basal teeth; 2—7 (n = 10) minute lat- eral teeth. Mandible (male): no specimens. Labrum (female) (Fig. 10G): 207-234 wm (n = 3) long, 32-40 pm (n = 4) wide; 15-21 (n = 3) spicules which are minute and widely spaced in a single row. Labrum (male): no specimens. Hypopharynx (female) (Fig. 12G): 209— 230 wm (n = 3) long, 29-48 pm (n = 5) wide; 29-48 (n = 5) short spicules on apex and lateral margins. Hypopharynx (male): no specimens. Palpal sensory organ (female) (Fig. 16H): 6—9 (n = 7) closely grouped sensilla 20-26 pm (n = 6) long, heads spatulate. Palpal sensory organ (male): no speci- mens. Palpomyia lineata. Mandible (female) (Figs. 2A, 4C): 120-239 wm (n = 8) long, 37—49 wm (n = Y) wide; tooth row 25% of total length; 7-11 (n = 14) large coarse me- dial teeth, uniform on distal three-fourths of row, then diminishing rapidly; 3—8 (n = 14) small basal teeth; 5—11 (n = 14) small lat- eral teeth. Mandible (male) (Figs. 6H, 9B): 120— 184 pm (n = 12) long, 21-31 pm (n = 11) wide; 3—10 (n = 16) large medial teeth, O— 2 (n = 16) minute lateral teeth. Labrum (female) (Fig. I1OH): 235-258 wm (n = 3) long, 33-40 pm (n = 4) wide; 18—23 (n = 2) short, widely spaced spicules on ventrolateral surfaces, closely spaced on subapical surface. Labrum (male) (Fig. 11H): 140-186 wm (n = 5) long, 23-33 wm (n = 9) wide; 9- 22 (n = 6) medium slender to coarse spic- 825 ules on ventrolateral and subapical ventral surfaces, absent from apex. Hypopharynx (female) (Fig. 12H): 269 wm (n = 1) long, 33-47 wm (n = 2) wide; 24 (n = 1) short, closely set spicules on rounded apex and margins. Hypopharynx (male) (Fig. 13H): 173- 186 wm (n = 5) long, 21-36 wm (n = 7) wide; 8-17 (n = 6) long, slender spicules on semirounded apex and distal margins. Palpal sensory organ (female) (Fig. 161): 11 (n = 1) longitudinally widely spaced sensilla 25—32 wm (n = 6) long, heads fu- siform. Palpal sensory organ (male) (Fig. 17H): 3-5 pm (n = YQ) longitudinally widely spaced sensilla 17-17 wm (n = 3) long, heads fusiform. Palpomyia _ plebeia. Mandible (female) (Figs. 2B, 4D): 146-171 wm (n = 19) long, 27-33 wm (n = 22) wide; tooth row 21% of total length; 6-8 (n = 21) large coarse medial teeth, largest proximal, their camber 55°, pitch 91°; 2-5 (n = 21) small basal teeth; 3—6 (n = 22) small lateral teeth. Mandible (male) (Figs. 7A, 9C): 110— 126 wm (n = 18) long, 17—29 wm (n = 18) wide; 3-11 (n = 18) large medial teeth, some bifurcate; 0-2 (n = 18) minute lateral teeth. Labrum (female) (Fig. 101): 110-197 wm (n = 8) long, 23-32 wm (n = 10) wide; 20-30 (n = 8) medium length, closely spaced spicules. Labrum (male) (Fig. 111): 130-153 pm (n = 7) long, 12-27 wm (m = 11) wide; 15—23 (n = 7) spicules which are long and slender and replace the midapical structures of females, short and coarse on ventrolat- eral surfaces. Hypopharynx (female) (Fig. 121): 161— 205 wm (n = YQ) long, 20-29 wm (n = 10) wide; 13—23 (n = 6) minute to short spic- ules on rounded apex and margins. Hypopharynx (male) (Fig. 131): 130-148 wm (n = 6) long, 16—20 wm (n = 10) wide; 10-16 (n = 10) short, slender or very short blunt spicules on rounded apex, minute ones on margins. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Palpal sensory organ (female) (Fig. 16J): 2 (n = 12) longitudinally widely spaced sensilla, 14-18 pm (m = 6) long, heads spatulate. Palpal sensory organ (male) (Fig. 171): 3-7 pm (n = 14) longitudinally widely spaced sensilla 17-20 wm (n = 4) long, heads spatulate. Palpomyia rufa. Mandibles (female) (Figs. 2C, 4E): 137—239 pm (n = 34) long, 32-50 pm (n = 34) wide; tooth row 21% of total length; 6—9 (n = 41) large coarse medial teeth, largest proximal, their camber 46°, pitch 90°; 0-7 (n = 41) small basal teeth; O—7 (n = 40) minute lateral teeth. Mandible (male) (Figs. 7B, 9D): 110— 162 wm (n = 15) long, 16—27 wm (n = 15) wide; 2—7 (n = 15) large medial teeth, no lateral teeth. Labrum (female) (Fig. 10J): 210—291 pm (n = 12) long, 27-57 pm (n = 16) wide; 12-20 (n = 10) minute, closely spaced spicules. Labrum (male) (Fig. 11J): 142-180 pm (n = 10) long, 20-40 pm (n = 10) wide; 10-18 (n = 8) spicules, long and slender and replace the midapical structures of fe- males, short and stout on subapical ventral surface, long and slender on ventrolateral surfaces. Hypopharynx (female) (Fig. 12J): 189- 310 pm (n = 14) long, 19-51 wm (1 = 16) wide; 9-18 wm (n = 6) short spicules on rounded apex and margins. Hypopharynx (male) (Fig. 13J): 142-192 wm (n = 8) long, 17—36 wm (n = 9) wide; O—18 (n = 8) short spicules, blunt on apex, sharp and widely spaced on lateral margins. Palpal sensory organ (female) (Fig. 16K): 1-12 (n = 18) longitudinally widely spaced sensilla, 16-19 wm (n = 6) long, heads spatulate. Palpal sensory organ (male) (Fig. 17J): 2-5 (n = 13) longitudinally widely spaced sensilla 16—20 wm (n = 5) long, heads spat- ulate. Palpomyia pseudorufa. Mandible (fe- male) (Figs. 2D, 4F): 145-173 wm (n = 28) long, 23—33 wm (n = 29) wide; tooth row 23% of total length; 6-8 (n = 35) large coarse medial teeth, largest midlength of row, their camber 68°, pitch 72°; 04 (n = 35) small basal teeth; O-5 (n = 35) minute lateral teeth. Mandible (male) (Figs. 7C, 9E): 101-130 wm (n = 22) long, 16-24 um (n = 16) wide; 4—6 wm (n = 22) large short medial teeth, no lateral teeth. Labrum (female) (Fig. 10K): 170-210 wm (n = 11) long, 12-31 pm (n = 14) wide; 10—24 (n = 9) short closely spaced spicules. Labrum (male) (Fig. 11K): 113-141 wm (n = 8) long, 13—20 pm (n = 8) wide; 7— 17 (n = 9) spicules which are long and slender and replace the midapical structures of females, short and coarse on ventrolat- eral surfaces. Hypopharynx (female) (Fig. 12K): 170— 210 wm (n = 12) long, 15—25 pm (n = 9) wide; | 1—26 (n = 8) medium, coarse, wide- ly spaced spicules on flattened apex and margins. Hypopharynx (male) (Fig. 13K): 118— 140 pm (n = 11) long, 12-17 wm (v = 9) wide; 10-18 wm (nm = 11) minute to short spicules on apex and margins. Palpal sensory organ (female) (Fig. 16L): 3-6 (n = 17) longitudinally widely spaced sensilla 20-21 jm (n = 5) long, heads spat- ulate. Palpal sensory organ (male) (Fig. 17K): 1—6 (n = 17) longitudinally spaced sensilla 17—21 wm (n = 5) long, heads spatulate. Palpomyia_ basalis. Mandible (female) (Figs. 2E, 5A): 146-157 wm (n = 4) long, 33-37 wm (n = 4) wide; tooth row 19% of total length; 6—6 (n = 8) medium, coarse, widely spaced medial teeth, uniform size, their camber 38°, pitch 103°; 0-3 (n = 8) small basal teeth; 3—5S (nm = 8) minute lateral teeth. Mandible (male) (Figs. 7D): 102—108 wm (n = 2) long, 20-20 pm (n = 2) wide; no data on teeth. Labrum (female) (Fig. 1OL): 180—187 wm (n = 2) long, 16-40 pm (n = 2) wide; VOLUME 97, NUMBER 4 no data on number of spicules which are short to minute. Labrum (male) (Fig. 11L): 144 wm (rn = 1) long, 16 wm (7 = 1) wide; 16 (” = 1) short, slender spicules on ventrolateral and subapical ventral surfaces. Hypopharynx (female) (Fig. 12L): 161— 192 wm (7 = 2) long, 13-15 pm (7 = 2) wide; 6—8 (n = 2) long, slender spicules on apex, absent on margins. Hypopharynx (male) (Fig. 13L): 141 pm (n = 1) long, 31 wm (n = 1) wide; 6-8 (n = 2) medium length, filiform spicules on pointed apex, absent on margins. Palpal sensory organ (female): no speci- mens. Palpal sensory organ (male): no speci- mens. Palpomyia flaviceps. Mandible (female) (Figs. 2E 5B): 125-172 pm (n = 22) long, 27-39 wm (n = 24) wide; tooth row 20% of total length; 4—9 (n = 33) medium coarse medial teeth, largest proximal, their camber 41°, pitch 78°; 0-3 (n = 33) small basal teeth; O-5 (n = 33) small lateral teeth. Mandible (male): no specimens mea- sured. Labrum (female) (Fig. 10M): 153-186 pm (n = 7) long, 24—64 pm (n = 13) wide; 9-29 (n = 10) short, closely spaced spic- ules on ventrolateral surfaces, sparce sub- apical ones. Labrum (male): no specimens. Hypopharynx (female) (Fig. 12M): 145— 188 pm (n = 12) long, 13—28 wm (n = 11) wide; 0—10 (n = 5) long, filiform spicules on rounded apex, absent on margins. Hypopharynx (male): no specimens. Palpal sensory organ (female) (Fig. 16M): 2—8 (n = 18) longitudinally widely spaced sensilla 24-29 wm (n = 5) long, heads fusiform. Palpal sensory organ (male): no speci- mens. Palpomyia_ scalpellifera. Mandible (fe- male) (Figs. 2G, 5C): 133—203 pm (n = 8) long, 28-44 pm (n = 8) wide; tooth row 15% of total length; 6-7 (n = 8) large coarse medial teeth, distal ones largest, di- 827 minishing slightly proximally, their camber 48°, pitch 90°; 1-3 (n = 8) small basal teeth; 3—7 (n = 7) small lateral teeth. Mandible (male) (Figs. 7E, 9F): 100-132 wm (n = 4) long, 21-24 wm (n = 4) wide; 3-6 (n = 4) large medial teeth; no (n = 4) lateral teeth. Labrum (female) (Fig. ION): 162—217 wm (n = 4) long, 25-32 wm (n = 4) wide; no data on number of spicules which are medium and closely spaced on subventral surfaces, sparse subapical ones. Labrum (male) (Fig. 11M): 133 wm (n = 1) long, 15 wm (7 = 1) wide; 11 (nm = 1) spicules, long and slender on tip, short and slender to stout on distal lateral and sub- apical ventral surfaces. Hypopharynx (female) (Fig. 12N): 165— 219 pm (n = 3) long, 20-28 wm (n = 3) wide; 9-11 (n = 2) long, slender spicules on rounded apex and distal margins. Hypopharynx (male) (Fig. 13M): 132 wm (n = 1) long, 11-20 pm (n = 2) wide; 4 (n = 1) short to medium length spicules on pointed apex and distal margins. Palpal sensory organ (female) (Fig. 16N): 6—14 (n = 5) longitudinally widely spaced sensilla 30-32 pm (n = 5) long, heads spatulate. Palpal sensory organ (male) (Fig. 17L): 6—10 (n = 2) longitudinally widely spaced sensilla 17—19 wm (n = 5) long, heads spat- ulate. Palpomyia hastata. Mandible (female) (Figs. 2H, 5D): 144-150 pm (n = 2) long, 32—34 wm (n = 2) wide; tooth row 17% of total length; 6—6 (mn = 2) large coarse me- dial teeth, distal ones uniform size, proxi- mal one largest, their camber 46°, pitch 97°; 1—2 (n = 2) small basal teeth; O-—3 (n = 2) small lateral teeth. Mandible (male): no specimens. Labrum (female) (Fig. 100): no data on length, 29 wm (nm = 1) wide; 18 (nm = 1) medium, closely spaced spicules. Labrum (male): no specimens. Hypopharynx (female) (Fig. 120): no data on length; 20 wm (n = 1) wide; 6 (n = 1) long slender spicules on rounded apex. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 'iypopharynx (male): no specimens. Palpal sensory organ (female) (Fig. 160): 12 (n = 1) longitudinally widely spaced sensilla 18-24 (n = 5) long, heads spatulate. Palpal sensory organ (male): no speci- mens. DISCUSSION Mandibles of female Palpomyiini resem- ble those of other entomophagous cerato- pogonids, e.g. Ceratopogonini (McKeever et al. 1991), Heteromyiini and Sphaeromiini (personal observations) Stenoxenini (Wirth and Ratanaworabhan 1972) of the Cerato- pogoninae, and ectoparasitic Atrichopogon meloesugans Kieffer and A. lucorum (Mei- gen) of the Forcipomyiinae (Glukhova 1981) in having large, widely-spaced re- trorse teeth. Mandibles of the subgenus 77ri- thecoides of Culicoides (Culicoidini), as il- lustrated by Glukhova for C. humeralis (1982), have large retrorse teeth that are similar in shape and approach the size of those of some Palpomyiini and Ceratopo- gonini. A second characteristic common to Pal- pomyiini and many members of Ceratopo- gonini, Heteromyiini and Sphaeromiini is the presence of antrorse teeth on the lateral edge of their mandibles. Lateral antrorse mandibular teeth also occur in Simulium venustum Say (Nicholson 1945) and Culi- coides (Trithecoides) anophelis Edwards (Glukhova 1982) in C. (T.) flavescens Mac- fie, and in C. (T.) paraflavescens Wirth and Hubert (Wirth and Hubert 1989). Their function in all of these groups is unknown, but they may assist with enlargement of the wound in the exoskeleton of prey. A third characteristic common to each of these four tribes is their reduced, unarmed laciniae, except for members of the anophelis group of Trithecoides that feed on mosquitoes and phlebotomine sand flies and that have strong teeth on the lacinae (Wirth and Hub- ert 1989). Mandibles of Palpomyiini function in the same manner as reported for Ceratopogon- ini (McKeever et al. 1991). The initial in- cision in the prey’s integument is made by antrorse distal mandibular teeth and en- larged by sawing action of the antrorse teeth. Mandibular action during this process is the same as that for Culicoides as de- scribed by McKeever et al. (1988) and Sut- cliffe and Deepan (1988), and for Simulium Latreille as described by Sutcliffe and Mclver (1984). Other elements of the pro- boscis enter the wound made by the man- dibles, but presence of antrorse spicules on the labrum and hypopharynx and absence of armature on the laciniae preclude these structures from functioning in producing the incision. Once the proboscis is inserted, saliva containing a strong protease flows through the salivary canal of the hypopharynx and digests the prey (Downes 1971). The di- gested material is drawn from the prey through the food canal formed by the la- brum dorsally, the mandibles ventrally and the laciniae laterally. Thus, the mandibles separate the food canal from the salivary canal. Although all reports indicate that Palpo- myiini usually feed on male Chironomidae and Ephemeroptera, Downes (1978) report- ed that his few records for B. glabra and P. lineata indicated that they select moderate to large prey species from these groups. This is the only specific reference to feed- ing by the species that we examined. Dur- ing a two-week study at Loch Lomond, Scotland in 1960, Downes (1978) found that four species of Palpomyia, in the fla- vipes group as defined by Grogan and Wirth (1979), fed exclusively on mayflies, while a species of Bezzia and another species of Palpomyia fed only on chironomids. Elements of the proboscis are basically similar in females of the 16 species exam- ined, but species specific variations exist in the mandibles, labrum and hypopharynx. Grogan and Wirth (1975, 1979) separated the Nearctic species of Palpomyia into four species groups, viz., tibialis, lineata, dis- tincta, and flavipes, on the basis of their VOLUME 97, NUMBER 4 male and female genitalia. Mandibular teeth of the first three of these groups (Figs. 4A— F) are longer and larger than those of either the flavipes group, viz. P. basalis, hastata, altispina, and flaviceps, (Figs. SA—D) or of species of Bezzia (Figs. 3A—E), with the ex- ception of B. glabra which has longer teeth than those of P. pseudorufa. Phaenobezzia opaca has the longest teeth (x = 6.2 um, n = 39) of any species studied except those of Palpomyia subaspera (xX = 8.0 wm, n = 2). Basic similarities probably are interre- lated with a common method of feeding, viz., sucking the saliva-digested contents of their prey, and their food specialization, i.e. males of their own species and males of Chironomidae and Ephemeroptera. Species specific differences in the various structures probably reflect minor differences in the re- stricted range of prey. In contrast to minor differences exhibited by elements of the proboscis of Palpomyi- ini, Glukhova (1981) reported basic simi- larity with marked differences in all ele- ments of the proboscis of species of Atri- chopogon. She attributed the differences to the diversity in prey (Coleoptera, Neurop- tera, and Lepidoptera) and to their feeding on different parts of the body (wing veins, thorax, and abdomen) of their prey. Male Nematocera belonging to species in which the females are entomophagous and hematophagous have unarmed mandibles that are smaller and less sclerotized than those of females and are thought to be nec- tar feeders (Downes 1971). In a later paper, Downes (1978) described the behavior of Bezzia feeding on honeydew and expressed the opinion that such feeding would be found to be a normal activity of both sexes of all predaceous midges. Mandibles of male Palpomyiini are fairly typical of en- tomophagous Ceratopogoninae. They differ from those of some other entomophagous groups, e.g. Ceratopogonini (McKeever et al. 1991) in having large antrorse, pointed teeth on their medial edge. These teeth may be as large as the retrorse teeth of females, 829 but their shape precludes their functioning as biting structures. The poorly sclerotized labrum of female Palpomyiini has antrorse spicules of various shapes, rather than teeth, and is therefore not adapted for active penetration of the prey. These spicules are most abundant in P. opaca. The labrum functions as the dor- sal wall of the food canal and the sensilla basiconica on its ventral surface probably function as chemoreceptors that monitor in- coming food. In Tabanus nigrovitattus Macquart, similarly located sensilla have four dendrites entering each sensillum and were considered to be chemosensory by Stoffolano and Yin (1983). The two termi- nal peg-like structures in species of Palpo- mylini resemble the pair of peg-shaped sen- silla immediately proximal to the terminal tricuspid teeth of Culicoides (McKeever et al. 1988). Sutcliffe (1994) states that a pair of “terminal labral pegs” is present in Cu- licidae, Simuliidae and Ceratopogonidae and that they are equipped with both chemosensory and mechanosensory ele- ments. The peg-like structures in Palpo- mylini probably are homologous to those in the foregoing three families and are che- moreceptors, mechanorecptors or both. The labra of females of other species of the family may be highly sclerotized and armored; e.g. four species of mammal-feed- ing Culicoides (McKeever et al. 1988) that have two tricuspid terminal teeth with a pair of sensilla at their base. The labra of species of Forcipomyia Meigen (Forcipomyiinae) are highly variable, with pointed apices with short lateral spicules in caterpillar feeders, smooth rounded apices in those that feed on Odonata, smooth apices rein- forced by longitudinal sclerotized thicken- ings in those that feed on amphibia, and apices with long bristles (spicules) on the edges and short spicules on the dorsal sur- faces in those that feed on hemolymph from wing veins of various insects (Glukhova 1981). The labra of male Palpomyiini are short- er, narrower, less sclerotized, and have lon- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and fewer spicules than conspecific fe- -s. The terminal structures present in fe- ales are smaller and reduced in number or absent. A similar relationship has been re- ported for Ceratopogonini (McKeever et al. 1991) and for two species of frog-feeding Corethrella Coquillett (McKeever 1986). Male P. opaca have a significantly greater number of spicules than any other species examined. Labra of males are species spe- cific, but differences among species are less pronounced than in labra of females. The hypopharynx of females is unarmed; its edges are either smooth or bear antrorse spicules as reported for Forcipomyia (Glu- khova 1981) and Ceratopogonini (Mc- Keever et al 1991). The shape and orien- tation of its spicules indicate that it has no function in penetrating the prey, but enters the incision made by the mandibles and conducts saliva into the wound through its salivary duct. The hypopharynx of females exhibits some generic differences in its shape and specific differences in distribution of its spicules. The hypopharynx of Phaenobez- zia opaca (Fig. 12F) differs from those of species of Bezzia (Figs. 12A—E) and Pal- pomyia (Figs. 12G—O) in being more point- ed and from species of Bezzia in lacking lateral spicules; males also have a more pointed hypopharynx (Figs. 13F) than spe- cies of Bezzia (Figs. 13A—E) and Palpo- myia (Figs. 13G—M). Females of the flavi- pes group of Palpomyia differ from those of other species of the genus in lacking lat- eral spicules (Figs. 12L—O), but do not dif- fer appreciably from each other. Terminal spicules are filiform in species lacking lat- eral spicules. The hypopharynx of males of all species has both terminal and lateral antrorse spic- ules; in most species they are larger than in conspecific females. The function of these spicules is unknown for either sex, and they are completely absent in some entomopha- gous Forcipomyia (Glukhova 1981). The weakly sclerotized, unarmored laci- niae of Palpomyiini, like those of Cerato- pogonini (McKeever et al. 1991), serve only as lateral boundaries of the food canal of the proboscis. They differ from those of some species of entomophagous Forcipo- myia which have strongly armored laciniae similar in degree of development to those of hematophagous Culicoides and are thought by Glukhova (1981) to serve in piercing the integument of and attachment to the flying prey. Sensilla in the maxillary palp sensory or- gan differ among species with respect to their number, points of origin on the palp of the individual sensilla, shape of their head, and their total length; a combination of these characters makes the sensory or- gans species specific. Thus, in females of all species of Bezzia (Figs. 16A—E) and in Phaenobezzia opaca, Palpomyia subaspera and P. cressoni (Figs. 16F—H), the sensilla originate in close proximity to each other and the heads are in approximately the same plane with respect to the long axis of the palpal segment. In the other seven spe- cies of Palpomyia (Figs. 16I—O) the points of origin of the individual sensilla may ex- tend over one-half the length of the palpal segment and proximal sensilla may not ex- tend to the point of origin of distal ones. Sensilla of females vary in mean length from 11.2 wm (n = 6) in B. glabra to 37.5 wm (n = 3) in B. imbifida. Heads of sensilla of male and female B. dorsasetula (Figs. 17E, 16E) and P. lineata (Figs. 17H, 161) are fusiform, heads of sensilla of male P. subaspera (Fig. 17G) are approximately the same diameter as the stalk and those of fe- males and males of all other species are spatulate. Palpal sensilla are reported to be olfac- tory receptors, including carbon dioxide de- tectors, in Culicoides (Rowley and Corn- ford 1972, Chu-Wang et al. 1975) and are thought to function as host detectors. How- ever, present evidence indicates that female Palpomyiini rely on visual rather than ol- factory stimuli when hunting and capturing their prey. Bezzia and Palpomyia are able to recognize visually distinctive areas that VOLUME 97, NUMBER 4 serve as swarm markers for their prey, even though no prey is present, and hover there for long periods until prey individuals ac- cumulate (Downes 1978). This indicates that in predaceous Ceratopogonidae the pal- pal sensilla serve little or no function in lo- cating prey. Therefore, the number of sen- silla should be approximately the same in both sexes of a given species and this was true for all species we examined. In con- trast, some species of female Culicoides, which locate their hosts by olfactory stim- uli, have four-fold increase in sensilla than do males of the same species (Rowley and Cornford 1972). For example, female C. nubeculosus Meigen have 12-17 sensilla per organ compared with 3—7 in males (Messaddeg et al. 1989). Their function in both sexes of Palpomyiini is unknown. LITERATURE CITED Brinson, FE J., S. McKeever, and D. V. Hagan. 1993. Comparative study of mouthparts of the phlebo- tomine sand flies Lutzomyia longipalpis, L. shan- noni, and Phlebotomus papatasi (Diptera: Psy- chodidae). Annals of the Entomological Society of America 86: 470-483. Chaika, S. Y. 1978. The fine structure of the chemo- receptors of the biting midges (Diptera: Cerato- pogonidae). Vestnik Moskovskoga Universiteta Seriya XVI Biologiya 2: 21—28 (In Russian with English summary.) Chu-Wang, I., R. C. Axtell, and D. L. Kline. 1975. Antennal and palpal sensilla of the sand fly Cul- icoides furens (Poey) (Diptera: Ceratopogonidae). International Journal of Insect Morphology and Embryology 4: 131-149. Dow, M. I. and E. C. Turner, Jr. 1976. A taxonomic revision of the Nearctic species of the genus Bez- zia (Diptera: Ceratopogonidae). Virginia Poly- technic Institute and State University Research Di- vision Bulletin 103: 1-162. Downes, J. A. 1971. The ecology of blood-sucking Diptera: an evolutionary perspective, pp. 232— 258. In Fallis, A. M., ed., Ecology and physiology of parasites. University of Toronto Press, Toronto, Canada. 1977. Evolution of feeding habits in Cera- topogonidae. Mosquito News 37: 279-280. 1978. Feeding and mating in the insectivo- rous Ceratopogonidae (Diptera). Memoirs of the Entomological Society of Canada 104: 1-62. Gad, A. M. 1951. The head capsule and mouth-parts in the Ceratopogonidae (Diptera-Nematocera). 831 Bulletin Societe Fouad I* Entomologique 35: 17— IBY Glukhova, V. H. 1981. A comparative morphological review of the mouthparts of the females and males in the subfamilies Dasyheleinae and Forcipomyi- inae. Entomologicheskoe Obozrenie USSR 60: 62-76 (in Russian; translation in Entomological Review 60: 59-72, 1981.) 1982. On the structure of the mouthparts in bloodsucking midges of the subgenus Trithecoides of the genus Culicoides (Ceratopogonidae). Para- zitologiya 16: 155-159 (In Russian with English summary.) Grogan, W. L., Jr. and W. W. Wirth. 1975. A revision of the genus Palpomyia Meigen of northeastern North America (Diptera: Ceratopogonidae). Uni- versity of Maryland Agricultural Experiment Sta- tion Miscellaneous Publication 875, College Park. 49 pp. 1979. The North American predaceous midges of the genus Palpomyia Meigen (Diptera: Ceratopogonidae). Memoirs of the Entomological Society of Washington 8: 1—125. . 1981. A new American genus of predaceous midges related to Palpomyia and Bezzia (Diptera: Ceratopogonidae). Proceedings of the Biological Society of Washington 94: 1279-1305. 1988. The predaceous midges of the world (Diptera: Ceratopogonidae; tribe Ceratopogonini). Flora and Fauna Handbook 4. Brill, New York. Lewis, D. J. 1973. Phlebotomidae and Psychodidae (sand flies and moth flies), pp. 155-179. Jn Smith, K. G. V., ed., Insects and other arthropods of med- ical importance. British Museum (Natural Histo- ry), London. McKeever, S. 1986. Mouthparts of the four North American Corethrella species (Diptera: Chaobor- idae), with detailed study of C. appendiculata. Journal of Medical Entomology 23: 502-512. McKeever, S., D. V. Hagan, and W. L. Grogan, Jr. 1991. Comparative study of mouthparts of ten species of predaceous midges of the tribe Cera- topogonini (Diptera: Ceratopogonidae). Annals of the Entomological Society of America 84: 93— 106. McKeever, S., M. D. Wright, and D. V. Hagan. 1988. Mouthparts of females of four Culicoides species (Diptera: Ceratopogonidae). Annals of the Ento- mological Society of America 81: 332-341. Messaddeq, M., M. Fabre, and M. Kremer. 1989. Scanning electron microscopy study of the sense organs in Culicoides nubeculosus (Diptera: Cera- topogonide). Annales de Parasitologie Humaine et Comparee 64: 224—237. Nicholson, H. P. 1945. The morphology of the mouth- parts of the non-biting blackfly, Eusimulium dac- otense D. and S., as compared with those of the biting species, Simulium venustum Say (Diptera: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Simuliidae). Annals of the Entomological Society if America 38: 281—297. ley, W. A. and M. Cornford. 1972. Scanning elec- tron microscopy of the pit of the maxillary palp of selected species of Culicoides. Canadian Jour- nal of Zoology 50: 1207-1210. Spinelli, G. R. and W. L. Grogan, Jr. 1985. Clastri- eromyia a new Neotropical genus of predaceous midges related to Palpomyia and Bezzia (Diptera: Ceratopogonidae). Proceedings of the Entomolog- ical Society of Washington 87: 329-334. Stoffolano; J. G:, Jr, and L. R..S: Yin. 1983: €om- parative study of the mouthparts and associated sensilla of adult male and female Tabanus_ni- grovitattus (Diptera: Tabanidae). Journal of Med- ical Entomology 20: 11-32. Sutcliffe, J. E 1994. Structure and possible functions of the labral tip complex in haematophagous members of Culicidae, Simulidae and Ceratopo- gonidae, pp. 216-217. In J. E. O’Hara [ed.], Third International Congress of Dipterology Abstract Volume. Biological Resources Division CLBRR Agriculture Canada, Ottawa. Sutcliffe, J. E and P. D. Deepan. 1988. Anatomy and function of the mouthparts of the biting midge Culicoides sanguisuga (Diptera: Ceratopogoni- dae). Journal of Morphology 198: 353-365. Sutcliffe, J. EK and S. B. McIver. 1984. Mechanics of blood-feeding in blackflies (Diptera: Simulidae). Journal of Morphology 180: 125-144. Wirth, W. W. 1983a. A review of the American pre- daceous midges of the Bezzia nobilis group (Dip- tera: Ceratopogonidae). Proceedings of the Ento- mological Society of Washington 85: 670-685. 1983b. The North American species of the cockerelli and dorsasetula groups of the preda- ceous midge genus Bezzia, subgenus Homobezzia (Diptera: Ceratopogonidae). Proceedings of the Entomological Society of Washington 85: 762— 782. Wirth, W. W. and W. L. Grogan, Jr. 1982. The pre- daceous midges of the genus Phaenobezzia in North America (Diptera: Ceratopogonidae). Mem- oirs of the Entomological Society of Washington 10: 179-192. . 1983. The Nearctic species of the Bezzia bi- vittata group (Diptera: Ceratopogonidae). Pro- ceedings of the Biological Society of Washington 96: 489-523. Wirth, W. W. and A. A. Hubert. 1959. Trithecoides, a new subgenus of Culicoides (Diptera; Cerato- pogonidae). Pacific Insects 1: 1-38. . 1989. The Culicoides of Southeast Asia (Dip- tera: Ceratopogonidae). Memoirs of the American Entomological Institute 44: 1-508. Wirth, W. W. and N. C. Ratanaworabhan. 1972. A revision of the tribe Stenoxenini (Diptera: Cera- topogonidae). Annals of the Entomological Soci- ety of America 65: 1368-1388. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 833-838 ENALLAGMA OPTIMOLOCUS, A NEW SPECIES OF DAMSELFLY FROM MONTANA (ODONATA: COENAGRIONIDAE) KELLY B. MILLER AND MICHAEL A. IVIE Department of Entomology, Montana State University, Bozeman, Montana 59717. Abstract.—Enallagma optimolocus n. sp. is described and diagnosed from streams in western Montana. Garrison’s 1984 key is modified to distinguish this species from the related E. carunculatum Morse and E. civile (Hagen). The competing aspects of two views of these specimens, 1.e. full species or hybrid individuals, is discussed. The occurrence of E. carunculatum and E. anna Williamson at the collecting sites is noted. The absence of E. civile from the areas surrounding the range of E. optimolocus is noted, with Roemhild’s 1975 (and Garrison’s 1984) record of E. civile for Flathead Co., Montana, corrected to E. carunclulatum. Illustrations of diagnostic features for the species are given. Key Words: The Odonata are perhaps the taxonomi- cally best known Order of North American insects, with virtually all species described, and the vast majority of both sexes and nymphs known (Kosztarab and Schaefer 1990, McCafferty et al. 1990). Indeed, in an analysis of the taxonomic status of the Odonata of North America north of Mexi- co, McCafferty et al. (1990) suggested that most of what remained to be discovered would be species range extensions along the southern boundaries of the region. The rate of description of new species from North America has slowed to an average of less than one per year, and at that rate, known collected-but-undescribed North American species may run out before the decade is over. This condition seems directly tied to the fact that the Odonata are second only to the butterflies and a few families of beetles in number of enthusiasts. Checklists and fau- nistic papers exist for most States and Prov- inces of the United States and Canada, in- cluding Roemhild’s (1975) list for Montana damselflies. Still, there are poorly sampled damselfly, Montana, taxonomy regions, especially in the central parts of the continent, where much remains to be done to document the fauna. Of the last three dragonflies described from North America (Carle 1992, Dunkle 1992, Vogt and Smith 1993) one came from the north central States, one from the Gulf Coast of Louisi- ana, and only one from the Texas—Mexico border area. Montana, with a huge and diverse area of relatively intact ecosystems and a very sparse human population (including ento- mologists and insect collectors), represents one of the most poorly known parts of the continent. Never-the-less, it was a surprise to discover an undescribed species of En- allagma while conducting a survey of the Odonata of Montana. Many Odonata are quite sensitive to wa- ter quality, and are susceptible to many pol- lutants (Carle 1979). McCafferty et al. (1990) list 30 North American species (of a total 415) as extinct, endangered, vulner- able, or rare. The presence of this new, rare, and possibly localized species in several of the world’s most famous fishing waters bes- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON peaks both a healthy environment and a ‘ontinuing need to protect those waters and their sources. The genus Enallagma contains approxi- mately 75 species world-wide (Garrison 1984), 36 of those in the United States and Canada (McCafferty 1990, plus a new rec- ord of a Mexican species in southern Ari- zona, Garrison in lit.). Garrison (1984) pro- vided the most recent revision of the spe- cies of western North America. The eastern species are covered in regional faunistic treatments by a combination of Walker (1953), Johnson (1972) and Dunkle (1990). The description below follows the format of Garrison (1984). Enallagma optimolocus, NEW SPECIES Holotype 6: MONTANA: Flathead County, Whitefish River 4.5 km SW White- fish; Swi 4, sw 74sec. 9. Ir 30 N-_R. 21 W. 27 July 1993, Kelly B. Miller leg. [Montana Entomology Collection, Bozeman (MTEC), on permanent loan to the California Acad- emy of Sciences, San Francisco]. Paratypes (16 36): 8—ibid.; 2—ibid., 07 August 1993; 2—ibid., 08 August 1991; 1—~ibid., 24 June 1992; I1—MONTANA: Madison County, Madison River, confluence with Chermy Creek-ssec.. 36, TE" 22S... 2. E14 July, 1992, Kelly B. Miller leg; 1—MON- TANA: Lewis and Clark Co., Beaver Creek, by river, 8/25/1993, D.W. Lundahl colr. [deposited in the MTEC (12) and the collections of K. B. Miller (2), T. Donnelly (1) and R. Garrison (1)]. All specimens were acetone-treated, and prepared in My- lar® envelopes. Etymology.—From the Latin optimum (best) and locus (place). A shortened form of the unofficial motto of Montana, ‘“The Last Best Place.” MALE. Head—Dorsum black with blue, subtriangular postocular spots between eyes and posterior margin of occiput; hind margin of occiput with mesal thin, blue line; rear of head light blue; face, genae blue; postclypeus black; anteclypeus, la- brum and labium blue; antennae black. Thorax—Pronotum black dorsally with lat- eral spots and hind margin blue; prothoracic pleura and sternum blue. Pter- othorax blue with longitudinal stripes black; mesostigmal plates black mesally, light blue laterally; mid-dorsal carina and antealar crests black; mid-dorsal stripe 1.8 the width of blue antehumeral stripe; average width of humeral stripe about 0.3 the width of mid-dorsal stripe, slight- ly wider at base (near legs) than at apex (near wings) [one paratype (from Madison Co.) with stripe constricted to a thin line medially]; black spot at upper end of sec- ond lateral suture; remainder of thorax blue. Legs—Femur black externally, pale internally; tibia pale posteriorly, black an- teriorly; tarsi pale with black apically on each tarsomere or, occasionally entirely dark; claws pale with black at tips; spines black. Wings hyaline, stigma black. Ab- domen (Fig. 1).—Segment I blue, occa- sionally with a small, lateral black spot apically; segment II blue with dorsal black transverse spot in posterior half of segment connected to black posterior annulus; seg- ment III blue with apical 0.50 to 0.70 black dorsally; segment IV blue with apical 0.50 to 0.75 black dorsally; segment V_ blue with apical 0.50 to 0.75 black dorsally; segment VI blue with apical 0.75 to 0.80 black dorsally; segment VII black dorsally with thin margin of blue proximally; seg- ment VIII and IX entirely blue, except for occasionally a small line of black apically, and sometimes a small, elongate black spot laterally on one or both segments; segment X black dorsally. All segments blue later- ally. Terminalia (Figs. 2, 3, 4, 5).—Supe- rior appendages black; inferior appendage pale except dorsally and apically black. In lateral view, superior appendage slightly longer than inferior appendage; pale tuber- cle visible between upper and lower arms of superior appendage; margin of juncture of arms of superior appendage at about right angle; in dorsal view, pale tubercle visible on interior of superior appendage as longitudinal mass, not visible beyond su- VOLUME 97, NUMBER 4 Figs. 1-5. Enallagma optimolocus. 1, Lateral view of abdomen. 2, Male abdominal terminalia, dorsal view, 3, left lateral view. 4-5, Right superior appendage of male genitalia, showing extremes of development of mesally pointed black tooth, lateral oblique view. perior appendage; mesally pointed, black tooth on lower arm of superior appendage variable, present and prominent in one- third of the types, absent in one-third of the types and intermediate in one-third of the types; inferior appendages evenly curved when viewed laterally. Length.—30.5—33.2 mm, abdomen, 23.9-27.0 mm, hind wing, 19.0-19.1 mm, hind femur 2.9—3.0 mm. Female and nymph.—unknown. DIAGNOSIS The form of the male genitalia will dis- tinguish this species from its congeners. Fe- males are not known. The color pattern of the abdomen (Fig. 1) of FE. optimolocus males is similar to that of E. carunculatum Morse, but E. optimolocus differs from that 336 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON species in having the superior appendage extending beyond the pale tubercle (Figs. 2, 4, 5). The terminalia of EF. optimolocus (Figs. 2, 3, 4, 5) are similar to those of E. civile (Hagen), but differ in the form of the superior appendage in lateral view (Fig. 3). The black tooth on the inner margin of the superior appendage is small, and usually surrounded by the pale tubercule (Figs. 4, 5). Lastly, in E. optimolocus the abdomen has tergites III, I1V, and V with 50% or more black along the dorsum. Geographically, E. optimolocus is sympatric and flies together with E£. carunculatum throughout its known range. Enallagma civile is rarely taken in Montana, to date only to the east of E. op- timolocus localities, with the closest record 45 km distant (K. Miller, unpubl.). Roem- hild’s (1975) record of E. civile from Flat- head Co., Montana (repeated on Garrison’s (1984) distribution map), is a misidentifi- cation of a specimen of E. carunculatum [MTEC]; In Garrison’s (1984) key E. optimolocus keys to couplet 7 where it fits neither choice. The following emendation of cou- plet 7 is required to accommodate E. opti- molocus in Garrison’s key: 7. In dorsal view, pale tubercle visible well be- yond the upper arm of superior appendage (Fig. 63b from Garrison 1984) Beal (aitag ts ra Rew Seen aac E. 7’. In dorsal view, pale tubercle not visible be- yond the upper arm of superior appendage (Figs. 2, 3, 4) 7a. Height of upper arm of superior appendage % to % total height of superior appendage; dorsal surface of abdominal segments III, TV, carunculatum Morse and V black in posterior one-quarter; lower superior appendage with a_ large mesally pointed black tooth (Fig. 64b from GarrisonpliO84) ies ceee ae ele E. civile (Hagen) 7a’. Height of upper arm of superior appendage approximately % total height of superior ap- arm of pendage; dorsal surface of abdominal seg- ments III, IV, and V black in posterior one- half or more; lower arm of superior append- age moderate to obsolete mesally pointed black: tooth ecsersceses ca ss E. optimolocus n. sp. DISCUSSION The discovery of these specimens has lead to a lively and very productive discus- sion among colleagues about the status of these individuals. The debate centers on 2 competing hypotheses: that the specimens represent an undescribed species, and that because they exhibit characteristics some- what intermediate between E. carunculatum and FE. anna Williamson, both of which oc- cur at all known localities of E. optimolo- cus, they represent a series of hybrids. An- other suspected parent, based upon mor- phology, is E. civile, but it is unknown in the areas where E. optimolocus has been found (see above). We have come down on the side of those who support the full spe- cies status, and provide the following ex- planation for that viewpoint. First, if non-overlapping characteristics intermediate between other species was grounds for hybrid status, several other spe- cies of Enallagma would have to be so con- sidered. Hybridization in damselflies is rare, but has been observed (Williamson 1906, Leong et al. 1992, Garrison pers. com., Donnelly pers. com.). All known cases fit the normal hybridization situation, i.e. ei- ther occurring commonly in a narrow hy- brid zone between 2 basically allopatric species, or very rarely in situations where sympatric species occur with one parent species very common, and the other ex- tremely rare. This makes good biological sense, since hybridization would be select- ed against in widely sympatric species, in- creasing the development of barriers to its occurrence. Neither of these expected situations fits this case. The locally available candidates for parents, Enallagma carunculatum and FE. anna, are sympatric over most of the western USA (Garrison 1984), including all areas where E. optimolocus have been taken. Thousands of specimens of these 2 species have been collected within this area of sympatry, and even if hybrids were very rare, they should have turned up throughout the range. At the Whitefish River site, some ¥s of male Enallagma specimens taken in 1993 were FE. optimolocus. Of the remain- ing *%, the vast majority were evenly divid- VOLUME 97, NUMBER 4 ed between E. carunculatum and E. anna. Only 3 damselflies were taken the day the Beaver Creek specimen was collected (D. Lundahl, pers. com.), and only a small number of specimens were taken the day the Madison River specimen was found. Because only the 1993 specimens were rec- ognized in the field, the others represent a relatively random sample mixed in with small samples of common species. Lack of females is explained by the fact that for conservation reasons only male Enallagma are taken during normal collecting, as fe- males are basically non-useful for docu- menting a species’ occurrence, and are therefore left to breed. The species has not been seen since the need for females for a type series was recognized. Another problem with resolving the hy- bridization hypothesis is the lack of phy- logenetic information. Based on arrange- ment of species in Garrison (1984), it would appear that the 2 possible parents are not closely related within the genus. If this is supported by a cladistic analysis, hybrid- ization would not be expected to occur be- tween such distantly related species. Lastly, from a conservation biology standpoint, it may be wisest to treat E. op- timolocus as a full species until the null hy- pothesis can be tested. However, the lack of females leaves the situation open to question, although female Enallagma are notoriously hard to distin- guish (Garrison 1984). Careful biological and genetic investigations are needed as a follow up test of the hypothesis that E. op- timolocus is indeed a good species. BIOLOGICAL NOTES The type locality is a slow, clear stream in a very broad, glacial valley in the Rocky Mountain Trench. The Whitefish River at that point has a silt bottom, with an abun- dant insect fauna, with at least twelve other Odonata species occurring at this site. Most common were EF. carunculatum, E. Calopteryx aequibilis (Say), Aeshna um- brosa Walker, Ophiogomphus severus Ha- anna, 837 gen, and Argia emma Kennedy. The Mad- ison River locality is at the confluence of a small stream with a broad river in an inter- mountain valley. The river is a freestone stream below a shallow reservoir, whereas Cherry Creek is silt-bottomed and clear. This area also possesses a very abundant insect fauna for a Montana freestone moun- tain stream, with six other Odonata known to occur in the immediate area. Most com- mon were EF. carunculatum, E. anna, Ophiogomphus severus Hagen, and Argia emma Kennedy. In general, the behavior of this species was observed to be similar to that of E. ca- runculatum at the type locality, flying to- gether with that species through the grasses and sedges emergent at the waters’ edge. Enallagma anna was also present, but tend- ed to fly over more Open water and rest on emergent water buttercup (Ranunculus aquatilis). Males flew very low over the stream often landing on grasses or emergent stream vegetation very near the water. They were wary and difficult to capture. ACKNOWLEDGMENTS We would like to thank D. Lundahl for providing specimens, R. S. Miller and D. L. Gustafson for discussions of species con- cepts and other issues, and B. C. Kondra- tieff, M. J. Westfall Jr., T. Donnelly, R. W. Garrison, J. Louton and an anonymous re- viewer for reviews of an earlier version of the manuscript. Donnelly and Garrison pro- vided extensive analysis of specimens, and valuable discussions of the possible options for a solution to the status of this popula- tion. This is contribution J-3026 of the Montana Agricultural Experiment Station. LITERATURE CITED Carle, EE: of the Odonata, with a list of rare and endangered 1979. Environmental monitoring potential Anisoptera of Virginia, United States. Odonato- logica 8: 319-323. 1992. Ophiogomphus (Ophiogomphus) aus- tralis spec. nov. from the Gulf Coast of Louisiana, with larval and adult keys to American Ophio- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gomphus (Anisoptera: Gomphidae). Odonatologi- ca 21: 141-152. Dunkle, S. W. 1990. Damselflies of Florida, Bermuda and the Bahamas. Scientific Publishers, Gaines- ville, Florida. 148 pp. . 1992. Gomphus (Gomphurus) gonzalezi spec. nov., a new dragonfly from Texas and Mexico (Anisoptera: Gomphidae). Odonatologica 21: 79— 84. Garrison, R. W. 1984. Revision of the genus Enallag- ma of the United States west of the Rocky Moun- tains and identification of certain larvae by dis- criminant analysis (Odonata: Coenagrionidae) University of California Publications in Entomol- ogy 105: 1-130. Johnson, C. 1972. The damselflies (Zygoptera) of Texas. Bull. Florida State Mus. Biol. Sci. 16: 1— 128. Kosztarab, M. and C. W. Schaefer. 1990. Conclusions, pp. 241-247. In Kosztarab, M., and Schafer, C. W., eds., Systematics of the North American In- sects and Arachnids: Status and Needs. Virginia Agricultural Experiment Station Information Se- ries 90-1. Virginia Polytechnic Institute and State University, Blacksburg. Leong, J. M. and J. E. Hafernik, Jr. 1992. Hybridiza- tion between two damselfly species (Odonata: Coenagrionidae): morphometric and genitalic dif- ferentiation. Ann. Entomol. Soc. Amer. 85: 662— 670. McCafferty, W. P., B. P. Stark, and A. V. Provonsha. 1990. Ephemeroptera, Plecoptera, and Odonata, pp. 43-58. In Kosztarab, M., and Schaefer, C. W., eds., Systematics of the North American Insects and Arachnids: Status and Needs. Virginia Agri- cultural Experiment Station Information Series 90-1. Virginia Polytechnic Institute and State Uni- versity, Blacksburg. Roemhild, G. 1975. The damselflies (Zygoptera) of Montana. Montana Agricultural Experiment Sta- tion Research Report 87: 1-53. Vogt, T. EF and W. A. Smith. 1993. Ophiogomphus susbehcha spec. nov. from north central United States (Anisoptera: Gomphidae). Odonatologica 22: 503-509. Walker, E. M. 1953. The Odonata of Canada and Alaska Vol. 1. Univ. Toronto Press. 292 pp. Williamson, E. B. 1906. Copulation of Odonata. En- tomol. News 17: 143-150. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 839-844 A NEW GENUS AND THREE NEW SPECIES OF CHEWING LICE (PHTHIRAPTERA: PHILOPTERIDAE) FROM PERUVIAN OVENBIRDS (PASSERIFORMES: FURNARITIDAE) ROGER D. PRICE AND DALE H. CLAYTON (RDP) Department of Entomology, Oklahoma State University, Stillwater, Oklahoma 74078 (Current address) 4622 Kinkead Avenue, Fort Smith, Arkansas 72903; (DHC) Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, England. Abstract.—The new genus Furnariphilus is described to include three new species from Peruvian hosts within the passerine family Furnariidae, subfamily Furnariinae: F. pagei, the type species of the genus, from Furnarius leucopus Swainson; F. griffithsi from Sclerurus mexicanus Sclater; and F. parkeri from Sclerurus caudacutus (Vieillot). Key Words: Clayton et al. (1992) published a survey of chewing lice collected in 1985 from a wide array of Peruvian bird taxa. During this project, a number of undescribed louse taxa were collected from hosts in the parv- orders Thamnophilida and Furnariida (Pas- seriformes). These taxa included a new spe- cies placed by Price and Clayton (1989) in a new genus of Menoponidae, Kaysius, and seven new species described by Price and Clayton (1993, 1994) in the existing phil- opterid genus Rallicola Johnston and Har- rison. We have recently examined addition- al philopterid lice collected during the Pe- ruvian project from members of the Tham- nophilida and Furnariida. Lice from three species of ovenbirds (Furnariidae: Furnari- inae) represent new species which are also members of an undescribed genus. The pur- pose of this paper is to name and charac- terize the new genus and to describe and illustrate the three new species that com- prise it. In the following descriptions, all mea- surements are in millimeters. Abbreviations for measured structures are explained the Ectoparasites, Peru, Furnariphilus, Furnariinae, Bird first time they are used. Host classification follows that of Sibley and Monroe (1990). Holotypes of the new species will be de- posited in The Field Museum (Chicago) and paratypes, as numbers allow, will be located in the collections of that museum and those of the National Museum of Nat- ural History (Washington, D.C.), Oklahoma State University (Stillwater), and the Uni- versity of Minnesota (St. Paul). Furnariphilus Price and Clayton, NEW GENUS Figs. 1-10 Type species: Furnariphilus pagei Price and Clayton, new species. Head (Figs. 1, 5) distinctly longer than wide, with preantennal region tapered to truncate hyaline margin; conus small; with single very short ocular seta; dorsoanterior head plate prominent, distinctly separated from remainder of head; male antenna with enlarged scape. Prothorax (Figs. 1, 5) quadrangular, with single seta near each lateroposterior corner. Metathorax posteriorly broadened, each 1) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ay au aS / 7 1 Figs. l-5. Furnariphilus page. 1, segment. 5, Female. side with row of medium to very long mar- ginal setae; prominent large rounded sternal plate with 4 setae. Abdomen (Figs. 1, 5, 6, 10) slender, with distinct partial median division of terga II(first apparent tergum)—VII for female, H-— IV for male. Male terga V—VIII much shorter than terga I-IV. Terga II with 4 me- =) i am Slee I. — > a —_ \ \ (( \ Ol Male. 2, Female ventral terminalia. 3, Male genitalia. 4, Male terminal dian setae, II-VI each with 2, [X without median setae. Only terga V—LX with lateral corner setae. Without pleural seta on II, with single short seta on III, medium seta on IV, long seta on V, and pair of long to very long setae on each of VI—VIII. Sterna II-VI prominent, undivided, each of II-V with medioposterior pair of setae, VI with VOLUME 97, NUMBER 4 Wie tc as D nna E a J g o) Figs. 6-10. 841 Bebe. oe << sae €) aaeee ° SF ITI — Furnariphilus griffithsi. 6, Male metanotum and abdomen. 7, Female ventral terminalia. 8, Male genitalia. 9, Male terminal segment. 10, Female metanotum and abdomen. 2 or 4 posterior setae. Subgenital plate of female posteriorly with regular to irregular row of short, stout spiniform setae along with finer setae posterior to them (Figs. 2, 7). Both sexes with prominent tergum IX followed by smaller sclerites associated with distinctly indented posterior margin; male with small accessory sclerite on each side of last tergal portion (Figs. 4, 9). Male genitalia (Figs. 3, 8) distinctive, with very short, stout widely-separated par- ameres each bearing median sensillum and terminal minute seta, complex mesomeral structures, and broad relatively short basal apodeme. Discussion.—This genus is easily rec- ognized by the combination of head shape, the sexually dimorphic antenna, the large thoracic sternal plate, the anterior abdomi- nal terga partially divided medially, the me- dian indentation of the terminal abdominal segment, the male with a small accessory piece on each side of the last segment, the unique male genitalia with the short broad parameres and complex mesomeral struc- tures, and the female lacking any evidence of a prominent seta-bearing tubercle later- oposterior to the subgenital plate. Of the philopterid lice associated with birds of these two host parvorders (Table 1), Rallicola, the most widely distributed genus with 30 species currently recognized, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cable 1. Host Parvorder Thamnophilida Family Thamnophilidae (typical antbirds) Parvorder Furnariida Superfamily Furnarioidea Family Furnariidae Subfamily Furnariinae (ovenbirds) Subfamily Dendrocolaptinae (woodcreepers) Superfamily Formicarioidea Family Formicariidae (ground antbirds) Family Conopophagidae (gnateaters) Family Rhinocryptidae (tapaculos) Classification of the parvorders Thamnophilida and Furnariida with their associated philopterid lice. Lice (No. of Species) Formicaphagus (12) Rallicola (1) Rallicola (12) Furnariphilus (3) Picicola (3) Brueelia (2) Rallicola (16) Formicaphagus (3) Formicaricola (8) Formicaphagus sp. Rallicola (1) shows certain affinities with the new genus described here. However, Rallicola females are characterized by the prominent seta- bearing tubercle posterior to each side of the subgenital plate, the males have geni- talia with long slender parameres and rela- tively simple mesomeral structures, and both sexes lack the marked medial inden- tation of the terminal abdominal portion. Additional lesser differences further sup- port the distinction between these two gen- era. The only other known _ philopterids from hosts within the superfamily Furna- rioidea are three species of Picicola Clay and Meinertzhagen and two species of Brueelia Keler (Table 1), all of which are quite different from Furnariphilus in gross head shape, genitalic features of both sexes, and other differences associated with struc- ture and chaetotaxy. Two other philopterid genera, Formicar- icola Carriker and Formicaphagus Carriker, are found on hosts within the superfamily Formicarioidea and the parvorder Tham- nophilida (Table 1). The former currently has eight recognized species and is restrict- ed to hosts in the family Formicariidae, with seven known only from the genus For- micarius. These lice are also of the Ralli- cola-type, but lack the prominent seta-bear- ing tubercles posterior to the female sub- genital plate. However, they differ from Furnariphilus in having both sexes with a distinct complete median division of ab- dominal terga II—VIII and without dimor- phic antennae; the males with genitalia of a grossly different type having ovoid para- meres, and with a rounded posterior abdo- men; and the females lacking a delineated terminal portion posterior to IX. The 15 recognized species of Formicaphagus are in both parvorders, with two additional unna- med series in our collection from the Con- opophagidae (Table 1); these lice differ from Furnariphilus in having characteristic male genitalia similar to those of Formi- caricola and both sexes with a distinctly broader head and abdomen, in addition to other differences. Furnariphilus pagei Price and Clayton, NEW SPECIES Figs. 1-5 Type host.—Furnarius leucopus Swain- son. Male.—As in Fig. 1. Preantennal head width (PAW), 0.30—0.33; temple width (TW), 0.38—0.39; head length (HL), 0.48— 0.50. Prothorax width (PW), 0.26—0.28. Metanotum with 6—8 (usually 7) medium to VOLUME 97, NUMBER 4 long setae posteriorly on each side; meta- thorax width (MW), 0.36—0.38. Abdominal terga with medium setae medially; sternal setae shorter, with 4 on sternum VI. Ab- domen width at V (AWV), 0.42—0.45. La- teroposterior corner of IX with 2 medium to very long setae on each side. Terminal portion with proportionately large accesso- ry piece on each side (Fig. 4). Genitalia (Fig. 3) with slightly curved parameres and complex of mesomeral structures as shown; genitalia width (GW), 0.10—0.11; genitalia length (GL), 0.23—0.25; genitalic paramere length (GPL), 0.04—0.05. Total body length (TL), 1.80—1.87. Female.—As in Fig. 5. Head (except for smaller antennal scape), thorax, and much of abdomen as for male. PAW, 0.34—0.36; TW, 0.41—0.43; HL, 0.53—0.55. PW, 0.29- 0.31; MW, 0.41—0.44. Differences from male associated primarily with posterior ab- dominal segments. With medium and very long lateroposterior ventral setae on each side of IX; AWV, 0.54—0.59. Subgenital plate (Fig. 2) posteriorly bearing submar- ginal row of 10—13 short spiniform setae on each side as well as total of 9-12 fine setae posterior to them; subgenital plate width (SPW), 0.29-0.31. TL, 2.14—2.27. Discussion.—This species is readily dis- tinguished from others of the genus by hav- ing the male terminalia as in Fig. 4, the male genitalia with features as in Fig. 3, the female with two medium to very long setae on each ventral lateroposterior corner of IX, the configuration of the setae of the female subgenital plate much as in Fig. 2, and the abdomen of both sexes with generally lon- ger median tergal and shorter sternal setae. Material examined.—Holotype male, ex F. leucopus, Peru: Dept. Cuzco: 20 km NW Pilcopata near Rio Tono, 750 wm, 25 il 985s. DH. +Clayton! #1157: Para- types: 17 males, 15 females, same data as holotype. Etymology.—This species is named for Roderic D. M. Page, University of Oxford, in recognition of his deep interest in and 843 contributions to the study of host-parasite cospeciation. Furnariphilus griffithsi Price and Clayton, NEW SPECIES Figs. 6-10 Type host.—Sclerurus mexicanus Sclater. Male.—Head and thorax much as for F. pagei (Fig. 1). PAW, 0.33; TW, 0.40-0.41; HL, 0.52—0.54. PW, 0.27—0.29; metanotum medially with short pair of setae (Fig. 6) and with 7—8 (usually 8) medium to very long posterior marginal setae on each side; MW, 0.40. Abdomen as in Fig. 6. With short median tergal setae; sternum VI with only single seta on each side. AWV, 0.46— 0.49. Terminal segment with short acces- sory piece on each side (Fig. 9). Genitalia (Fig. 8) with mesomeral posterior boundary bilobed and extending near end of para- meres; remainder of mesomeral structures as shown; GW, 0.10—0.11; GL, 0.23—0.24; GPE;.0:055) TE, 1-99=2710: Female.—Head and thorax much as for F. pagei (Fig. 5). PAW, 0.35—0.38; TW, 0.44—0.47; HL, 0.55—0.59. PW, 0.29-0.31; metanotum medially with pair of minute se- tae (Fig. 10) and marginally each side with 5—7 (usually 6) medium to very long setae; MW, 0.42—0.49. Abdomen as in Fig. 10. With only short medial setae on terga II— VIII; sternal setae on II-VI longer, with those on III-IV 0.06—0.09 long and extend- ing well over following sternum. AWYV, 0.56—0.66. Lateroposterior corner of IX each with only single medium ventral seta. Subgenital plate (Fig. 7) with highly irreg- ular row of 7—12 short submarginal spini- form setae on each side, along with total of 14-17 fine setae posterior to them; SPW, 0.28—0.30. TL, 2.38—2.59. Discussion.—This species is readily sep- arated from F. pagei by the male with dif- ferent genitalic parameral and mesomeral structures and the smaller accessory lateral portion of the terminal abdominal segment; the female with an irregular alignment of the submarginal short spiniform setae on the subgenital plate, only a single medium PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘tral seta on each side of abdominal seg- nent LX, and longer sternal setae on II-VI; ind both sexes with a pair of median me- tanotal setae and markedly shorter median abdominal tergal setae. Material examined.—Holotype male, ex S. mexicanus, Peru: Dept. Madre de Dios: Cerro de Pantiacolla, 20.vii1.1985, D. H. Clayton. Paratypes: 2 males, 3 females, same data as holotype. Etymology.—This species is named for Richard Griffiths, University of Oxford, in recognition of his work on the molecular phylogenetics of lice and their hosts. Furnariphilus parkeri Price and Clayton, NEW SPECIES Type host.—Sclerurus caudacutus (Vieil- lot). Male.—Mtuch as for F. griffithsi, except for consistently smaller body dimensions. PAW 5029 IW ,20:36; 4, OAS IPW 025: MW, 0.36. AWV, 0.41. TL, 1.90. Female.—Much as for F. griffithsi, ex- cept for consistently smaller body dimen- sions and shorter setae on sterna IIJ-—IV. PAW, 0.33; TW, 0.41—0.42; HL, 0.53-0.54. PW, 0.27—0.28; MW, 0.40—0.41. Sterna HI— IV with each seta only 0.03—0.04 long, at most extending only slightly over following sternal plate. AWV, 0.50—0.52. SPW, 0.24— 0:25. TL,.2.28=2.31. Discussion.—The non-overlapping di- mensional differences between the smaller F. parkeri and the larger F. griffithsi, cou- pled with the shorter sternal setae on HI-— IV for females of the former, enable ready separation of these two species. Admittedly, these differences are not as profound as we would prefer, indicating that these series from two different host taxa within the same genus are indeed closely related, but we believe them sufficient to justify rec- ognition of two distinct species. Material examined.—Holotype female, ex S. caudacutus, Peru: Dept. Madre de Dios: Cerro de Pantiacolla, 1030 m, above Rio Palotoa, 31.viii.1985, D. H. Clayton. Paratypes: | male, 1 female, same data as holotype. Etymology.—This species is named in honor of Theodore (Ted) Parker, the world’s most gifted field ornithologist, tragically killed during a 1993 field trip in Ecuador. ACKNOWLEDGMENTS Funds were provided to the junior author by The Field Museum, the Latin American Studies Center of the University of Chica- go, and NSF grant BSR-8508361 to J. W. Fitzpatrick for Peruvian faunal inventory work. This manuscript has been approved for publication by the Director, Oklahoma Agricultural Experiment Station, Stillwater. LITERATURE CITED Clayton, D. H., R. D. Gregory, and R. D. Price. 1992. Comparative ecology of Neotropical bird lice (In- secta: Phthiraptera). Journal of Animal Ecology 61: 781-795. Price, R. D. and D. H. Clayton. 1989. Kaysius emer- soni (Mallophaga: Menoponidae), a new genus and new species of louse from the wedge-billed woodcreeper (Passeriformes: Dendrocolaptidae) of Peru. Annals of the Entomological Society of America 82: 29-31. 1993. Review of the species of Rallicola (Phthiraptera: Philopteridae) from the woodcree- pers (Passeriformes: Dendrocolaptinae). Journal of Medical Entomology 30: 35—46. 1994. Review of the species of Rallicola (Phthiraptera: Philopteridae) from the antbirds, ovenbirds, and tapaculos (Passeriformes). Journal of Medical Entomology 31: 649-657. Sibley, C. G. and B. L. Monroe, Jr. 1990. Distribution and taxonomy of birds of the world. Yale Univer- sity Press, New Haven, CT. xxiv + 1111 pp. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 845-855 NOMENCLATURAL CHANGES IN THE PENTATOMIDAE (HEMIPTERA-HETEROPTERA) D. A. RIDER AND L. H. ROLSTON (DAR) Department of Entomology, North Dakota State University, Fargo, North Dakota 58105; (LHR) Department of Entomology, Louisiana Agricultural Experiment Station, Louisiana State University Agricultural Center, Baton Rouge, Louisiana 70803. Abstract.—Numerous nomenclatural problems in the Pentatomidae (Heteroptera) are discussed and corrected. These corrections have necessitated one generic replacement name, Keleacoris for Kelea Schouteden, 1958, and seven specific replacement names: Aelia chinensis for A. bifida Hsiao and Cheng, 1978; Aeliomorpha viridescens for A. viridis Azim and Shafee, 1987; Antestia dollingi for A. adspersa (Fabricius, 1803); Dic- tyotus walkeri for D. aequalis (Walker, 1867); Holcostethus mcdonaldi for H. piceus (Dallas, 1851); Menida signoreti for M. parvula (Signoret, 1858); and Sciocoris sahelensis for S. australis Linnavuori, 1975. Ten new combinations are also recognized: Basicryptus costalis ugandana (Linnavuori, 1982), B. striatus (Linnavuori, 1982), B. upembanus (Lin- navuori, 1982), Dalsira dallasi (Schouteden, 1912), D. kocki (Schouteden, 1962), D. ma- bokeana (Linnavuori, 1982), D. mulunguana (Linnavuori, 1982), D. niemboana (Linna- vuori, 1982), Keleacoris congolensis (Distant, 1910), and Stalius castaneus (Distant, 1893). One new generic synonym is proposed: Paposia China, 1962 as a junior synonym of Trincavellius Distant, 1900b. Key Words: Pentatomidae, Heteroptera, nomenclature, systematics While preparing a catalog of the Penta- tomidae of the world, we have discovered Herrich-Schaffer’s Pentatoma marginale predates Westwood’s usage of the binomen a number of nomenclatural problems that need to be corrected. We wish to keep the number of taxonomic changes in the cata- log to a minimum, so these corrections are made herein, prior to the publication of the catalog. I. ASOPINAE Oplomus festivus Dallas, 1851 Pentatoma marginalis Westwood, 1837: 37. [not Herrich-Schaffer, 1836, Roferta, (Pentatominae)| Oplomus marginalis: Dallas, 1851: 83; Thomas, 1992: 54, 60-61. Oplomus rutilis Dallas, 1851: 83-84. Oplomus festivus Dallas, 1851: 85. by a year. Pentatoma marginale Herrich- Schaffer has at one time or another been placed in the following genera: Rhaphigas- ter Laporte, Strachia Hahn, Nezara Amyot and Serville, and Acrosternum Fieber, and is currently a valid species in the genus Ro- ferta Rolston. See Rolston and McDonald (1981) for the complete synonymy of Ro- ferta marginalis. Nine different junior syn- onyms of P. marginale Westwood are avail- able, the oldest being Oplomus rutilis Dal- las, 1851: 83-84, and O. festivus Dallas, 1851: 85. There has been some confusion surrounding the name rutilis; the main source of which stems from the female type specimen. The type specimen of festivus is PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Le Right genital plates, caudo-ventro-lat- Figs. 1-2. eral view. 1, Stalius castaneus (Distant). 2, Stalius tar- tareus (Stal). a male, and there has been no problems as- sociated with the identity of this taxon. Therefore, we select Oplomus festivus Dal- las, 1851, as the replacement name for Pen- tatoma marginale Westwood, 1837. II. DISCOCEPHALINAE: OCHLERINI Stalius castaneus (Distant, 1893), NEW COMBINATION Melanodermus castaneus Distant, 1893: AS: This species was omitted from the junior author’s (Rolston 1992) revision of Ochler- ini. The species is known only from the fe- male holotype, which was collected in Nic- aragua. It is morphologically similar to Stalius tartareus (Stal) excepting the form of the basal plates, and these differ consid- erably (Figs. 1—2). III. PENTATOMINAE Aelia chinensis, NEW NAME Aelia bifida Hsiao and Cheng, 1978: 326, a2] SZOnnes. 4..5.. 10,413. [mneteCosta: 1847, Neottiglossa] Costa (1847) described, as Aelia bifida, a relatively common _ circum-Mediterranean species now in the genus Neottiglossa Kir- by. The primary homonymy resulting from Hsiao and Cheng’s (1978) description of Aelia bifida as a new species necessitates the above replacement name. Aelia chinen- sis occurs in China and Inner Mongolia. Aeliomorpha viridescens, NEw NAME Aeliomorpha (Distantiella) viridis Azim and Shafee, 1987: 422-424, figs. E-H. [not Reuter, 1887, Aeliomorpha| Reuter (1887) described Aeliomorpha viridis from Madagascar. Although Azim and Shafee (1987) placed their Indian A. viridis in a different subgenus, it is still a primary homonym and requires a replace- ment name. Amyntor unicolor (Walker, 1867), REVISED STATUS Halys (Dichelops) obscura Dallas, 1849: 188, pl. 19 fig. 3. [not Westwood, 1837, Sarju; not Herrich-Schaffer, 1839, Bro- chymena parva| Amyntor obscurus: Stal, 1867: 519; Stal, 1876: 107; Atkinson, 1888b: 154; Leth- ierry and Severin, 1893: 183; Distant, 1902: 144-145, fig. 85; Kirkaldy, 1909: 147; Zhang, 1985: 80-81, pl. 42 fig. 122; Hua, 1989: 43. Bolaca unicolor Walker, 1867: 251; Atkin- son, 1888a: 70; Lethierry and Severin, 1393: 161. Amyntor unicolor: 431. Distant, 1900a: 425, The binomen, Halys obscura, has been used on at least three separate occasions to represent three different species. Westwood (1837) first used the name for a species now contained in the halyine genus Sarju. The second species, described by Herrich- Schaffer (1839), now belongs in the halyine genus Brochymena, and was given the new name B. parva by Ruckes (1947). Finally, Dallas (1849) used Halys obscura to rep- resent a species now residing in the genus Amyntor. The next available name is uni- color Walker, 1867; the proper combina- tion, Amyntor unicolor, has been used only once (Distant 1900a). Antestia dollingi, NEW NAME Cimex adspersus Fabricius, 1803: 175. [not Thunberg, 1784: 53, Coreidae] Antestia adspersa: Stal, 1868: 34. VOLUME 97, NUMBER 4 Thunberg’s (1784) usage of the binomen Cimex adspersus easily predates Fabricius’ (1803) usage. Cimex adspersus Thunberg is probably a member of the family Coreidae. There are no currently known junior syn- onyms of Cimex adspersus Fabricius, 1803. Berecynthus hastator (Fabricius, 1798), REVISED STATUS Cimex delirator Fabricius, 1794: 103. [not Fabricius, 1787, Coreidae] Cimex hastator Fabricius, 1798: 532. Berecynthus delirator: Stal, 1868: 26. Fabricius described two different species under the binomen Cimex delirator. The first (1787) is a member of the family Co- reidae and a junior synonym of Zicca ni- gropunctata (DeGeer); the second (1794) is a neotropical pentatomid. The next avail- able name for this neotropical pentatomid is Cimex hastator Fabricius, 1798. Cosmopepla lintneriana Kirkaldy, 1909 Cimex carnifex Fabricius, 1798: 535; Co- quebert, 1801: 81; Fabricius, 1803: 117. [not Fabricius, 1775, Lygaeidae] Pentatoma bimaculata Thomas, 1865: 455. [not Montrouzier, 1855: 98, Hyrmine] Cosmopepla lintneriana Kirkaldy, 1909: 80 [New name for Pentatoma bimaculata Thomas, 1865]; McDonald, 1986: 4, 5, figs. 15-27. Cosmopepla bimaculata: Van Duzee, 1917: 49; Stoner, 1920: 96—97; Blatchley, 1926: 152-153; Leonard, 1928: 82; McDonald, 1966: 25, 51; McPherson, 1982: 73-74; Froeschner, 1988: 574. Kirkaldy (1909) considered the name Pentatoma bimaculata Thomas, 1865, to be preoccupied, but did not indicate the older synonym. He proposed the new name Cos- mopepla lintneriana. No one followed Kir- kaldy’s recommendation except McDonald (1986) who indicated that P. bimaculata Thomas, 1865, was preoccupied by P. bi- maculata Westwood, 1837: 8, 35. Froesch- ner (1988) argued correctly that P. bima- culata Westwood, 1837, was in fact a no- 847 men nudum. However, Montrouzier (1855) (see Hyrmine sexpunctata bimaculata, be- low) also described a P. bimaculata, pre- dating Thomas’ description by ten years. The next available name for Cimex carnifex Fabricius, 1798, is Cosmopepla lintneriana as was proposed by Kirkaldy (1909). Dictyotus walkeri, NEW NAME Pentatoma aequalis Walker, 1867: 310— 311. [not Say, 1831, Hymenarcys ae- qualis| Dictyotus aequalis: Distant, 1900a: 388. Say (1831) described as Pentatoma ae- qualis a North American species currently in the genus Hymenarcys Amyot and Ser- ville. Later, Walker (1867) also described as P. aequalis an Australian species now placed in the genus Dictyotus Dallas. There is no recognized junior synonym of Walk- er’s binomen to serve as the valid name. We therefore propose Dictyotus walkeri, new name, for Pentatoma aequalis Walker. Halyomorpha hasani, NEW NAME Halyomorpha punctata Hasan, 1993: 210, 214-215, figs. SA—H. [not Cachan, 1952, Halyomorpha| Cachan (1952) described as Halyomor- Pha punctata a Madagascaran species which is still regarded as a valid species in Halyomorpha Mayr. Hasan (1993) appar- ently overlooked this earlier description in describing his new species, Halyomorpha punctata; thus necessitating the above new name. Holcostethus Fieber, 1860 Holcostethus Fieber, 1860: 79 [nomen nu- dum]; Fieber, 1861: 333. Peribalus Mulsant and Rey, 1866: 237, 262. Dryocoris Mulsant and Rey, 1866: 237, 267. Ribes and Schmitz (1992) recently divid- ed the well-known genus Holcostethus Fie- ber into two genera, based primarily on the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ngth and shape of the juga. Those species in which the juga do not meet in front of ihe tylus were retained in Holcostethus, while the remaining species were placed in Dryocoris Mulsant and Rey. Not only is the use of the generic name Dryocoris in error, but we also believe there is little merit in splitting the genus based upon the form of the juga. Dryocoris was first proposed by Amyot (1845) in his mononomial system to rep- resent the single species, Cimex sphacelatus Fabricius, 1794. Ribes and Schmitz (1992) were mistaken in their claim that Amyot (1845) initially included three species in Dryocoris. Included within the synonymy of Dryocoris were sphacelatus Fabricius, 1794; vernalis Wolff?, 1804; and baccarum Amyot and Serville, 1843. Amyot (1845) tentatively (note question mark) placed ver- nalis as a junior synonym of sphacelatus, and baccarum Amyot and Serville is not an original description. The inclusion of bac- carum Amyot and Serville indicates that Amyot believed Amyot and Serville had misidentified the true baccarum Linnaeus, 1758 (a member of the genus Dolycoris). One problem lies in the fact that apparently Amyot (1845) based his Dryocoris on a misidentified type species. He states that Dryocoris is similar to Pentatoma Olivier, except that the head is larger and more rounded, and the lateral lobes meet beyond the median lobe. The true sphacelatus has the jugal lobes shorter, not meeting beyond the tylus. At any rate, this initial use of Dry- ocoris is invalid because the entire work (Amyot 1845) has been officially placed upon the list of rejected works because it is largely mononomial. The first valid use of Dryocoris in a bi- nomial is generally credited to Mulsant and Rey (1866), who used the name in place of Holcostethus Fieber, 1861, apparently rec- ognizing Amyot’s prior use of Dryocoris. Mulsant and Rey (1866) followed Amyot (1845) in including only one species within Dryocoris, that being Cimex sphacelatus Fabricius, 1794. Dryocoris should be cred- ited to Mulsant and Rey, 1866, with Cimex sphacelatus Fabricius, 1794, the type spe- cies by monotypy. Holcostethus was first described by Fie- ber in 1860 when it was included in a key to genera; no species were included, how- ever, so this use is a nomen nudum. He re- described Holcostethus in 1861, and includ- ed three species: jani Fieber, sphacelatus (Fabricius), and congenor Fieber. Although he (1861) did not designate a type species for Holcostethus, Mulsant and Rey (1866) effectively fixed the type for both Dryocor- is and Holcostethus when they treated the two names as synonyms, and included only sphacelatus within Dryocoris. Most recent workers have considered sphacelatus as the type species of Holcostethus (Kirkaldy 1909, McDonald 1974, Froeschner, 1988). Dryocoris is a junior synonym of Holcos- tethus. Mulsant and Rey (1866) also described Peribalus, placing in it three species: Cimex vernalis Wolff, Cimex distinctus Fieber, and Pentatoma inclusus Dohrn. Stal (1872a) considered Dryocoris to be a junior syn- onym of Peribalus. This has led most re- cent workers to consider both Dryocoris and Peribalus as junior synonyms of Hol- costethus. Kirkaldy (1909) fixed vernalis as the type species of Peribalus. If Holcostethus is divided into two gen- era, based on the form of the juga, Peri- balus should be used for those species in which the juga are contiguous anterior to the tylus. We believe, however, that the ge- neric separation based upon this character is unwarranted. The form of the juga in some species is quite variable with different individuals exhibiting either character state. Holcostethus should remain as the valid name for the genus with Dryocoris and Per- ibalus as junior synonyms. Holcostethus macdonaldi, NEW NAME Pentatoma picea Dallas, 1851: 236. [not Palisot de Beauvois, 1817, Antiteuchus, (Discocephalinae)] Peribalus piceus: Uhler, 1886: 7; Gillette VOLUME 97, NUMBER 4 and Baker, 1895: 16; Van Duzee, 1904: 34-eeVan'Duzee) 1917: 33; Blatchley, 1926: 106. Holcostethus piceus: Kirkaldy, 1909: 48; McDonald, 1974: 247, 252, figs. 36-43; McDonald, 1982: 5; McPherson, 1982: 48, 50; Froeschner, 1988: 581—582. Pentatoma picea Palisot de Beauvois, 1817, easily predates Dallas’s usage of the name. Palisot de Beauvois’s species is a valid species in the discocephaline genus Antiteuchus Dallas. There is no available junior synonym for P. picea Dallas, 1851; therefore, we propose Holcostethus mac- donaldi, new name. Hyrmine sexpunctata bimaculata (Montrouzier, 1855) Pentatoma bimaculatum Montrouzier, 1855: 98. Hyrmine 6-punctata var. montrouzierana Kirkaldy, 1909: 115. [Unnecessary new name for Pentatoma bimaculatum Mon- trouzier, 1855] Kirkaldy (1909) apparently believed Pentatoma bimaculatum Montrouzier, 1855 to be preoccupied by P. bimaculata West- wood (1837). As Froeschner (1988) argued, P. bimaculata Westwood, 1837, is a nomen nudum appearing in a list on page 8, and a simple note that it should be deleted as a variety of P. obscurus [the preceding spe- cies] on page 35. Keleacoris, NEW NAME Kelea Schouteden, 1958: 128—130; Linna- vuori, 1982: 153. [not Merrem, 1818, Aves] Schouteden’s (1958) usage of the generic name Kelea is preoccupied by the Avian genus Kelea Merrem, 1818. There is no available junior synonym, therefore we pro- pose Keleacoris, new name. Keleacoris is currently monotypic, containing only K. congolensis (Distant, 1910), new combina- tion. 849 Menida signoreti, NEW NAME Rhaphigaster parvulus Signoret, 1858: 289. [not Dallas, 1851, Acrosternum| Antestia parvula: Stal, 1865: 211; Walker, 1867: 281. Menida parvula: Stal, 1876: 99; Lethierry and Severin, 1893: 174; Kirkaldy, 1909: 1537 Linnavuorl, 1982-158, 160) figs! 258e, ty 2olc; Linnavuon, 1986: 131; 135, figs. 5a, 12c—f, 13a-f. Eurymenida parvula: Ahmad and Moham- mad, 1982: 12. Dallas (1851) described Rhaphigaster parvulus, which now belongs in the genus Acrosternum. Thomas and Yonke (1990) speculated that it may actually be a syn- onym of the African species, A. heegeri Fieber. Because there are no available ju- nior synonyms for R. parvulus Signoret, 1858, we propose Menida signoreti as a re- placement name. Menida signoreti is dis- tributed in the tropical areas of west Africa (Cameroon, Guinea, Liberia, Ivory Coast, Nigeria). Neococalus germari, NEW NAME Cimex leucogrammus Germar, 1838: 179- 180. [not Gmelin, 1790: 2131, Ancyro- soma (Podopinae); not Gmelin, 1790: 2165, Miridae] Sciocoris leuacogrammus: Herrich-Schaffer, 1844: 788-89, fig. 756; Dallas, 1851: 134; Walker, 1867: 174. Cocalus leucogrammus: Stal, 1861: 200; Stal, 1865: 119-120; Stal, 1876: 54; Reu- ter, 1884: 6; Lethierry & Severin, 1893: 11; Distant, 1898: 316; Distant, 1901: 25; Schouteden, 1910: 80, Schouteden, 1912: 107; Villiers, 1952: 303, Villiers, 1954: 916. Neococalus leucogrammus: Linnavuori, 1975: 31, 32, figs. 14c, 17bd; Linnavuori, 1982: 75, figs. 85b, 86d. Gmelin (1790) actually used the binomen Cimex leucogrammus twice in his 13th edi- tion of Systema Naturae. First, he proposed Cimex leucogrammes as a replacement PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘me for Cimex albolineatus Fabricius, (781, which was preoccupied by Cimex al- bolineatus Goeze, 1778. Goeze’s taxon 1s now a species in the Miridae; Fabricius’ and Gmelin’s taxon is a member of the po- dopine genus Ancyrosoma Amyot & Ser- ville. Second, he used Cimex leucogrammus as the name for a new species of Miridae. Germar (1838) was apparently unaware of Gmelin’s earlier uses of this binomen. Distant (1898) placed Sciocoris clausus Walker, 1867, in the synonymy of Neoco- calus leuacogrammus. All modern workers, however, have consider N. clausus and N. leucogrammus as distinct species (Leston 1952, 1953; Schouteden’ 1957, 1963, Lin- navuori 1975, 1982). There are no other known junior synonyms of N. leucogram- mus, thus necessitating the above new name. Piezodorus flavulus (Stal, 1853), REVISED STATUS Cimex pallescens Germar, 1838: 175. [not Gmelin, 1790, Miridae; not Donovan, 1794, Miridae] Rhaphigaster pallescens: Herrich-Schiaffer, 1845: 7, 11-12; Walker, 1867: 363. Rhaphigaster flavulus Stal, 1853: 221. Nezara (Piezodorus) pallescens: Stal, 1865: 198. Piezodorus pallescens: Stal, 1876: 100; Lethierry and Severin, 1893: 176; Kir- kaldy, 1909: 136; Linnavuori, 1975: 123; Linnavuori, 1982: 147-148, figs. 235c, 2360, 2372. The binomen Cimex pallescens has been used at least three separate times for the descriptions of three different heteropteran species. The first two (Gmelin 1790, Don- ovan 1794) are both members of the family Miridae, and both predate Germar’s (1838) usage of the binomen for a pentatomid now placed in the genus Piezodorus Fieber. The next available synonym is Rhaphigaster fla- vulus Stal, 1853. Sciocoris sahelensis, NEW NAME Sciocoris australis Linnavuori, 1975: 26— 27, figs. 12a, b; Linnavuori, 1982: 65, 66, figs. a, b. [not Dallas, 1852, Eribotes] Linnavuori (1975) apparently overlooked Dallas’s (1852) original usage of the bino- men Sciocoris australis when he described a new species by that name from Sudan. Dallas’s species, described from Australia, now resides in the genus Eribotes Stal. There are no known junior synonyms of Linnavuori’s species; therfore, we propose Sciocoris sahelensis as a replacement name. Thyanta humilis Bergroth, 1891 Pentatoma patruelis Stal, 1859: 226—227; Walker, 1867: 289. [not Stal, 1853, Eu- dryadocoris goniodes}| Thyanta patruelis: Stal, 1862: 58; Stal, 1872b: 35; Lethierry and Severin, 1893: 148; Kirkaldy, 1909: 95. Thyanta humilis Bergroth, 1891: 225-226. Thyanta (Argosoma) patruelis: Rider and Chapin, 1991: 5, 6, 37-39, figs. 184-198. Stal (1853) described Pentatoma patruel- is, which is now a junior synonym of the African Eudryadocoris goniodes Dallas. He (1859) later described as P. patruelis a South American species that is now placed in the genus Thyanta Stal. The next avail- able name is Thyanta humilis Bergroth, 1891. Trincavellius Distant, 1900 Trincavellius Distant, 1900b: 163. Paposia China, 1962: 57-59. New synon- ymy. Examination of Distant’s (1900b) and China’s (1962) descriptions, and included illustrations, leaves litthe doubt that Trin- cavellius Distant and Paposia China are synonyms. Although two of the three taxa belonging in Trincavellius were originally described in Sciocoris Fallén, this genus has generally been included within the nomi- nate tribe of the Discocephalinae, rather than the pentatomine tribe Sciocorini. This VOLUME 97, NUMBER 4 is probably because of the rather convex ab- dominal venter that differs from most scio- corines. Other characters (shape of head, explanate anterolateral pronotal margins), however, are characteristic of the Sciocori- ni. Additionally, most typical discocepha- line characters are lacking. Trincavellius should, at least tentatively, be placed within the Sciocorini. At present, three specific taxa belong in Trincavellius, all of which are probably conspecific. We have examined a number of specimens from various locations in Chile and Peru, as well as a male syntype of Scio- coris kingi Reed. All specimens examined are conspecific and differ in no appreciable manner from any of the species descriptions given by Butler, 1877 (galapagoénsis), Reed, 1898 (kingi), or China, 1962 (rucke- si). Any synonymization of specific names, however, should wait until appropriate type material can be examined. IV. PHYLLOCEPHALINAE Basicryptus Herrich-Schaffer, 1844 and Dalsira Amyot and Serville, 1843 Dalsira Amyot and Serville, 1843, orig- inally contained two species, both new and both eligible as type species: marginata and affinis. Stal (1876) left only one of these two species in Dalsira by transferring mar- ginata to Basicryptus Herrich-Schaffer, 1844 (type species Cimex costalis Germar, 1838, by monotypy). Kirkaldy (1909) cre- ated lasting confusion by selecting margin- ata as the type species of Dalsira and cre- ating a new generic name, Metonymia (type species Dalsira affinis Amyot and Serville, 1843, by original designation), overlooking the fact that Distant (1902) had earlier fixed affinis as the type species of Dalsira. The effect of Kirkaldy’s invalid action was to replace the name Basicryptus with Dalsira, and to apply the name Metonymia to those species previously placed in Dalsira. How- ever, since both Metonymia and Dalsira have the same type species, Metonymia is an objective junior synonym of Dalsira, 851 while Dalsira as erroneously used by Kir- kaldy is synonymous with Basicryptus. The genera Basicryptus and Dalsira need revising, and the two lists that follow are based on literature. These species and subspecies appear to belong in Basicryptus: 1. albidicosta (Walker, 1868) 2. angulatus Schouteden, 1909 3. antennatus Distant, 1892 4. atricostata (Distant, 1910) 5. bohndorffi Distant, 1890 6. brunneus Jensen-Haarup, 1931 7. costalis (Germar, 1838) [type sp.] 8. costalis ugandana (Linnavuori, 1982), new combination 9. distinctus (Signoret, 1851) 10. eburnea Jeannel, 1913 11. elongata Distant, 1892 12. frenchi Bergroth, 1895 13. gibbosa (Dallas, 1851) 14. hutereaui Schouteden, 1916 15. irroratus (Westwood, 1837) 16. maindroni Jeannel, 1913 17. marginatus (Amyot and Serville, 1843) 18. masaicus Jeannel, 1913 19. nigrocinctus Jeannel, 1913 20. nigromaculatus Schouteden, 1904 21. pictus Schouteden, 1916 22. plicatus (Reiche and Fairmaire, 1847) 23. projectus Distant, 1898 24. rugicollis (Westwood, 1837) 25. rugosus (Fabricius, 1803) 26. striatus (Linnavuori, 1982), new com- bination 27. strigosa (Burmeister, 1835) 28. subtruncatus (Walker, 1868) 29. upembanus (Linnavuori, 1982), combination new These species appear to belong in Dal- Sira: 1. affinis Amyot and Serville, 1843 [type sp.] 2. alata (Distant, 1898) 3. angolana (Schouteden, 1962) 4. asperata Distant, 1889 5. atricostata Distant, 1910 6. bigemmis Bergroth, 1891 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 7. brunni Schouteden, 1904 8. crassa Distant, 1898 9. dallasi (Schouteden, 1912), new com- bination 10. humeralis (Walker, 1868) 11. icterica (Gerstacker, 1892) 12. kocki (Schouteden, 1962), new combi- nation 13. lentiginosa (Stal, 1852) 14. longiceps (Schouteden, 1910) 15. mabokeana (Linnavuori, 1982), combination 16. maura Distant, 1898 17. modesta (Fabricius, 1803) 18. mulunguana (Linnavuori, 1982), new combination 19. niemboana_ (Linnavuori, combination 20. otjimbora (Hesse, 1925) 21. overlaeti (Schouteden, 1962) 22. scabrata Distant, 1901 23. wagneri Leston, 1952 new 1982), new ACKNOWLEDGMENTS We are sincerely grateful to all of those who have provided help in tracking down much of the literature needed for this paper and for the upcoming pentatomid catalog. We thank Mrs. J. Margerison-Knight, The Natural History Museum, for the opportu- nity to examine the type specimen of Me- lanodermus castaneus. We also thank Ger- ald Fauske, North Dakota State University, for his helpful comments on an early ver- sion of the manuscript. LITERATURE CITED Ahmad, I. and F A. Mohammad. 1982. A new species of the genus Eurymenida Berg. (Pentatomidae: Pentatominae: Menidini) from Pakistan with notes on their relationships. Proceedings of the Ento- mological Society of Karachi 11-12: 7-12. (1981.) Amyot, C. J. B. 1845. Entomologie Frangaise. Rhyn- chotes. Hémipteres. Homoptéres. Phthiroptéres. Pseudopteres. Méthode mononymique. Annales de la Société Entomologique de France 14: 369— 492 + 8-9 pls. Amyot, C. J. B. and J. G. A. Serville. 1843. Histoire Naturelle des Insectes. Hémipteres. Librairie En- cyclopédique de Roret, Paris, Ixxvi + 681 pp. + 12 pls. Atkinson, E. T. 1888a. Notes on Indian Rhynchota: Heteroptera, No. 3. 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Revision of the genus Holcostethus in North America (Hemiptera: Pentatomidae). Jour- nal of the New York Entomological Society 82: 245-258. 1982. Description of the male genitalia of Holcostethus hirtus (Van Duzee) with a revised key to North American species (Hemiptera: Pen- tatomidae). Journal of the New York Entomolog- ical Society 90: 5—7. . 1986. Revision of Cosmopepla Stal (Hemip- tera: Pentatomidae). Journal of the New York En- tomological Society 94: 1-15. 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 and Edwardsville. 240 PP: Merrem, 1818. /n Ersch and Gruber, eds., Allgemeine Encyklop. Wissenschaften Kiinste, Section 1 (1), p. 338. Montrouzier, P. 1855. Essai sur la faune de I’Ile de Woodlark ou Moiou. d’ Agriculture, Sciences et Industries de Lyon (2)7: 1-114. Mulsant, E. and C. Rey. 1866. 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Chapin. 1991. Revision of the genus Thyanta Stal, 1862 (Heteroptera: Pentatom- idae) I. South America. Journal of the New York Entomological Society 99: 1-77. Rolston, L. H. 1992. Key and diagnoses for the genera of Ochlerini (Hemiptera: Pentatomidae: Discoce- phalinae). Journal of the New York Entomological Society 100: 1-41. Rolston, L. H. and E J. D. McDonald. 1981. Con- spectus of Pentatomini genera of the Western Hemisphere—Part 2 (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 88(4): 257-272 (1980.) Ruckes, H. 1947. Notes and keys on the genus Bro- chymena (Pentatomidae, Heteroptera). Entomolo- gica Americana 26: 143-238. Say, T. 1831. Descriptions of new species of heter- opterous Hemiptera of North America. New Har- mony Press, Indiana, 39 pp. Schouteden, H. 1910. Wissenschaftliche ergebnisse der schwedischen zoologischen expedition nach dem Kilimandjaro, dem Meru und den Umgeben- den Massaisteppen Deutsch-Ostafrikas 1905—1906 unter leitung von Prof. Dr. Yngve Sjéstedt. 12. Hemiptera. 6. Pentatomidae. /n Sjéstedt’s Kili- mandjaro-Meru Expedition. P. Palmquists Aktie- bolag, Stockholm 12: 73-96. . 1912. Cimicidae, Coreidae, Reduviidae de la Region du Zoutpansberg (Transvaal). Revue Zool- ogique Africaine 2(1): 101-114. . 1957. Contributions a I’ étude de la faune en- tomologique du Ruanda-Urundi (Mission P. Bas- ilewsky 1953). CXXVI. Heteroptera, Platispidae, Acanthosomatidae, Pentatomidae et Coreidae. An- VOLUME 97, NUMBER 4 nales du Musée Royal du Congo Belge 58: 269— 310. 1958. Pentatominae novae africanae: tl. = Kelea Schout. et Euryaspis Sign. Revue de Zool- ogie et de Botanique Africaines 48: 129-143. 1963. Pentatomides de la Céte d’Ivoire—I. Revue de Zoologie et de Botanique Africaines 68(3-4): 397-402. Signoret, V. 1858. 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STAINES 3302 Decker Place, Edgewater, Maryland 21037. Abstract.—A biographical sketch, bibliography, and list of proposed taxa are presented for E A. Monros. Key Words: Francisco Monros was one of the most prolific chrysomelid taxonomists of the middle 1900’s. His research interests cov- ered most of the family, they have contrib- uted greatly to chrysomelid systematics, es- pecially many of the smaller subfamilies. His monographs on various subfamilies for Argentina are still standard works. Little of his work has been tested by modern system- atic techniques, so that his longterm impact is not known. BIOGRAPHICAL SKETCH Francisco Monros was born in Barce- lona, Spain on 6 June 1922. He began his entomological studies at the Barcelona Mu- seum under Francisco Espanol Coll. In 1938 Monros, his parents and brother left Spain due to the Civil War and arrived in Argentina, where he began his career as an agronomy engineer with the Faculty of Agronomy of the National University of Buenos Aires. In 1948, Monroés began his association with the Miguel Lillo Institute of the Na- tional University of Tucuman. During his tenure in Tucuman he held the following positions: Interim Director of the Entomol- ogy Institute, Technical Secretary of the Academy of Biological Sciences, Vice Dean, as well as various professorships. In 1950 Monr6s was awarded the Juana Petrocchi Award from the Argentine As- Monros, biography, bibliography, proposed taxa sociation for the Progress of the Sciences for his work “‘Revision Sistematica de los Hispidae Argentinos.” In 1952-1953 the John Simon Guggenheim Memorial Foun- dation awarded Monros a scholarship to visit the principal museums in the United States. A similar scholarship was awarded in 1955—1956 by the government of France, which allowed study in the Natural History Museum of Paris, the British Museum, the Frey Museum in Munich, and other muse- ums in Europe. Blake (1958) and Wygodzinsky (1959) wrote obituaries which talk of Monros as a person rather than a scientist. The picture which emerged was of a modest, self as- sured, quiet person with high ideals, who handled his responsibilities efficiently. He did, however, have definite scientific opin- ions which were expounded in the appro- priate arena. Monro6s was a keen observer of natural history. He kept field notes and sketches from his field trips. His lectures were pop- ular and interesting and were usually ac- companied by his ambidextrously executed drawings. Monros published 67 articles, all but five on chrysomelids. His interests covered the entire family but he did not publish on the subfamily Galerucinae. Most of his works are illustrated with his own drawings. Mon- ros was interested in geographical variation VOLUME 97, NUMBER 4 of species as many of his papers discuss variation in detail. Monros amassed a large collection through semiannual collecting trips and ex- changes with other museums. His personal collection, including his types, is now held by the United States National Museum of Natural History (Blake 1961). Much Mon- rds material is also present in the collection of the Instituto Fundaci6n Migel Lillo of Tucuman University (Willink per. comm). Monros died on 3 May 1958. He was survived by his wife, the former Maria Muntanola, his daughter Silvia, and his brother Antonio. BIBLIOGRAPHY The author is EK Monros unless otherwise specified. The numbering system is_ that used by Wygodzinsky (1959). Both Halffter (1958) and Wygodzinsky (1959) mentioned that Monroés had the Clytrinae part of the Coleopterorum Catalogus Supplementa in press, but to my knowledge this part was never published. 1943 1. La familia Sagridae en Sud America: El genero Atalasis Lac. (Col., Chrysomelidae). Revista de la Sociedad Entomologica Argentina 11(5): 411-— 422, 3 figs., 1 pl. 1944 2. Algunos cole6pteros de interés forestal observados en la Isla Victora (Gobernacion del Neuquen). Re- vista de la Facultad de Agronomia y Veterinaria Buenos Aires 10(3): 537-543, 2 figs. 3. Archiopactus bruchi Heller = Archiopactus ni- veopectus Hust. Notas sinonimica. Revista de la Sociedad Entomologica Argentina 12(3): 181- 183, 3 figs. 4. Descripcion de un nuevo clitrido argentino (Col., Chrysomeloidea). Revista de la Facultad de Agronomia y Veterineria Buenos Aires 11(1): 148-152, 1 fig. 1945 5. Tres interesantes confusiones en Chrysomeloidea neotropicales. Revista de la Sociedad Entomolo- gica Argentina 12(5): 410-415, 2 figs. 6. A propoésito de algunos Chrysomeloidea neotrop- icales (Coleoptera). Notas del Museo de La Plata 10: 143-155, 5 figs. 16. IZ 19: 20. 857 1947 . Revision de los Megalopidae Argentinos (Col. Chrysomelidae). Revista de la Sociedad Entomo- logica Argentina 13: 150—217, 51 figs., 5 pls. . Revision del género Plectonycha Lac. (Col. Chrysomelidae). Anales de la Sociedad Cientifica Argentina 144: 46-64, 10 figs. . Notas s6bre Crioceridae argentinos (Col. Chryso- meloidea). Arthropoda 1(1): 78-88, 9 figs. . Descripciones y comentaries relativos a ‘‘Chry- someloidea”’ (Coleoptera). Acta Zoologica Lil- loana 4: 155-174, 2 figs, 1 pl. . Revision sistematica de los Hispidae argentinos (Insecta, Coleop., Chrysomeloid.). Anales del Mu- seo Argentino Ciencias Naturales “‘Bernardino Ri- vadavia” 42: 125-324, 10 figs., 27 pls. [with M. J. Viana]. 1948 . Descripcion de diez nuevas especies de “‘Lam- prosoma”’ neotropicales (Col., Chrysomelidae). Acta Zoologica Lilloana 5: 81—95, 3 figs. . Notas sébre “‘Eupoda”’ Argentinos (Col., Chrys- omelidae). Acta Zoologica Lilloana 5: 203—220, 9 figs. . Descripcion de diez nuevos “‘Camptosoma’”’ neo- tropicales (Col., Chrysomelidae). Acta Zoologica Lilloana 6: 171—200, 38 figs. 1949 . Descripcion de seis nuevas Chlamisinae neotrop- icales (Col., Chrysomelidae) (2a contribuci6n al conocimiento de Chlamisinae). Revista de Ento- mologia, Rio de Janeiro 20(1—3): 617-629, 14 figs. Descripcion de las metamorfosis de *‘Lamproso- ma chorisiae’’ Monros y consideraciones taxon- omicas sdébre “‘Lamprosominae’’ (Col. Chryso- melidae). Acta Zoologica Lilloana 7: 449-466, 12 figs., 1 pl. El género ‘‘Mylassa”’ Stal (Col., Chrysomelidae, Cryptocephalinae). Acta Zoologica Lilloana 7: 489-525, 27 figs. . SObre la posicion sistematica de algunos ‘‘Eupo- da”’ dudosos (Col., Chrysomelidae). Acta Zoolo- gica Lilloana 7: 545-574, 32 figs. Revision de las especies Argentinos de “‘Dory- notini’’ (Col. Cassidinae) (la contribuci6n al con- ocimiento de ‘‘Cassidinae’’). Acta Zoologica Lil- loana 8: 391-426, 14 figs. [with M. J. Viana] 1950 Descripciones y comentarios s6bre ““Chlamisinae”’ neotropicales (Coleoptera, Chrysomelidae). Re- vista Brasileira de Biologia 10(4): 409-424, 24 figs. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1951 Notes on chrysomelid beetles of the subfamily Chlamisinae, with descriptions of new species. Proceedings of the United States National Muse- um 101(3283): 451-463, figs. 85—90. . Diez nuevas espécies de Chlamisus neotropicales (Col., Chrysomelidae) (6a contribuci6n al cono- cimiento de Chlamisinae). Dusenia (Curitiba) 2(4): 255-272, 19 figs. 23. Nuevas especies de “‘Euryscopa’”’ Sudamericanas 27. Consideraciones sObre 28. 29) a2: 34. ee) Nn 36. (Col., Chrysomelidae, Clytrinae). Acta Zoologica Lilloana 9: 293-305, 9 figs. . Descripci6n de diez especies nuevos de Babiini neotropicales (Col. Chrys.). Revista de la Socie- dad Entomologia Argentina 15: 149-161, 12 figs. . Descriptions of new Neotropical Clytrinae (Col., Chrysomelidae). Annals and Magazine of Natural History (12)4: 1146-1158, 13 figs. . Nuevo género de Megascelinae de la Patagonia. Revista Chilena de Entomologia 1: 171-173, 2 figs. ““Leasia australis” Jac. (Coleoptera). Revista Brasileira de Biologia 11(4): 451-456, 13 figs. 1952 Diez nuevas Babiini Sudamericanas (Col., Chrys- omelidae, Clytriniae). Dusenia (Curitiba) 3(2): 118-130, 10 figs. Las Cassidinae de la Seccion ‘‘Hemisphaerotina”’ con revision de las especies Argentinos (Col. Cas- sidinae) (2a contribucion al conocimiento de Cas- sidinae). Acta Zoologica Lilloana 11: 367-395, 8 figs. [with M. J. Viana]. . Notas s6bre Criocerinae. Acta Zoologica Lilloana 11: 467-482, 6 figs. . Las especies argentinas de Cupedidae (Coleop- tera). Anales de la Sociedad Cientifica Argentina 154: 19-41, 30 figs. [with M. M. de Monrés]. Notas sdbre algunas Eumolpinae neotropicales (Coleoptera Chrysomelidae). Revista Chilena de Entomologia 2: 187—196, 5 figs. . Seis nuevas especies de Criocerinae del Noreste Argentino (Col. Chrysomelidae). Acta Zoologica Lilloana 10: 33-49, 29 figs. Revision de las especies argentinas de Chlamisi- nae (5a contribuci6n al conocimiento de Chlami- sinae). Acta Zoologica Lilloana 10: 489-672, 177 figs. . Descripciones de “‘Clytrinae’’ Americanas (Col., Chrysomelidae). Revista Brasileira de Biologia 12(4): 349-362, 17 figs. 1953 Aulacoscelinae, eine neue Chrysomeliden-Unter- familie, mit Beschreibung einer neuen bolivian- Sis 38. 39), 40. 41. 42. 44. ischen Gattung (Col.). Entomologische Arbeiten aus dem Museum G. Frey 4: 19-25, 7 figs. Some corrections in the nomenclature of Clytrinae (Chrysomelidae). Coleopterists Bulletin 7(6): 45— 50. 1954 Notes and synonyms in Chrysomelidae (Coleop- tera). Proceedings of the Entomological Society of Washington 56(1): 23-26, 3 figs. Some new synonyms in Clytrinae (Coleoptera: Chrysomelidae). Psyche 60: 148-150. Revision sistematica de las especies de “‘Clytri- nae” de la Argentina, Paraguay, Uruguay y Chile (Col., Chrysomelidae). Acta Lilloana 14: 5—274, 404 figs. Megalopus jacobyi, nueva plaga de Solanaceas en el Noroeste Argentino, con notas sobre biolo- gia y taxonomia de Megalopinae (Col., Chryso- melidae). Revista Agronomica Noreste Argentino 1(2): 167-179, 19 figs. Revision of the chrysomelid subfamily Aulacos- celinae. Bulletin of the Museum of Comparative Zoology 112(4): 321-360, 77 figs. . Descripciones y comentarios s6bre Chlamisinae (7 a contribucién) (Col. Chrysomelidae). Acta Zoologica Lilloana 13: 77—95, 29 figs. General remarks on the subfamily Clytrinae, as suggested by the second edition of the Coleopter- orum Catalogus (Chrysomelidae). Coleopterists Bulletin 8(3—4): 62, 68. 1955 . Remarques sur les affinités des familles de Cer- ambycoidea. Bulletin de I’Institut Royal des Sci- ences Naturelles de Belgique 31(31): 1-7, 2 figs. 46. Biologia y descripcion de la larva de Atalasis sagroides (Col., Chrysomelidae). Revista Agron- Omica Noroeste Argentino 1(3): 275-281, 9 figs. 47A. Book review- Insects of Micronesia by J. Linse- ly Gressitt. Ciencia e Investigacion 11: 267. 47B. Book review- Insects: The Yearbook of Agricul- 48. 49. Syl ture. Revista Agronoémica del Noroeste Argentino 1(2): 220. On some new genera of nearctic Chrysomelinae (Chrysomelidae). Coleopterists Bulletin 9(4): 53— 60, 34 figs. 1956 Sur le genre Megamerus MacLeay [Col. Chrys- omelidae]. Revue Frangaise d’Entomologie 23(2): 104-115, 13 figs. . Revision generica de Lamprosominae con descrip- cion de algunos géneros y especies nuevas (Col., Chrysomelidae). Revista Agronémica Noroeste Argentino (Tucuman) 2(1): 25-77, 62 figs. Sur quelques Clytrinae du Muséum de Paris (Col., VOLUME 97, NUMBER 4 Nn WwW 54. ae 59: 60. 61. 62. 66. Chrysomelidae). Revue Frangaise d’Entomologie 23(3): 161-164, 4 figs. . Notas s6bre Criocerinae del subgénero Guasilema (Col. Chrysomelidae). Revista de la Sociedad En- tomologia Argentina 18: 35-44, 16 figs. . Uber einige verkannte Chrysomeliden-Namen. Entomologische Arbeiten aus dem Museum G. Frey 7(3): 1118-1137. [with J. Bechyné]. 1957 Las especies Lema del subgénero Pachylema Monros (Col., Chrysomelidae). Boletin de la Ac- ademia Nacional de Ciencias 40: 49—54, 3 figs. 1958 Notas y descripciones de Criocerinae (Col. Chrys- omelidae). Boletin de Academia Nacional Cien- cias Cordoba 40: 171—187, 13 figs. . Enumeracion de las Donaciinae africanas con no- tas a algunas especies (Coleoptera: Chrysomeli- dae). Diamang (Companhia di Diamantes de An- gola) 38: 87-92, 6 figs. . Die Gattung Microdonacia Blackburn (Col. Chrysomelidae). Entomologischen Arbeiten aus dem Museum G. Frey 9(3): 742-749, 7 figs. . Descripcion de una nueva tribus de Clytrinae. Acta Zoologica Lilloana 15: 35—39, 4 figs. Consideraciones sobre la fauna del Sur de Chile y revision de la tribus Stenomelini (Coleoptera, Chrysomelidae). Acta Zoologica Lilloana 15: 143-153, 6 figs. 1959 Diez nuevas especies de Lema del subgénero Quasilema (Col., Chrysomelidae). Acta Zoologi- ca Lilloana 16: 23-31, 10 figs. Coleopterorum Catalogus, Supplementa, Chryso- melidae: Sagrinae. pars 51(1): 1-19. W. Junk. Gravenhage. Notes on Lamprosomatinae (Chrysomelidae). Co- leopterists Bulletin 12: 29-33, 4 figs. 1960 . Los géneros de Chrysomelidae. Opera Lilloana (Tucuman) 3: 5-337, 194 figs., 3 pls. . Notas s6bre Chrysomelidae. Acta Zoologica Lil- loana 17: 1—24, 28 figs. . Illustraciones de ‘“‘Hispinae”’ (Chrysomelidae, Co- leoptera). Acta Zoologica Lilloana 17: 25—32, 24 figs. Coleopterorum Catalogus, Supplementa, Chryso- melidae: Lamprosomatinae. pars 51(10): 1-16. W. Junk. Gravenhage. 859 OBITUARIES AND INFORMATION ON THE MONROS COLLECTION Blake, D. H. 1958. Francisco de Asis Monros, 1922— 1958. Proceedings of the Entomological Society of Washington 60: 188-189. Blake, D. H. 1961. A note on the Monros collection. Proceedings of the Entomological Society of Washington 63: 207-208. Halffter, G. 1958. Francisco Monros 1922-1958. Ciencia, Mexico. Revista hispano-americano de ciencias puras y aplicadas 18: 152-153. Martinez, A. 1958. Francisco de Asis Monros (1922- 1958). Neotropica. Notas zoologicas sudamerican- as. Buenos Aires. 4: 64. Wygodzinsky, P. 1959. Francisco de Asis Monrés 1922-1958. Acta Zoologica Lilloana 17: XII-XXI (obituary and biblography). TAXA PROPOSED BY MONROS Alticinae Microdonacia schmidti 57: 747 truganina 57: 748 Aulacoscelinae Aulacoscelis confusca 42: 336 vogti 64: | Janbechynea n. gen. 36: 21 (Bothroscelis) n. subgen. 42: 345 inverosimilis 42: 350 paradoxa 36: 21 Cassidinae Paranota n. gen. 19: 396 Chlamisinae Aulacochlamys n. gen. 34: 657 minuta 34: 663 pygidialis 34: 661 radiata 34: 662 rectecarinata 34: 658 ultima 34: 665 Chlamisus acalay 34: 551 adustus 14: 172 aeronauticus 34: 594 aerosus 14: 177 aerosus silenciosus 43: 83 annalipes 22: 256 bandeirante 14: 189 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bechynei 15: 626 bicornutus 43: 92 binomis 43: 77 bolivianus bowditchi 20: 415 borgmeieri 15: 627 bryanti 43: 88 bryanti colombianus 43: 89 caribe 43: 85 chalcomicanus 43: 84 claraplex 34: 618 colima 22: 257 coya 34: 604 cribricollis 21: 460 discalceatus 34: 634 erinaceus 21: 457 erodus 14: 182 goeldii 20: 421 guarani 34: 627 hispidulus llajtamaucanus 21: 455 igneus 14: 174 inca 43: 87 inopinatus 34: 629 insolitus 21: 456 integrithorax 34: 620 jabaquarensis 22: 629 kurkuncho 34: 632 longicornis 34: 625 malaisei 22: 265 mariae 22: 260 melzeri 22: 267 melochiae 34: 551 mendesi 22: 263 mimicus 20: 420 mourei 43: 90 obidensis 15: 623 ocellatoides 43: 92 octomaculatus 20: 418 oralis 43: 91 oropedion 20: 416 paulista 22: 270 perforatus 34: 606 pilaga 34: 576 pilicollis 34: 635 podtiaguini 43: 90 proseni 34: 630 pseudoimpressus 20: 423 rogaguanus 21: 458 sidae 34: 596 simillimus 14: 179 sordidulus 34: 614 tarijanus 14: 187 taurus 43: 94 trifolium 15: 620 tucumanus 34: 622 vianai 34: 611 weisei 22: 261 xanthopyga 43: 93 yaguar 21: 462 zikani 22: 271 zischkai 43: 83 Diaclaspis n. gen. 64: 3 Diplacaspis chlamysoides 43: 81 pectoralis 43: 80 pici 64: 3 Exema carinipennis 34: 653 serijaniae 34: 650 variopicta 34: 655 Melittochlamys n. gen 14: 192 achardi 15: 619 exscula 14: 194 freyi 21: 452 nicki 15: 617 Pseudochlamys bellicosus 43: 78 Chrysomelinae Acalligrapha n. gen. 48: 55 Bidensomela n. gen. 48: 54 bidenticola meridionalis 48: 54 Calligramma n. gen. 48: 56 Coreopsomela n. gen. 48: 55 Graphicallo n. gen. 48: 57 Oreina n. gen. 53: 1129 Clytrinae Aratea gigantea 35: 360 Babia (Heterobabia) n. subgen. 24: 158 repetita 28: 123 Clytra binominata 37: 49 oblita 37: 49 weisei 37: 49 Clytrasoma balyi 37: 49 VOLUME 97, NUMBER 4 Coscinoptera atypica 40: 116 bahiana 25: 1146 brittoni 25: 1147 euryscopoides 40: 127 heterotricha 35: 355 humeralis 40: 116 murina 35: 354 pseudobliqua 25: 1149 Cylindrodachrys n. gen. 4: 148 cleroides 4: 149 Dachrys balyi 24: 155 bryanti 24: 156 carioca 28: 121 elongaticeps 35: 358 impressicollis 20: 119 kuscheli 24: 154 malleri 28: 118 muriense 35: 357 punctipennis 24: 152 simulans 24: 157 wittmeri 24: 152 yungana 28: 120 Diapromorpha hottentota 37: 49 Dinophthalma amazonica 24: 150 amplicollis 40: 114 anchora 35: 356 coroicana 24: 149 Eoclytra n. gen. 58: 36 freudii 58: 36 Eoclytrini new tribe 58: 35 Euryscopa albopilosa 40: 106 coscinopteroides 23: 301 denieri 40: 110 guerini 40: 109 leechi 35: 352 metropolitana 23: 299 nana 23: 303 nigerrima 23: 300 ordinata 23: 304 sanguinipennis 23: 295 similaris 23: 301 tosta 23: 293 vagabunda 23: 296 vagabunda seriatopunctata 35: 354 Gyandrophthalma tetraspilota 35: 349 Heterobabia n. gen. 24: 158 megistops 24: 159 Ischiopachys empyrea smaragdina 40: 266 noctivaga 35: 361 reticulicollis 51: 161 Labidostomis mannerheimi 37: 49 Lacordairella n. gen. 37: 47 Megalostomis balyi 25: 1152 bubalus bubaloides 40: 99 chuncho 25: 1151 dynamica 35: 351 mariae 25: 1154 platyceros 25: 1156 robustipes 40: 67 runa 25: 1158 univittata oblita 40: 64 univittata pacifica 40: 66 vianai 10: 170 weyrauchi 35: 350 Pnesthes instabilis minuta 40: 151 Protoclytra (Lacordairella) n. subgen. 40: 47 Saxinis blakeae 24: 160 meridionalis 40: 258 Stereoma laevicollis bosqi 40: 225 laevicollis orophila 40: 224 obesa 40: 225 sagittifera 40: 230 Tellenina n. gen. 40: 47 Temnodrachys n. gen. 24: 150 (Eudachrys) n. subgen. 40: 49 amazonica 24: 150 cisplatina 40: 187 complexa pallipes 40: 207 decolorata 40: 204 haywardi 40: 184 hybrida 40: 161 impressifrons 40: 174 lacordairei 40: 200 monticola 40: 195 oyagvaca 40: 19] 861 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pauperrima 40: 160 bosqi 9: 80 yuntana 40: 192 bucki 55: 184 pygmaea 40: 205 chanchamaya 55: 182 guichua 40: 169 chapadensis 60: 24 sympathica 40: 202 chiriguana 33: 47 taeniatoides 40: 199 chromatopus 30: 477 trifoliae 40: 185 cochabamba 54: 52 vianai 40: 175 commelinarum 9: 85 willinki 40: 188 constrictithorax 60: 23 wygodzinskyi 40: 180 curaca 60: 30 xerophila 40: 201 devota 10: 161 Urodera diabroticoides 10: 157 coripatana 28: 124 discospilota 63: 219 cryptocephala 35: 359 dolichosoma 10: 165 dachrymorpha 28: 126 elegans 10: 167 duplicata 40: 249 elegantissima 30: 477 fallaciosa 40: 246 erythrobasalis 30: 477 hamatifera densepunctata 40: 243 fastidiosa 10: 170 inornata 40: 253 felix 10: 158 laevicollis orophila 40: 222 garimpeira 60: 27 languginosa 40: 239 germari 9: 83 mariameliae 28: 125 guanabara 52: 41 martinezi 28: 129 guerini 30: 474 piluloides 28: 128 heinzei 63: 193 : homonyma 10: 169 CEC SMES huiracocha 60: 30 Crioceris hupiracocha 63: 337 (listed in text as ignorata 9: 78 pupiracocha p. 214) natalensis 63: 179 ichilana 52: 43 piliceps 55: 174 jacobyana 63: 221 Elisabethana jangadeira 60: 25 gambiensis 55: 182 lacordaireana 52: 44 Lema langei 52: 35 (Hapsiddemoides) n. subgen. 30: 470 lebiana 33: 46 (Neolema) n. subgen. 30: 472 lima 52: 40 (Pachylema) n. subgen. 30: 473 malleri 52: 36 (Parhapsidolema) n. subgen. 30: 472 mediovittata 63: 187 (Quasilema) n. subgen. 30: 468 melodiosa 33: 42 amnesia 10: 170 mimica 10: 160 amplipalpa 60: 28 monoleptoides 10: 163 annuliantennata 30: 476 multichroma 30: 477 autoptera 33: 45 oglobini 9: 87 azureidorsis 63: 190 pachamana 60: 31 basicostata 10: 166 plaumanni 63: 215 bechynei 55: 185 polichroma 33: 33 bicolora 63: 184 pseudoviolacea 54: 50 bilineata chola 52: 39 quadrivittata xanthogramma 13: 213 binomis 10: 170 riparia 9: 82 bispilota 30: 477 rufibasis 63: 220 VOLUME 97, NUMBER 4 schadei 30: 471 simplissima 30: 472 solanophaga 52: 42 strangulicollis 63: 202 togoana 63: 204 tradescantiae 13: 214 tuberculifera 63: 189 tuberculosa 30: 477 tumapasa 60: 215 urbana 13: 215 usambariensis 63: 204 varayoj 60: 28 variolosa 52: 36 vianai 30: 468 vivax 10: 155 weiseana 63: 204 weyrauchi 52: 38 wittmeri 54: 54 woytkowskii 52: 40 Lilioceris (Chujoita) n. subgen. 63: 148 bondari 55: 175 fairmairei 63: 174 ignorata 9: 78 Metopoceris (Heinzenella) n. subgen. 55: 177 mariae 55: 181 wenzeli 55: 180 Oulema tanganyikana 63: 227 testaceitarsis 63: 227 Plectonycha juncta 8: 57 meditabunda 13: 216 melanoptera 8: 63 vorax 13: 218 xibixibi 33: 48 Pseudocrioceris brachydactylus 55: 172 Cryptocephalinae Mylassa chachallasi 17: 513 discariana 17: 507 frigens 17: 518 Donaciinae Donaciasta cupensis 56: 91 goeckii 56: 89 quioca 56: 9] Sominaella n. gen. 53: 1121 Eumolpinae Bohumiljania n. gen. 59: 149 antiqua 59: 150 Crowsonia n. gen. 32: 194 tetradactyla 32: 195 Cubispini new tribe 38: 26 Eubrachys n. gen. 53: 1127 Habrophora gemella 32: 194 mutila 32: 193 ornata 32: 192 picturata 32: 193 simplex 32: 191 Hornibiini new tribe 18: 551 Lio n. gen. 18: 554 Paraulaca n. gen. 53: 1125 Philippimolpus n. gen. 32: 196 Phytospinthera n. gen. 53: 1124 Stenomelini new tribe 59: 143 Hispinae Anisostena (Neostena) n. subgen. 11: 203 missionensis | 1: 203 Anoplitis difficilis 11: 210 Cephaloleia bondari 5: 414 Demotispa argentina 11: 158 Nanocthispa n. gen. 11: 312 Oediopalpini new tribe 11: 150 Parabaliosus n. gen. 11: 254 ogloblini 11: 254 Probaenia crenata interrupta 11: 273 Stenispa gemignanii 11: 167 proxima 11: 169 Sternostena laeta uhmanni | 1: 198 Sternostenoides n. gen. 11: 267 daguerrei | 1: 268 Uhmannispa n. gen. I|1: 172 maculata 11: 172 863 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON roplata ogloblini 11: 288 reducta 11: 297 Xenochalepus bajulus chacoensis 11: 234 Lamprosomatinae Dorisina n. gen. 50: 58 Guggenheimia n. gen. 50: 52 sumatrana 50: 53 Histerogaster n. gen. 14: 198 phaleroides 14: 200 Lamprosoma acaciae 12: 93 auricaudatum 50: 64 byrrhoides 12: 85 chorisiae 12: 92 chorisiae chaguancum 12: 93 dissectum 12: 83 incredibilis 50: 66 indigaceum 12: 91 minimum 14: 196 nassutum 50: 63 oomorphum 12: 88 podtiaguini 10: 173 subelongatum 12: 87 subhumerale 12: 89 subnitidium 12: 90 zariateguii 12: 94 Lamprosomoides n. gen. 62: 33 monticola 62: 33 Neochlamysini new tribe 62: 29 Oomorphoides n. gen. 50: 54 Oomorphus alvargengai 62: 32 amazonicus 62: 33 caledonicus 62: 32 darlingtoni 50: 48 goiasensis 62: 33 microbius 62: 32 wittmeri 50: 50 Xenoomorphus n. gen. 50: 40 bicornutus 50: 41 cavifrons 62: 31 draconianus 62: 29 gingindhlovuanus 62: 31 innominatus 62: 31 Megalopidinae Agathomerus (Agathomeroides) n. subgen. 7: 185 (Euagathomerus) n. subgen. 7: 190 (Mesagathomerus) n. subgen. 7: 182 (Trichagathomerus) n. subgen. 7: 196 bichito 6: 146 varians 6: 146 Bothromegalopus n. gen. 7: 204 Mastostethus martinezi 7: 178 minutus 7: 172 Megalopus jacobyi apicalis 7: 214 jacobyi inhumeralis 7: 214 jacobyi pallidus 7: 214 jacobyi testaceus 7: 213 schaeferi 7: 211 vespa 7: 208 vespa inornatus 7: 211 Plesioagathomerus n. gen. 6: 149 canus 6: 152 vittatus 6: 150 Megascelidinae Mariamela n. gen. 26: 171 wittmeri 26: 171 Megascelis lemoides 64: 14 picturata 64: 16 yungarum 64: 15 Sagrinae Atalasis sagroides confluens 13: 208 sagroides nigripennis |: 422 Coolgardica kellerberrina 63: 45 Diaphanopsidini new tribe 63: 7 Duboulaya n. gen. 53: 1120 Mecynodera papuana 63: 51 Megamerus alvarengai 49: 108 Neodiaphanops pectoralis 63: 56 Sagrinola n. gen. 53: 1120 VOLUME 97, NUMBER 4 ACKNOWLEDGMENTS I thank S. L. Staines for the Spanish translations. T. N. Seeno, California De- partment of Food and Agriculture, com- mented on an earlier draft of this manu- 865 script. Abraham Willink, Instituto Superior de Entomologica, Universidad Nacional de Tucuman, commented on the biographical sketch and shared helpful insights as a friend and coworker of Monros. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 866-871 DESCRIPTION OF NEOCOLOCHELYNA HAKUSANA, SP. NOV., AND ITS LARVA (HYMENOPTERA: TENTHREDINIDAE) FROM JAPAN ICH TOGASHI |-chome, Honmachi, Tsurugi-machi, Ishikawa, 920-21, Japan. Abstract.—The adult and larva of Neocolochelyna hakusana, sp. nov., from Honshu (Japan Sea coast), Japan, is described and illustrated. Larvae feed on Actinidia arguta. Life history notes are given. Key Words: In late June of 1994, I found some adults of Neocolochelyna Malaise ovipositing in the leaves of Actinidia arguta Plauch on Mt. Hakusan, Ishikawa Prefecture, Japan, and captured three specimens. After com- paring these specimens with N. itoi Takeu- chi which occurs on the Pacific coast, I be- lieve they represent a new species. I also found several larvae of this new species feeding on the leaves of A. arguta, and three larvae were collected for rearing in early July, 1994. This new species is de- scribed below, including description of the larva and biological notes. KEY TO THE JAPANESE SPECIES Ie Remial G.2a05 Me pcus enh Seer ever neue atta 3) aoe 2 Male 2. Eye in dorsal view nearly as long as head be- hind the eyes (Fig. 1); lancet with 23 serrulae (Figs. 10 and 12) — Eye in dorsal view slightly longer than head be- hind the eyes (ratio about 1.2:1.0); lancet with 25 Senmlde (Hiss: I1-and 13); 225.2 itoi Takeuchi 3. Apical portion of subgenital plate slightly an- gulated (Fig. 14); apical portion of harpes rath- er bluntly rounded (Fig. 16); penis valve as in Pig S o ei ae ee ae Se hakusana sp. nov. — Apical portion of subgenital plate rounded eee eee hakusana sp. nov. (Fig. 15); apical portion of harpes rather sharp- ly rounded (Fig. 17); penis valve as in Fig. 19 Rice ies oan) sop iay Macy even Maer egal oes even ttcas itoi Takeuchi sawfly, Neocolochelyna, larva, food plant, Actinidia arguta, Japan Neocolochelyna hakusana Togashi, NEW SPECIES Figs: 1—5,, 10; 12, 14:16:18 Female.—Length 17-19 mm. Robust species. Body ferruginous, with following parts dark brown to black: apical portion of mandible, lower portion of frons except for supraclypeal area, pronotum except for la- tero-posterior corners, triangular maculae on praescutum, posttergite, and posterior margin of propodeum. Antenna dark brown with basal three segments ferruginous. Legs: coxae and femora black, trochanters and tibiae dark brown, tarsi dirty yellow. Head seen from above transverse, dilated behind eyes (Fig. 1); eye in dorsal view nearly as long as head behind eyes (Fig. 1); postocellar area with rather short median furrow (Fig. 1); OOL:POL:OCL = 2.8:1.0: 3.3; postocellar, lateral, and interocellar fur- rows distinct (Fig. 1); circumocellar furrow distinct but lower portion interrupted (Fig. 1); antenna stout, nearly % as long as costa of forewing (or slightly longer than thorax, ratio about 1.0:0.9), relative lengths of seg- ments about 2.2:1.0:3.6:1.8:1.4:1.4:1.1:0.7: 0.8; pedicel longer than its apical width (ra- tio about 1.0:0.7). Thorax: normal; hind basitarsus slightly VOLUME 97, NUMBER 4 867 a Figs. 1-9. 9 g 1-5. Neocolochelyna hakusana sp. nov. 1, head, dorsal view, 2, antenna, lateral view, except for scape, 3—5, sawsheath, lateral view. 6—9. Sawsheath of N. itoi, lateral view. Scale for 1—3 and 6: 1 mm. Figs. 10-13. 10—11. 9th to 10th serrulae of lancet, 10, N. hakusana sp. nov., 11, N. itoi. 12-13. Basal two serrulae of lancet, 12, N. hakusana sp. nov. 13, N. itoi. shorter than following 4 segments com- bined (ratio about 1.0:1.1—1.2). Abdomen: sawsheath as in Figs. 3-5. Lancet with 23 serrulae (Figs. 10 and 12). Punctation. Head strongly, coarsely and reticulately punctured. Thorax strongly and reticulately punctured. Male.—Length 17 mm. Coloration and structure similar to female. OOL:POL = 2.0:1.0; apical portion of subgenital plate slightly angulated (Fig. 14); apical portion of harpes rather bluntly rounded (Fig. 16); penis valve as in Fig. 18. Distribution.—Japan (Japan Sea coast of Honshu)(Fig. 20). Holotype: female, Mt. Hakusan (altitude about 1300 m), Ishikawa Pref., 30. VI. 1994, I. Togashi leg. Preserved in the col- lection of the Department of Zoology, Na- tional Science Museum (Nat. Hist.), Tokyo. Paratypes: one female and one male, Mt. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Le | Figs. 14-19. 14-15. Apical margin of subgenital plate, 14, N. hakusana sp. nov. 15, N. itoi. 16-17. Male genitalia, ventral view, 16, N. hakusana sp. nov. 17, N. itoi. 18-19. Penis valve, lateral view, 18, N. hakusana sp. nov. 19, N. itoi. Scale for 14-17: 1 mm, scale for 18-19: 0.5 mm. E132" E134" 8136" = g138° = E140" Fig. 20. Distributional map of Neocolochelyna spp. (Closed circles = N. itoi; open circles = N. hakusana sp. nov.) VOLUME 97, NUMBER 4 869 Figs. 21-26. dorsal view, 24, labrum, 25, maxilla, 26, labial palpus. Scale for 21—23: 1 mm. Arashima, Fukui Pref., 6. VI. 1982, T. Mu- rota leg.; two females, same data as for ho- lotype. One paratype female is deposited in the National Museum of Natural History, Washington, D.C., one paratype male is de- posited in the Department of Zoology, Na- tional Science Museum (Nat. Hist.), Tokyo, and others in my collection. Remarks.—This new species is very closely allied to N. itoi in coloration and structure. However, it is distinguished from the latter by the ratio between OOL and POL (in itoi, the ratio between OOL and POL is about 1.7:1.0), by the ratio between the eye and the head behind the eyes (in itoi, the eye is slightly longer than the head behind the eyes, ratio about 1.2:1.0), by the number of serrulae of the lancet (in itoi, the number of serrulae is 25), and by the char- acters of the male genitalia (see Figs. 14— 19): Larva of N. hakusana sp. nov. 21, head, frontal view, 22, head, lateral view, 23, mandibles, DESCRIPTION OF LARVA OF N. hakusana, NEw SPECIES Figs. 21—30 Final instar—Length 45-50 mm. Head pale yellow; eye and eye spot black; man- dible black. Body uniformly milky white, covered with thin layer of white wax. Head: vertical furrows distinct (Figs. 21 and 22); antenna 6-segmented; frons rather triangular in form, with 14 setae (Fig. 21); clypeus with 2 long setae (Fig. 21) on each side; labrum with shallow longitudinal fur- row (Fig. 21) and with 2 long and 24 short setae on each side (Fig. 24); mandibles as in Fig. 23; maxillary palpus 3-segmented, relative lengths of segments about 1.0:0.7: 1.8, galea digit-like (Fig. 25), lacinia with 13 strong setae (Fig. 25); labial palpus 3- segmented, relative lengths of segments about 1.0:1.4:1.7 (Fig. 26). Prothorax 3 annulate; meso- and meta- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 27-30. 30, final instar larva. thorax each with 5-annulate; first to ninth abdominal segments each 7-annulate. Food plant: Actinidia arguta Plauch. Notes on biology.—This species is uni- voltine. Adults appeared in June in 1994 and the female oviposited in many rows along the edge of foliage (Fig. 27). Larvae were present from early July to early Au- gust. First to fourth instar larvae form a col- ony on the under side of the foliage when resting (Figs. 28 and 29). When eating, they go to the edge of the foliage. Final instar larvae are not found in a colony but feed singly. When mature, the larvae drop to the ground and penetrate into the soil where they pass the autumn and winter seasons. Neocolochelyna itoi Takeuchi Figs:.6—9), 1,13, 15;,17,.19 Neocolochelyna itoi Takeuchi, 1951, p. 62; Togashi, 1955, p. 154; Togashi, 1965, p. Eggs and larvae of N. hakusana sp. nov. 27, foliage of Actinidia arguta with eggs in typical oviposition pattern, 28, resting colony of Ist to 2nd instar larvae, 29, resting colony of 3rd to 4th instar larvae, 246; Okutani, 1992, p. 238. Tsuruta and Shinohara, Distribution.—Japan (Pacific coast of Honshu, Shikoku and Kyushu)(see Fig. 20). Food plant.—Actinidia arguta Plauch. Specimens examined.—Kyushu—l &, Mt. Hikosan, Fukuoka, 23. V. 1950, N. Fu- kuhara; 2 2, Mt. Hikosan, Fukuoka, 3. VI. 1971, I. Togashi; | 2, Mt. Kuju, Oita, 14— 15. V. 1986, A. Shinohara. Shikoku—1 °, Mt. Ishizuchi, Ehime, 18. VI. 1978, N. Ya- shiro. Honshu—1 2 1 3, Mt. Odaigahara, Nara, 28-29. V. 1977, K. Mizuno; 3 2 5 3d, Mts. Ohminesan, Nara, 7. VI. 1981, K. Mizuno; | 2, Mt. Kunimiyama, Nara, 5. VI. 1988, K. Mizuno; 1 2, Ohara, Kyoto, 21. VI. 1984, T. Matsumoto; 1 2, Mt. Ku- rotakiyama, Gumma, 20. V. 1987, T. Mat- sumoto. VOLUME 97, NUMBER 4 Supplemental description of adult.— OOL:POL:OCL = 1.7:1.0:2.1; eye in dor- sal view slightly longer than head behind the eyes; sawsheath as in Figs. 6—9; lancet with 25 serrulae (Figs. 11 and 13). Male: apical portion of subgenital plate nearly rounded (Fig. 15); apical portion of harpes rather sharply rounded (Fig. 17); penis valve as in Fig. 19. According my observation at Mt. Hiko- san in 1971, female of this species ovipos- ited in many rows along the edge of the foliage of Actinidia arguta. But I do not found the larvae. ACKNOWLEDGMENTS I cordially thank Dr. David R. Smith, USDA, Washington, D.C., for the review of 871 my manuscript. Also, I thank Dr. A. Shi- nohara, National Science Museum (Nat. Hist.), Tokyo, and Mr. T: Murota, Fukui Prefecture, for the loan of material used in this work. LITERATURE CITED Okutani, T., K. Tsuruta, and A. Shinohara. 1992. Re- cords of Neocolochelyna itoi Takeuchi (Hymenop- tera, Tenthredinidae) in Honshu. Japanese Journal of Entomology, 60: 238. Takeuchi, K. 1951. New and unrecorded sawflies from Shikoku, Japan 1). Transactions of the Shi- koku Entomological Society, 2: 57-62. Togashi, I. 1955. (Occurrence of Neocolochelyna itoi Takeuchi in Kyushu). Kontyt, Tokyo, 23: 154 (In Japanese). . 1965. Tenthredinidae. /n Asahina et al., eds., Iconographia Insectorum Japonicorum Colore Na- turali Edita, 3: 246. Hokuryukan, Tokyo (In Jap- anese. ) PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 872-878 LEPIDOPTERA ASSOCIATED WITH GREENLEAF MANZANITA, ARCTOSTAPHYLOS PATULA E. GREENE (ERICACEAE), IN SHASTA COUNTY, CALIFORNIA MICHAEL A. VALENTI! AND RICHARD S. ZACK Department of Entomology, Washington State University Pullman, Washington 99164- 6382. Abstract.—Fifty lepidopteran taxa were collected in association with greenleaf manzanita. Of these, 31 were collected from manzanita foliage as larvae and 29 were reared to adult- hood. A listing of taxa is presented including method of collection, life stage(s) encountered in the field, collecting locations, and parasitoids reared from collected specimens. Key Words: Greenleaf manzanita, Arctostaphylos pa- tula E. Greene (Ericaceae), is a broadleaf evergreen shrub that commonly occurs in the Sierra Nevada mountain range of the western United States (Hickman 1993). This particular manzanita species grows in mesic habitats associated with montane for- est zones generally above 303 m (1000 ft) in elevation (Ball et al. 1983). In general, information concerning insects and brush- field ecosystems (e.g. montane chaparral communities) is currently lacking in the lit- erature (Force 1990); this is certainly the case concerning herbivorous insects asso- ciated with greenleaf manzanita (e.g. Haws et al. 1988). In response to the growing concern over a limited number of control options for greenleaf manzanita shrubs which compete with more economically im- portant timber species, a study was con- ducted from 1989 to 1994 to evaluate the potential of using native insects to alter vegetative composition (Valenti 1994). More than 500 insect species were identi- fied in association with greenleaf manzani- ' Present address: Delaware Department of Agricul- ture, 2320 S. Dupont Hgwy., Dover, DE 19901-5515. Send reprint requests to RSZ. Lepidoptera, Arctostaphylos patula, greenleaf manzanita, parasitoids ta. In this paper we present results of the Lepidoptera inventory. This list includes important host information on many species for which virtually nothing is known. In ad- dition, records of parasitoids and other nat- ural enemies, when available, are presented for a number of species. MATERIALS AND METHODS Site descriptions.—Bear Wallow. This 20+ ha, old-growth greenleaf manzanita brushfield (elevation 1524 m [5000 ft]) is located 3 km (by dirt road) from CA Route 89 in Old Station (approximately 6.6 km south southwest of the Logan Lake site) (T32N R4E S10). Greenleaf manzanita, which averaged 1.5—2.0 m in height, ac- counted for approximately 85% of the total vegetation. Hat Creek. Located adjacent to the USDA Forest Service Work Center, Hat Creek, CA (T34N R4E S16) (elevation 1018 m [3340 ft]), this site consists of a variety of woody plant species. Greenleaf manzanita is fairly common but accounted for less than 30% of the total vegetation. Other plant species present at this site in- cluded: sagebrush (Artemesia tridentata VOLUME 97, NUMBER 4 Nuttall) (Asteraceae); curl-leaf mountain- mahogany (Cercocarpus ledifolius Nut- tall), birch-leaf mountain-mahogany (C. betuloides Torrey & A. Gray), and ante- lope bitterbrush (Purshia_ tridentata [Pursh] de Candolle) (Rosaceae); Califor- nia black oak (Quercus kelloggii Newber- ry) (Fagaceae); and a mixture of conifers including ponderosa pine (Pinus pondero- sa Lawson), sugar pine (P. lambertiana Douglas), white fir (Abies concolor [Gor- don & Glendinning] Lindley), Douglas-fir (Pseudotsuga menziesii [Mirbel] Franco), incense cedar (Calocedrus decurrens |Tor- rey] Florin), and western juniper (Junip- erus occidentalis Hooker) (Pinaceae). Logan Lake. This site (elevation 1512 m [4960 ft]) is located 3.2 km (by dirt road) northwest of CA Route 89 in Old Station, approximately 24 km south of Hat Creek, CA (T32N R4E S2&3). Dominant vegeta- tion consisted of greenleaf manzanita (74%), tobacco brush (Ceanothus velutinus Hooker) (Rhamnaceae) (11%), and ponder- osa pine (9%). The 50+ ha site is located 1.6 km west of Logan Lake on an east-fac- ing slope and was mechanically cleared of all standing vegetation in 1976 and planted to ponderosa pine the following year. Greenleaf manzanita shrubs averaged 1.5— 2.0 m in height. Tamarack Swale. In 1974, this 25+ ha site was mechanically cleared and ponder- osa pine seedlings were planted in 1975. The site is located in a valley 6.4 km (by dirt road) southwest of CA Route 89 ap- proximately 5 km south of Hat Creek (T33N R4E S4&9) (elevation 1646 m [5400 ft]). Tobacco brush (16%) and pon- derosa pine (8%) were interspread with the dominant species, greenleaf manzanita (61%). Greenleaf manzanita shrubs were removed again in 1984 during a second me- chanical clearing treatment, however, shrubs subsequently became reestablished and averaged 0.75 m in height. Other location. In May 1991, a localized outbreak of a tussock moth (Lymantriidae) was discovered in Redding, California, at 873 the junction of Hilltop Drive and California Route 44. Larvae were actively feeding on an ornamental manzanita, Arctostaphylos densiflora Baker, and several dozen were subsequently transferred to caged A. patula plants at the Hat Creek Forest Insect Lab- oratory. Arthropod survey.—Lepidoptera were sampled by visually searching and hand picking individuals from plants, sweeping foliage with a canvas net, and Malaise trap- ping (at the Logan lake site only). Gener- ally, collecting began in May and ended in August for the years 1989 to 1994. Adult specimens collected in the field were trans- ported to the USDA Forest Service, Forest Insect Laboratory, Hat Creek, CA, for prep- aration. All adult specimens were pinned and labelled with complete collecting infor- mation. Voucher specimens are deposited in the Maurice T. James Entomological Col- lection, Department of Entomology, Wash- ington State University, Pullman. Attempts were made to rear all encoun- tered immatures to adults. Individual larvae were placed in small plastic containers or screen cages with host material and allowed to develop and pupate. Many pupae re- quired a cold treatment (e.g., 90 d at 4°C) before adults emerged. Parasitic flies (Dip- tera) and wasps (Hymenoptera) were col- lected from these larval rearings. RESULTS Following the scientific name of each species is the method of collection (Hp— hand picking, Sw—sweeping foliage, or Mt—Malaise trap), stage(s) encountered (E—egg, L—larva, P—pupa, A—adult, and rA—reared adult), and collection lo- cation (BW—Bear Wallow, HC—Hat Creek, LL—Logan Lake, TS—Tamarack and Swale, and OL—other location). Greenleaf manzanita is the host for 31 of the species collected and, unless otherwise noted, apparently represent new host re- cords. None of the hymenopteran parasit- oids encountered in this study are listed in the comprehensive catalog by Krombein et PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (979) and just two tachinid (Diptera) isitoid records (as noted) have previous- y been reported (Arnaud 1978). Accounts of spider (Araneae) predation on larvae of a geometrid are also new. Distributional re- cords are included for an additional 19 spe- cies of Lepidoptera collected in association with greenleaf manzanita brushfield com- munities (18 as adults and one as a chrys- alis and reared to adulthood). GREENLEAF MANZANITA HOST RECORDS Psychidae Hyaloscotes fumosa Butler: Hp; L, rA; HC Gracillariidae undet. genus and species (serpentine mine):--Hps BWC. LL, as undet. genus and species (blotch mine): Hp Bw. EES Ss Coleophoridae Coleophora glaucella Walsingham: Hp; L tA BW. HE Leas Related host record: Arctostaphylos glaucella Lindley (Walsingham 1882) This casebearer is also known to feed on other manzanita species in- cluding A. insularis E. Greene, A. viscida C. Parry, and A. patula (J.-E Landry, pers. comm.). Parasitoids reared from larvae: Agathis sp. (Hymenoptera: Bracon- idae): HC Chelonus sp. (Hymenoptera: Bra- conidae): HC Gelechiidae Gelechia panella Busck: Hp; L, rA: HC, LE Pseudochelaria manzanitae (Keifer): Hp; LerAsiG. LL Is Related host record: Arctostaphylos sp. (Duckworth 1964) Parasitoid reared from larvae: Erynnia tortricis (Coquillett) (Dip- tera: Tachinidae): HC Tortricidae Amorbia cuneana (Walsingham): Hp; L, rA: HC Choristoneura sp. (rosaceana_ |Harris] species complex): Hp; L, rA; HC Parasitoids reared from larvae (*also listed in Arnaud [1978]): Erynnia tortricis (Coquillett) (Dip- tera: Tachinidae):+ HC Eumea caesar (Aldrich) (Diptera: Tachinidae):*+ HC Nilea or Lespesia sp. (Diptera: Ta- chinidae): HC Epinotia arctostaphylana (Kearfott): Hp; bec Ac ele Related host record: Arctostaphylos uva-ursi L. (Dyar 1904) E. miscana (Kearfott): Hp; L, rA; LL, TS E. subplicana (Walsingham): Hp; L, rA; JEL, Related host record: Arctostaphylos manzanita C. Parry (Heinrich 1923) E. terracoctana (Walsingham): Hp; L, CAS EE Lycaenidae Incisalia augustus iroides (Boisduval): Hp, 2A; He, Ev Related host record: Arctostaphylos uva-ursi L. (Scott 1986) Geometridae Aethaloida packardaria (Hulst): Hp; E, LE TAs HEEL Several generations of this species were also reared through on kin- nikinnick, Arctostaphylos uva- uGsiles Anacamptodes_ clivinaria (Guenée): Hp; strAsrr Eupithecia sp:2 Hp; EsrAsHe Hesperumia fumosaria impensa Rindge: Hpbi rAsBWaiCGe Ek ats Larval habits were previously un- known. Late larval instars occur in three distinct color morphs which mimic host stems. A yel- low-green morph with a dorsal crimson stripe and a uniformly crimson morph occur on green- leaf manzanita. The third morph is mottled gray and occurs on an- telope bitterbrush. clivinaria VOLUME 97, NUMBER 4 Parasitoids reared from larvae: Deopalpus sp. nr contiguus (Rein- hard) (Diptera: Tachinidae): HE Madremyia saundersii (Williston) (Diptera: Tachinidae): HC Phryxe pecosensis (Townsend) (Diptera: Tachinidae): HC Aleiodes nolophanae (Ashmead) (Hymenoptera: Braconidae): He Nemoria glaucomarginaria (Barnes & McDunnough): Hp; L, rA; LL Porter (1986) reported that wild col- lected larvae are unknown. He collected ova from wild female moths and reared larvae on Cali- fornia live oak, Quercus agrifolia Née, California white oak, Q. lob- ata Née, and cork oak, Q. suber L. (Fagaceae). Larvae we collect- ed from greenleaf manzanita plants have dorsolateral projec- tions, and late instars mimic greenleaf manzanita inflorescence stems both in coloration (yellow, green, and red) and physical ap- pearance. Synaxis cervinaria (Packard): Hp; E, L, rA; BW, HC, LL, TS Larval habits were previously un- known. Mature larval coloration varies from dark gray to crimson (often mottled). Larvae mimic stems and twigs of greenleaf manzanita shrubs (see Valenti [1994] for complete descriptions of all life stages and life history). Parasitoids reared from eggs: Trichogramma_ sp. (Hymenoptera: Trichogrammatidae): HC, LL Telenomus alsophilae Viereck (Hy- menoptera: Scelionidae): HC, [ere Parasitoids reared from larvae: Campylochaeta sp. (Diptera: Ta- chinidae): LL Aleiodes n. sp. (Hymenoptera: Bra- conidae): LL 875 Meteorus rubens (Nees) (Hymenop- tera: Braconidae): LL Dusona nigritibialis (Viereck) (Hy- menoptera: Ichneumonidae): Le bs Euplectrus sp. (Hymenoptera: Eu- lophidae): LL Parasitoid observed in the field attack- ing larval instar III: Goniozus gracilicornis (Kieffer) (Hymenoptera: Bethylidae): HC Predators observed in the field attack- ing larvae: Misumenops celer (Hentz) (Arane- ae: Thomisidae): LL Xysticus sp. (Araneae: Thomisidae): cE Metaphidippus sp. (Araneae: Salti- cidae) LE Lasiocampidae Malacosoma californicum (Packard): Hp Le tA nei M. constrictum (Henry Edwards): Hp; L, rA; HC M. disstria Hiibner: Hp; L, rA; HC Phyllodesma americana (Harris): Hp; L, rA; LL Lymantriidae Orgyia cana Henry Edwards: Hp; L, rA; OL Larvae were collected from Vine Hill manzanita, Arctostaphylos densi- flora, in Redding, California, and reared to adulthood on greenleaf manzanita at Hat Creek; a subse- quent generation was also reared through to adulthood with green- leaf manzanita as host. Saturniidae Hemileuca’ eglanterina shastaensis (Grote) Hp: Lb, ,rA: BE. EL Related host record: Arctostaphylos sp. (Packard 1914) Hyalophora euryalus (Boisduval): Hp; E, Ie. tA He. EAS Related host record: Arctostaphylos sp. (Collins and Weast 1961) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON saturnia mendocino Behrens: Hp; E, L, rAsLieats Related host record: Arctostaphylos to- mentosa (Pursh) Lindley (Ed- wards 1880) Parasitoid reared from larvae: Lespesia sp. (Diptera: Tachinidae): bie Sphingidae Sphinx vashti Strecker: Hp; L, rA; HC Noctuidae Acronicta ? perdita Grote: Hp; L, rA; LL Apharetra californiae McDunnough: Hp; Re rAviel Parasitoid reared from larvae: Periscepsia helymus (Walker) (Dip- tera: Tachinidae): LL Aseptis ethnica Smith: Hp; L, rA; HC, LL Parasitoid reared from larvae: Periscepsia helymus (Walker) (Dip- tera: Tachinidae): LL DISTRIBUTIONAL RECORDS Hepialidae Hepialis hectoides Boisduval: Hp; A; HC Pyralidae Tulsa ? oregonella (Barnes & Mc- Dunnough): Mt; A; LL Hesperiidae Hesperia sp.: Mt; A; LL Nymphalidae Nymphalis californica (Boisduval): Hp; PtAAy BW. Hey LL. TS This species is known to feed on Ce- anothus spp. and has been re- corded on other shrubs including Arctostaphylos (Furniss and Barr 1975). Adults were fairly com- monly encountered and a single chrysalis was found attached to a leaf in a clump of isolated green- leaf manzanita shrubs growing in a gravel pit at the Hat Creek site. Geometridae Apodrepanulatrix litaria (Hulst): Sw; A; Ek Chlorosea nevadaria Packard: Sw; A; HC Cyclophora dataria (Hulst): Sw; A; LL Drepanulatrix rectifascia (Hulst): Sw; A; LE D. unicalcararia (Guenée): Sw; A; LL Itame guenearia (Packard): Sw; A; HC I. quadrilinearia (Packard): Mt; A; LL Nemoria darwiniata (Dyar): Sw; A; HC, LL Sabulodes edwardsata (Hulst): Mt; A; ele Semiothisa signaria dispuncta (Walker): Mt; A; LL Sericosema juturnaria (Guenée): Sw; A; HC S. wilsonensis (Cassino & Swett): Mt; A; | BF, Sphingidae Hyles lineata (FE): Mt; A; LL Paonias myops (J.E. Smith): Hp; A; HC Several adults were collected on the inside of an 8 X 8 m bird exclo- sure (with 1.5 xX 1.5 cm open- ings) containing only greenleaf manzanita. It is doubtful these adults came from outside the ex- closure because they were larger than the net openings. It is quite conceivable that larvae fed on greenleaf manzanita and pupated inside the exclosure. Adults that subsequently emerged became trapped inside the exclosure net- ting. Arctiidae Spilosoma vestalis Packard: Sw; A; HC, LL DISCUSSION The information presented here repre- sents the first in-depth study of the lepi- dopterous fauna associated with a manza- nita species. In two general treatments of insects affecting important western trees and shrubs (Furniss and Barr 1975, Furniss and Carolin 1977), only four species of Lepidoptera are recorded feeding on Arc- tostaphylos spp.; Nymphalis californica, Hemileuca eglanterina, Hyalophora eury- alus, and a species not encountered in our VOLUME 97, NUMBER 4 study, Orgyia vetusta gulosa (Henry Ed- wards) (Lymantriidae). The collection and rearing of 29 species substantiates that a significant number of Lepidoptera utilize greenleaf manzanita as a host. Some of these species are certainly generalists (e.g., Nymphalis californica, Malacosoma cali- fornicum, M. disstria, and Phyllodesma americana) and manzanita may not nec- essarily be the primary host. For most cases, we have presented new records of larval rearings and host associations. Ad- ditionally, a number of new larval/parasit- oid relationships were discovered through the rearings. ACKNOWLEDGMENTS This study would not have been possible without the assistance of many cooperating taxonomists whose efforts and expertise are gratefully appreciated: R. W. Carlson, D. C. Ferguson, R. W. Hodges, P. M. Marsh, A. S. Menke, R. W. Poole, M. E. Schauff, D. L. Vincent, and N. E. Woodley (USDA-ARS, Systematic Entomology Laboratory, Beltsville, MD), J. Shepard (Washington State University, Pullman, WA), J. A. Powell (University of Califor- nia. Berkeley, ‘CA), K. B: Bolte, J. Huber, J.-F Landry, L. Masner, J. E. O’Hara, and M. Sharkey (Centre for Land and Biolog- ical Resources Research, Agriculture Can- ada, Ottawa, Ontario), R. Robertson (Cal- ifornia Academy of Sciences, San Francis- co, CA), R. Crawford (The Burke Muse- um, University of Washington, Seattle, WA), and D. L. Wagner (University of Connecticut, Storrs, CT). Alan A. Berry- man (Washington State University) and George T. Ferrell (USDA Forest Service) provided support for this study which rep- resents, in part, the results of a cooperative agreement between the Department of En- tomology, Washington State University, Pullman, WA, and the USDA Forest Ser- vice, Pacific Southwest Forest and Range Experiment Station, Redding, CA (Agree- ment No. PSW-89-0017CA). We appreci- ate the helpful comments made by M. A. 877 Solis and an anonymous reviewer on an earlier draft of this manuscript. LITERATURE CITED Arnaud, P. H., Jr. 1978. A host-parasite catalog of North American Tachinidae (Diptera). USDA Miscellaneous Publication No. 1319. 860 pp. Ball, C. T., J. Keeley, H. Mooney, J. Seemann, and W. Winner. 1983. Relationship between form, func- tion, and distribution of two Arctostaphylos spe- cies (Ericaceae) and their putative hybrids. Acta Ecologica 4: 153-164. Collins, M. M. and R. D. Weast. 1961. Wild silk moths of the United States. Collins Radio Co., Cedar Rapids, Iowa. 138 pp. Duckworth, W. D. 1964. North American Stenomidae (Lepidoptera: Gelechioidea). Proceedings of the United States National Museum 116: 23-72. Dyar, H. G. 1904. The Lepidoptera of the Kootenai district of British Columbia. Proceedings of the United States National Museum 27: 779-938. Edwards, H. 1880. Transformations of some species not hitherto recorded. Pacific Coast Lepidoptera 27: 2-3. Force, D. C. 1990. Ecology of insects in California chaparral. USDA Forest Service Research Paper PSW-201. 5 pp. Furniss, M. M. and W. E Barr. 1975. Insects affecting important native shrubs of the Northwestern Unit- ed States. USDA Forest Service General Techni- cal Report INT-19. 64 pp. Furniss, R. L. and V. M. Carolin. 1977. Western forest insects. USDA Forest Service Miscellaneous Pub- lication No. 1339. 654 pp. Haws, B. A., A. H. Roe, and D. L. Nelson. 1988. Index to information on insects associated with western wildland shrubs. USDA Forest Service General Technical Report INT-248. 296 pp. Heinrich, C. 1923. Revision of the North American moths of the subfamily Eucosminae of the family Olethreutidae. Bulletin 123, United States Nation- al Museum. 298 pp. Hickman, J. C. 1993. The Jepson manual: Higher plants of California. University of California Press, Berkeley. 1400 pp. Krombein, K. V., P. D. Hurd, Jr, D. R. Smith, and B. D. Burks. 1979. Catalog of Hymenoptera in America north of Mexico. Smithsonian Institution Press, Washington, DC. 2735 pp. Packard, A. S. 1914. Monograph of the Bombycine moths of North America, part ILI, (T. D. A. Cock- erell, ed.). Memoirs of the National Academy of Science. 12: 1-276, + 503-516. Porter, A. H. 1986. Life history of Nemoria glauco- marginaria (Barnes & McDunnough) and larval taxonomy of the tribe Nemoriini (Geometridae: Geometrinae). Journal of the Lepidopterists’ So- ciety 40: 304-314. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ou, J. A. 1986. The butterflies of North America: for vegetation management. Ph.D. dissertation, , natural history and field guide. Stanford Uni- Washington State University, Pullman. 139 pp. versity Press, Stanford, CA. 583 pp. Walsingham, Lord. 1882. North American Coleopho- Valenti, M. A. 1994. Insects associated with greenleaf rae. Transactions of the Entomological Society of manzanita, Arctostaphylos patula E. Greene (Eri- London. 30: 429-442. caceae), and their potential as silvicultural tools PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 879-883 XANTHAPANTELES, A NEW GENUS OF MICROGASTRINAE (HYMENOPTERA: BRACONIDAE) FROM SOUTH AMERICA JAMES B. WHITFIELD Department of Entomology, University of Arkansas, Fayetteville, Arkansas 72701. Abstract.—Xanthapanteles, a new genus of Microgastrinae, is described and illustrated, with X. cameronae, n. sp., from Argentina, as type species. A brief discussion and tabular comparison of the features of Xanthapanteles and several somewhat similar microgastrine genera is presented. Key Words: and species In the process of examining a large bulk of unplaced material of Microgastrinae to prepare a key to the genera for an upcoming manual of New World Braconidae, speci- mens of an extremely unusual undescribed species were encountered that could not be placed, even with some difficulty, into any known genus. The new species is described below and a new genus is erected for it. A comparison of critical features of the new genus with those of related genera is pre- sented along with the generic description. The biological habits of the new genus are unknown. As the new genus is at this time based on only a single species, only a brief dis- cussion of characters distinguishing the ge- nus from other known and presumably re- lated genera will be presented. Accompa- nying this discussion is a tabular compari- son of the states a number of characters of Xanthapanteles and those of a number of the most obviously similar genera (Table 1). Because the phylogeny of microgastrine genera has been a subject of much (and still unresolved) controversy (Mason 1981, Wil- liams 1985, Walker et al. 1990, Austin 1990, Austin and Dangerfield 1992), it would be premature at this time to attempt Parasitoids, Neotropical, Apanteline, Xanthapanteles cameronae new genus a phylogenetic placement of the genus within the subfamily. Nevertheless it is hoped that the character comparisons pre- sented will ultimately contribute to the res- olution of some of the remaining phyloge- netic questions. Terms for wing venation features follow Mason (1986) and a new venational no- menclature (modified slightly from that dis- cussed by Sharkey, 1994) being developed for the upcoming identification manual to the New World genera of Braconidae. Mi- crosculpture terms follow the usage of Har- ris (1979), except that some intermediates between the states discussed by Harris are indicated by compound designations. Xanthapanteles, NEW GENUS Figs. 1-3 Type species: Xanthapanteles cameronae n. sp. (described below). Monobasic. Xanthapanteles possesses a unique com- bination of features that will easily distin- guish it from any known genus of Bracon- idae. Many features of this genus such as the wing venation (the configuration of forewing veins IRs and IM and the absence or reduction to spectral traces of veins 2r- m, 4Rs and 4M, as well as the well-defined PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ible 1. Comparison of several selected morphological characters across a range of microgastrine genera might, on the basis of one or more of the characters, be confused with Xanthapanteles. Characters and states are presented in a simplified fashion; see text and description for further detail. Genus Antennal Placodes Forewing Areolet Propodeal Pattern* Alphomelon two ranks open, 2r-m absent areola, 2 anterior carinae Apanteles two ranks open, 2r-m absent areola (variable) Austrocotesia two ranks closed, 2r-m present areola, | ant. carina Choeras two ranks variable medial carina Dasylagon two ranks closed, 2r-m present areola, | or 2 ant. carinae Dolichogenidea two ranks open, 2r-m absent areola (variable) Exoryza two ranks open, 2r-m absent areola (poorly defined) Hygroplitis irregular or three ranks closed, 2r-m present medial carina Microgaster two ranks closed 2r-m present medial carina Miropotes two ranks closed, 2r-m present areola, | anterior carina Papanteles two ranks closed, 2r-m present areola, usu. I ant. carina Parapanteles two ranks open, 2r-m absent areola (variable) Pholetesor (some)** two ranks open, 2r-m absent areola, | anterior carina Promicrogaster two ranks usually closed by 2r-m —_ medial carina or groove Rhygoplitis two ranks open, 2r-m absent medial carina Xanthapanteles irregular open, 2r-m absent areola, 2 ant. carinae Xanthomicrogaster two ranks closed, 2r-m present medial carina * The propodeal pattern is, in several genera, poorly defined or obscured in some species; the predominant clear pattern is presented. ** Several species groups of Pholetesor would never be confused with Xanthapanteles due to structure of the metasoma and to absence of a propodeal areola. vannal lobe of the hindwing—see Fig. 3), the 18-segmented antennae and the separa- tion of the spiracle of the first metasomal tergum from its central tergite, all clearly indicate that the new genus belongs to the Microgastrinae as currently defined (Austin and Dangerfield 1992, Whitfield and Mason 1994). Xanthapanteles possesses perhaps the most complete (although perhaps some- what exaggerated) pattern of propodeal are- olation of any microgastrine (Fig. 2), sug- gesting at least that it does not belong to one of the lineages possessing derived or reduced propodeal carination patterns such as a medial longitudinal carina or complete- ly rugose surface. The hypopygium is me- dially desclerotized, membranous and ex- pandible through a series of submedial pleats, as in genera such as Apanteles, Dol- ichogenidea, Choeras and Promicrogaster. Austin (1990), in contrast to Mason (1981), considers this to be a possible synapomor- phy for the above group of genera plus per- haps several others. The distribution of sol- id (evenly sclerotized) hypopygia, not only among outgroup taxa but also among puta- tively basal lineages of Microgastrinae, would tend to support Austin’s view. In ad- dition, the structure of the pleated hypopy- gium in the Microgastrinae listed above and in Xanthapanteles is different from superfi- cially similar hypopygia in Cardiochilinae and other related groups in lacking an apical sclerotized bridge. The ovipositor sheaths are moderately long, fairly hairy throughout, and attached about halfway or more up the second valvifers—again as in those genera (as also in many others—see discussions by Austin 1990, Williams 1985). Unlike any of those genera, however, the first metasomal tergite is strongly enlarged, entirely covering the dorsum of the first tergum and it, along with the second and third tergite, has a finely pebble-grained surface unlike that of any other known microgastrine (this appearance is only approximated in figure 2; it perhaps most resembles the sculpturing in some weakly sclerotized hormiines). Lastly, the antennae are unique: the first flagellomere is shorter than the second or third, and sub- VOLUME 97, NUMBER 4 Table 1. Continued. Metasomal Tergite | moderately broad usu. posteriorly narrowing usu. posteriorly narrowing moderately broad very broad moderately broad very broad very broad very broad usually narrow moderately broad moderate to narrow moderate to broad moderately broad moderate to broad extremely broad very broad Tergite Sculpture smooth to punctate smooth to aciculorugose smooth to aciculorugose smooth to aciculorugose smooth to punctate smooth to aciculorugose aciculorugose to rugose aciculorugose to rugose aciculorugose to rugose smooth to aciculorugose aciculorugose aciculorugose aciculorugose to rugose smooth to aciculorugose aciculate to rugose smooth to finely colliculate punctate to aciculorugose Hypopygium evenly sclerotized medially pleated evenly sclerotized medially pleated medially pleated medially pleated medially pleated evenly sclerotized usually pleated medially pleated medially pleated evenly sclerotized evenly sclerotized medially pleated medially pleated medially pleated evenly sclerotized 881 equal with the length of the fourth, and the longitudinal placodes on all longer flagel- lomeres are arranged irregularly rather than in even single or double ranks. This irregular placement of the placodes is known from several other micrigastrine genera, but not from any others with a propodeal areola. The general appearance of the wasp is also slightly unusual, as the mesosoma is some- what more slender and straight-backed than is typical for microgastrines, the color is pri- marily orangish yellow, and the wings are brownish and unusually slender (Fig. 1). In several of these body shape features Xantha- panteles somewhat resembles the presum- ably distantly related Rhygoplitis and Hygro- plitis. Xanthapanteles cameronae, NEW SPECIES Figs. 1-3 Holotype female.—Body length 3.8 mm; forewing length 4.3 mm. Lateral habitus— Fig. 1 Head: Pale honey-orange throughout ex- cept darker antennae and area between ocelli. Frons 2.0 as broad at midheight as long medially, weakly punctate; inner mar- gins of eyes converging towards clypeus, and slightly indented just above antennal bases. Antennae medium brown through- out, 1.2 as long as forewing; all but distal flagellomeres with scattered placodes, oc- casionally forming 3 indistinct ranks on placodes; flagellomere 2 3.75 as long as broad; flagellomere 14 2.3 as long as broad. Palpi light yellow-golden throughout visible portions. Head in dorsal view 1.7 as broad as medially long. Vertex relatively smooth and extremely finely granular, near- ly appearing polished. Mesosoma: Entire mesosoma pale or- ange-brown except darker dorsal portion of pronotum, lateral portions of metanotum and posteromedial section of propodeum. Pronotum with strong, broad, crenulate up- per and lower grooves. Mesoscutum evenly, finely granular but appearing superficially smooth and polished; width just anterior to tegulae slightly less than that of head. Scu- toscutellar scrobe composed of approxi- mately 5—6 partly confluent crenulations, nearly straight medially. Scutellar disc slightly longer than anteriorly broad, simi- larly sculptured to mesoscutum. Mesopleu- ron finely granular, convex, without obvi- ous longitudinal depression. Metanotum an- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. teriorly appressed to scutellum; sublateral setiferous lobes not clearly distinct. Propo- deum (Fig. 2) with strongly raised areolate pattern of carinae, the areola being hexag- onal and bearing 2 anterior carinae leading to anterior margin of propodeum; _ back- ground sculpturing not obvious. Legs: Coloration entirely honey-orange except darker terminal tarsomeres (fore- and mid-legs), apices of hind femora and tibiae, and most of hind tarsi. Spines on outer faces of hind tibiae tiny, inconspicu- ous, irregularly scattered. Apical spurs of hind tibiae subequal in length, the inner spur about 0.4 length of hind basitarsus. Xanthapanteles cameronae n. sp. 1, Lateral habitus. 2, Propodeum and anterior metasomal tergites. 3, Wings. Scale lines: 1—1 mm; 2—0.5 mm; 3—1 mm. Wings: More slender (Fig. 3) than in most microgastrines. Tegulae pale honey- yellow. Forewing venation moderate yel- low-brown; wings tinged slightly brownish. Vein 2r weakly arched, somewhat longer than 2 + 3RS, meeting it at about 100° an- gle. Vein 2r-m absent. Vein RI (metacarp) much longer than stigma, approximately 4X as long as distance from its distal end to end of 4RSb fold along wing edge. Stig- ma 4X as long as broad. Hindwing with vannal lobe subapically flattened, sparsely fringed with hairs. Metasoma: Entirely honey-orange except darker ovipositor sheaths. Anterior ter- VOLUME 97, NUMBER 4 gites—Fig. 2. Tergite I fused to tergite II posteriorly, although not clearly so lateral- ly; broadening posteriorly, slightly longer than posteriorly broad, with extremely pre- cipitous and short anterior excavation; an- terolateral portions strongly raised over edges of tergite; entire surface finely gran- ular. Tergite II sculptured similarly to 1, fused posteriorly to II, subquadrate and just Over twice as broad as long. Tergite III sculptured similarly to I and II, much short- er than II and with weakly rounded pos- terolateral corners. Succeeding terga of nor- mal, unsculptured, overlapping form. Hy- popygium moderately long, acute apically, desclerotized medially into a series of ex- pandable pleats. Ovipositor sheaths about 0.75 as long as hind tibiae, deep brown, slender-fusiform, hairy over most of length. Ovipositor weakly decurved, bladelike. Males: Unknown. Variation: Only two virtually identical females known. Cocoons: Unknown. Material examined.—Holotype female: ARGENTINA: Buenos Aires, La Plata (Fac. Agronomia), X-XI-1968, C. Porter (MCZ). Paratype: 1 female, same data. Ho- lotype and paratype in Museum of Com- parative Zoology, Harvard University. Hosts.—Unknown. Comments.—The species is named for Dr. Sydney Cameron, for her unfailing sup- port for my own efforts, for her many col- lections of braconid wasps over the years, and for her keen interest in the South Amer- ican fauna. ACKNOWLEDGMENTS I would like to acknowledge the helpful notes that the late Bill Mason (Ottawa) placed with the specimens described above—although he did not provide many details, he clearly recognized this species as unusual and had planned to describe it him- self as a new genus. A collection improve- ment grant from the National Science Foun- dation (BSR-82-03845) to Edward O. Wil- 883 son helped support Mason’s initial studies of these specimens. This and other studies of New World braconid genera have been supported by a grant from the National Sci- ence Foundation (DEB-9300517). Chris Carlton, Sydney Cameron and Scott Shaw provided useful comments on the manu- script; Scott Shaw also provided useful in- formation on the curatorial history of these specimens. LITERATURE CITED Austin, A. D. 1990. Revision of the enigmatic Aus- tralasian genus Miropotes Nixon (Hymenoptera: Braconidae: Microgastrinae), with comments on the phylogenetic importance of the female ovi- positor system. Systematic Entomology 15: 43— 68. Austin, A. D. and P. C. Dangerfield. 1992. Synopsis of Australasian Microgastrinae (Hymenoptera: Braconidae), with a key to genera and description of new taxa. Invertebrate Taxonomy 6: 1-76. Harris, R. A. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology, No. 28. State of California, Department of Food and Agricul- ture, Sacramento. 31 pp. Mason, W. R. M. 1981. The polyphyletic nature of Apanteles Foerster (Hymenoptera: Braconidae): a phylogeny and reclassification of Microgastrinae. Memoirs of the Entomological Society of Canada 115: 1-147. . 1986. Standard drawing conventions and def- initions for venational and other features of wings of Hymenoptera. Proceedings of the Entomologi- cal Society of Washington 88: 1-7. Nixon, G. E. J. 1965. A reclassification of the tribe Microgasterini (Hymenoptera: Braconidae). Bul- letin of the British Museum (Natural History) En- tomology Supplement 2: 1—284. Sharkey, M. J. 1994. Another look at wing vein/cell nomenclature. Ichnews 14: 2—S. Walker, A. K., I. J. Kitching, and A. D. Austin. 1990. A reassessment of the phylogenetic relationships within the Microgastrinae (Hymenoptera: Bracon- idae). Cladistics 6: 291—306. Whitfield, J. B. and W. R. M. Mason. 1994. Mende- sellinae, a new subfamily of braconid wasps (Hy- menoptera: Braconidae) with a review of relation- ships within the microgastroid assemblage. Sys- tematic Entomology 19: 61—76. Williams, D. J. M. 1985. The New World genus Lath- rapanteles n. gen.: phylogeny and placement in the Microgastrinae (Hymenoptera: Braconidae: Cotesiini). Canadian Journal of Zoology 63: 1962-1981. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 884-886 NOTES ON PARABLASTOTHRIX NEARCTICA (HYMENOPTERA: ENCYRTIDAE) ROBERT L. ZUPARKO Laboratory of Biological Control, University of California, Berkeley, 1050 San Pablo Avenue, Albany, California 94706. Abstract.—New host records and a range extension are reported for Parablastothrix nearctica, as well as inaccuracies in the original description. Only one adult emerges from each host, suggesting the species is monoembryonic. Parablastothrix has been placed in the Copidosomatini, but its mode of reproduction contrasts with the other genera of the tribe, all of which appear to consist of polyembryonic species. Key Words: matini, polyembryony Parablastothrix nearctica Miller (Hy- menoptera: Encyrtidae) was described from specimens collected in the eastern United States and reared from two leafminers, a Coptodisca sp. (Lepidoptera: Heliozelidae) and an Obrussa sp. (Lepidoptera: Nepticu- lidae) (Miller 1965). In parts of this and some subsequent papers, this species was also referred to as P. nearcticus: this suffix is incorrect because the generic name is based on the Greek feminine thrix. In March 1994, I collected five cocoons from the underside of leaves of a coast live oak (Quercus agrifolia Nee) at our experi- ment station in Albany, California. Three of these were formed by Stigmella variella (Braun) (Lepidoptera: Nepticulidae), from each of which emerged a single Parablas- tothrix adult (two females and one male). A male Paradelius rubra Whitfield (Hy- menoptera: Braconidae) emerged from the fourth cocoon. The Parablastothrix adults exhibited a frenetic activity, typical of many hyperpar- asitoids. But when I dissected the four emp- ty cocoons, I found no more than one me- conial pellet in each cocoon, indicating both Parablastothrix nearctica, Paradelius rubra, Stigmella variella, Copidoso- species were solitary, primary parasitoids. The fifth cocoon held a dead, well-formed P. rubra male pupa. In October 1994, I col- lected 10 more Parablastothrix adults (nine females and one male) from the foliage of the same trees. The Parablastothrix specimens were identical to those reported in a study on par- asitoids of leafmining lepidopterans on oaks in northern California, dating back to 1961 (Green 1979). All specimens had yellow legs with darkened metatibiae, and a green body with a bronze hue, but which other- wise matched the description of P. nearc- tica. Green (1979) reported this encyrtid emerged individually from cocoons of S. variella, and from undetermined stages of Coptodisca powellella Opler, Bucculatrix albertiella Busck (Lepidoptera: Lyoneti- idae) and Phyllonorycter sandraella (Opler) (Lepidoptera: Gracillariidae), while still in their leaf mines. Miller (1965) reported that P. nearctica had white legs, and the overall body color was black with bluish and greenish reflec- tions. Reexamination of the holotype and two paratypes of P. nearctica revealed that VOLUME 97, NUMBER 4 the original description was in error. All three type specimens have dark metatibiae and a green body, identical to the California specimens. Thus the latter represent a range extension of P. nearctica. | have seen ad- ditional specimens of P. nearctica collected from Arizona and central and southern Cal- ifornia. Hosts have been recorded for only 8 of the 17 described Parablastothrix species— all are leaf-mining Lepidoptera from the families Gracillariidae, Lyonetiidae, and Nepticulidae (Hedqvist 1976, Logvinov- skaya 1981, Khan 1983, Trjapitzin 1989). An undescribed Parablastothrix sp. was re- ported as a solitary, primary parasitoid on a nepticulid, a heliozelid, and a gracillariid on pecan in Georgia (Dutcher and Heyer- dahl 1988). Hoffer (1955) placed Parablastothrix in the tribe Microteryini (subtribe Pentacne- mil), with Calometopia Mercet and Pentac- nemus Howard. Noyes and Hayat (1984) synonymized Pentacnemus with Copidoso- ma Ratzeburg, and Trjapitzin (1989) syn- onymized Calometopia with Parablasto- thrix. Trjapitzin and Gordh (1978) placed Parablastothrix in the subtribe Parablasto- thrichina within the Copidosomatini, but re- marked on the absence of proven polyem- bryony. Noyes and Hayat (1984) consid- ered the genera in this subtribe to be mor- phologically closer to the Aphycina (tribe Aphycini); however they did not formally place these genera in their scheme of sys- tematic relationships. Recent work by G. Zolnerowich (pers. comm.) retains Para- blastothrix as a more basal lineage within the Copidosomatini. Within the Copidosomatini, polyembry- ony appears to be the rule, having been re- ported in all species whose biologies have been investigated, including species of Ap- silophrys De Santis, Copidosoma (=Lito- mastix Thomson), Copidosomopsis Girault, Paralitomastix Mercet, Ageniaspis Dahl- bom, Holcothorax Mayr and Paraleuroce- rus Girault (Noyes 1980, Noyes and Hayat 1984). Polyembryony may also occur in 885 Coelopencyrtus Timberlake (Timberlake 1919, Trjapitzin 1960, Taylor 1961, Anne- cke 1968), but this mode of reproduction has never been conclusively demonstrated. Solitary parasitism presupposes mon- oembryony. However, such evidence is not conclusive, because the polyembryonic spe- cies Macrocentrus ancylivorus Rohwer (Hymenoptera: Braconidae) is known to de- velop as a solitary parasitoid (Daniel 1932). Embryological studies have not been con- ducted on any Parablastothrix species. Nevertheless, the solitary habit of P. nearc- tica and the absence of any mention of mul- tiple emergences from all other Parablas- tothrix rearing records, suggests that poly- embryony is absent in the entire genus. Such a lifestyle would then biologically separate this genus from the remainder of the Copidosomatini. Material examined.—ARIZONA. CO- CHISE Co.: 10 mi. W. Portal, on Juniperus. CALIFORNIA. ALAMEDA CO.: Albany, Stigmella variella on Quercus agrifolia; Berkeley Hills, S. variella on Q. agrifolia and Phyllonorycter sandraella,; Patterson Reserve, Del Valle Lake, Stigmella sp.; CONTRA COSTA Co.: Antioch, S. variella on Q. agrifolia; 3 km E of Antioch, S. var- iella on Q. agrifolia; Cowell, S. variella on Q. agrifolia; Oakley, S. variella on Q. agri- folia; Russellmann Park; Russell Tree Farm, 6 km NE of Orinda, S. variella; MARIN Co.: Novato, Coptodisca powellella on Q. agrifolia; Sausalito, on Q. agrifolia; MON- TEREY Co.: 10 km SE of Big Sur; SAN MATEO Co.: Woodside, Bucculatrix alber- tiella on Q. agrifolia,; SANTA BARBARA Co.: Santa Cruz Island, Canada de la Cues- ta, Stigmella sp. on Q. dumosa; SONOMA Co.: Sonoma, Stigmella sp. on Q. agrifolia. MICHIGAN. MIDLANDS Co.. VIRGINIA. Falls Church. MISSISSIPPI. HINDS Co.: Jackson, Coptodisca sp. on Vaccinium ar- boreum Marsh, holotype (no. 8800, depos- ited in Canadian National Collection, Ot- tawa, Ontario) and paratype (deposited in United States National Museum, Washing- ton, D.C.). WEST VIRGINIA, MONON- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘LIA Co.: Morgantown, Obrussa_sp., ratype (deposited in U.S.N.M.). ACKNOWLEDGMENTS I thank Jerry Powell, University of Cal- ifornia, Berkeley, for identifying the S. var- iella cocoons, and Michael Schauff (U.S.N.M.), Vince Lee (California Acade- my of Sciences) and Steven Heydon (U.C., Davis) for providing access to material. Gary Gibson of the Biosystematics Re- search Centre (Agriculture Canada) kindly reexamined the holotype of P. nearctica. Ken Hagen and Leo Caltagirone (U.C., Berkeley) and Gregory Zolnerowich (Texas A&M _ University) provided valuable dis- cussions and advice. LITERATURE CITED Annecke, D. P. 1968. Records and descriptions of Af- rican Encyrtidae 4 (Hymenoptera: Chalcidoidea). Journal of the Entomological Society of South Af- rica 31: 249-264. Daniel, D. M. 1932. Macrocentrus ancylivorus Roh- wer, a polyembryonic braconid parasite of the Ori- ental fruit moth. New York State Agricultural Ex- periment Station, Technical Bulletin #187. 101 pp. Dutcher, J. D. and R. Heyerdahl. 1988. Parasitic Hy- menoptera of the leafmining guild of pecan, pp. 445-458. In Gupta, V.K., ed., Advances in Para- sitic Hymenoptera Research, E. J. Brill, Leiden. Green, D. S. 1979. Ecology and host-specificity of parasitoids of leaf-mining Lepidoptera on Quercus agrifolia (Fagaceae) in California. Ph.D. Thesis, University of California, Berkeley. 204 pp. Hedqvist, K.-J. 1976. Descriptions of new chalcid flies (Hym., Chalcidoidea, Encyrtidae and Eulo- phidae) reared from Nepticula species collected in Sri Lanka. Entomologisk Tidskrift 97: 50-54. Hoffer, A. 1955. The phylogeny and taxonomy of the family Encyrtidae (Hym., Chalcidoidea). Sbornik Narodniho Museu v Praze 11: 1—22. Khan, M. A. 1983. A new species of Parablastothrix (Hymenoptera: Encyrtidae) from India. Journal, Bombay Natural History Society 80: 180-182. Logvinovskaya, T. V. 1981. A review of Holarctic species of the genus Parablastothrix Mercet (Hy- menoptera, Encyrtidae). Entomological Review 60: 147-153. Miller, C. D. FE 1965. A Nearctic species of Para- blastothrix Mercet (Hymenoptera: Encyrtidae). Canadian Entomologist 97: 750-753. Noyes, J. S. 1980. A review of the genera of Neo- tropical Encyrtidae (Hymenoptera: Chalcidoidea). Bulletin of the British Museum (Natural History) 41: 107-253. Noyes, J. S. and M. Hayat. 1984. A review of the genera of Indo-Pacific Encyrtidae (Hymenoptera: Chalcidoidea). Bulletin of the British Museum (Natural History) 48: 131-395. Taylor, J. S. 1961. A note on some insects associated with Xylocopidae in the eastern Cape Province, South Africa. Pan-Pacific Entomologist 37: 220— 222: Timberlake, P. H. 1919. Descriptions of new genera and species of Hawaiian Encyrtidae (Hymenop- tera). Proceedings of the Hawaiian Entomological Society 4: 197-231. Trjapitzin, V. A. 1960. A Palaearctic species of the genus Coelopencyrtus Timb. (Hymenoptera En- cyrtidae). Entomological Review 39: 500-502. 1989. Parasitic Hymenoptera of the Family Encyrtidae of Palaearctics. Leningrad, “‘Nauka,” Leningrad Division. 488 pp. Trjapitzin, V. A. and G. Gordh. 1978. A review of the Nearctic Encyrtidae (Hymenoptera, Chalcidoi- dea). Communication II. Entomological Review 57: 437-448. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, p. 887 NOTE A New Name For A Homonym In Jembrana (Homoptera: Cercopoidea, Aphrophoridae) In 1992 Chou, Yuan, and Liang pub- lished an article in which the name Jem- brana forcipenis Chou and Liang was used for two different new species. This error was made by the printer and was not cor- rected by the authors because they did not see proofs of the article before publication. To correct this unfortunate mistake I pro- pose the following replacement name for the second species. Jembrana wangi Liang, NEw NAME Jembrana forcipenis Chou and Liang, In Chou, I., Yuan, F and Liang, A.-P, 1992: 235, 236 (Chinese), 242 (English), fig. 2 In Chen, S. X., ed., Insects of the Heng- duan Mountains Region, Science Press, Beijing, pp. 234-242. [preoccupied by Jembrana forcipenis Chou and Liang, Jn Chou, I., Yuan, E and Liang, A.-P, 1992: 235 (Chinese), 241, 242 (English), fig. 1 In Chen, S. X., ed., Insects of the Heng- duan Mountains Region, Science Press, Beijing, pp. 234—242.] Jembrana wangi (as J. forcipenis Chou and Liang, /n Chou, I., Yuan, FE and Liang, A.-P., 1992: 235,, 236, 242, fig. 2) was de- scribed from three males from Sichuan (Nanping: Jiuzhaigou) and Shaanxi (Wug- ong) in China. The replacement name is a patronym to honor the collector of the ho- lotype of this spittkebug, Mr. Suiyong Wang. The holotype is housed in the Insect Collection of the Institute of Zoology, Ac- ademia Sinica, Beijing, China. Acknowledgments.—I thank Dr. Lee Herman of the Department of Entomology, American Museum of Natural History, New York, for his kind help during the prepara- tion of this report. Ai-Ping Liang, Roosevelt Research Fel- low, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024, USA and Institute of Zoology, Aca- demia Sinica, 19 Zhongguancun Lu, Beijing 100080, PR China. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, p. 888 NOTE Range Extensions for the Ant Leptothorax pergandei (Hymenoptera: Formicidae): A Mesic Forest Species Discovered in the Chihuahuan Desert Leptothorax pergandei Emery was re- cently evaluated, and determined to be a single, highly variable species (MacKay, W. 1993, Sociobiology 21: 287-297). This spe- cies normally occurs in shaded deciduous forests or shaded prairie sites of eastern United States, as far west as Nebraska, Kansas, Oklahoma and Texas, and as far north as lowa (MacKay 1993, loc. cit.). We were thus very surprised to find this species in the Chihuahuan Desert in Arizona and in New Mexico, the first records of this spe- cies from both states and from such arid habitats. This species was collected at two locali- ties in Arizona. The first site is the Chiri- cahua Mountains, Cochise Co., 6.4 km WNW Jct. National Forest road 42 with National Forest road 42B, elevation 1580 m, 11-vii-1992 (Silver Creek Valley near Paradise). The area consists of a grazed, grassy meadow surrounding a wash with scattered oaks (Quercus gambelii) and ju- nipers (Juniperus monosperma) up to 5 m tall. Other shrubs include rabbitbush (Chrysothamnus sp.) and Hopi tea (Thele- sperma megapotamicum). The soil is fine textured with moderate to heavy clay. The L. pergandei nests are cryptic and fully ex- posed to the sun [vouchers in Museum of Comparative Zoology]. It is common in this area; 45 nests were collected and far more nests were seen. The second Arizona site is Graham Co., 14 mi. S Safford, 1x-13-1976, leg. D. S. Chandler, from sweeping low vegetation [1 specimen in Univ. of Ariz., Tucson, collection]. The first site of two New Mexico collec- tions is Hidalgo Co., San Simon Valley, 1 km W Jct. Rt. 80 on Rt. 533 (Portal Rd.), elevation 1250 m, 2-vili-1988 and 15-vii- 1992. The area consists of open, grazed Chihuahuan Desert scrub with scattered mesquite, Ephedra sp., patches of Hilaria mutica, creosotebush (Larrea tridentata) and mesquite (Prosopis glandulosa). The soil has a moderate to heavy clay compo- nent and parts of the area tend to accumu- late water temporarily during summer rain events. Leptothorax pergandei nests are cryptic, consisting of tiny, bare holes in bare soil or in grass clumps, in areas fully exposed to the sun [vouchers in MCZ]. The second site in southern New Mexico is Dona Ana Co., 45 km NE Las Cruces. It is a typical creosotebush (Larrea tridentata) scrub community. A single worker was col- lected in a pitfall trap, 24-vi-1984 [voucher in Laboratory for Environmental Biology, UTEP]- This species is apparently not common in the Chihuahuan Desert. We have done extensive collecting in Arizona and New Mexico for over 17 years and these are the only Chihuahuan Desert collections we have for this species. In two of the localities (first and third listed), this species is abun- dant, nesting in open, sun-baked desert sites as if it were the “normal” habitat for this species. We have not collected it in more mesic sites in the two states, despite exten- sive collecting in such habitats. William P. MacKay, Laboratory for En- vironmental Biology, The University of Tex- as, El Paso 79968; Stefan P. Cover, Muse- um of Comparative Zoology Laboratories, Harvard University, Cambridge, MA 02138; Jiirgen Heinze and Bert H6élldobler Theo- dor-Boveri-Institut, (Biozentrum der Univ- ersitdt), LS Verhaltensphysiologie und So- ziobiologie, Am Hubland, D-97074, Wiirz- berg, F. R. Germany. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 889-890 NOTE Seasonal Flight Activity of Vanhornia eucnemidarum Crawford (Hymenoptera: Vanhorniidae) in the Mid-Atlantic States I have been collecting with Malaise traps at various sites in Maryland, Virginia, and West Virginia from the coastal plain to the West Virginian and western Maryland Ap- palachians for more than ten years. The traps were in operation for the full season, from March or April to October or mid- November, depending on the site, and I have been able to accumulate seasonal flight data on an array of groups. I have always kept specimens and records of Van- hornia eucnemidarum, the only described North American species of the family. It has been considered rather rare and is scarce in collections. Most collection re- cords of V. eucnemidarum are in spring, and the data presented here, based on col- lections of 398 specimens, verify it as a uni- voltine, spring-flying species. Deyrup (1985. Great Lakes Entomologist 18: 65— 68) gave a comprehensive account of the biology and taxonomy of this species from his work in Indiana. Specimens of V. eucnemidarum are not common, but I have found them at seven collecting sites where I have trapped. Spec- imens have been taken from mid-May to to late July, with the peak flight during the last third of May to the end of June (Fig. 1). The earliest record was V-8-14-1985 from Fairfax Co., Virginia, and the latest record was VII-19-28-1992 from Tucker Co., West Virginia. Vanhornia eucnemidarum is known from Quebec west to Michigan, south to Georgia and Kentucky (Deyrup 1985). Localities for specimens I have examined are as follows, with the earliest and latest dates of collec- tion, years of trapping, (number of speci- mens trapped, and number of traps used per year): MARYLAND: Prince Georges Co., Beltsville Agricultural Research Center, -- 16-23 to VII-11-24, 1991-1993 (32; 3 to 4 traps). VIRGINIA: Clarke Co., University of Virginia Blandy Experimental Farm, 2 mi. S. Boyce, VI-1-11 to VII-6-19, 1990—1994 (25; 5 to 11 traps); Fairfax Co., near An- nandale, V-8-14 to VII-17-23, 1982-1994 (247; 1 trap); Essex Co., 1 mi. SE Dunns- ville, V-13-29 to VII-10-27, 1991-1994 (75; 10 to 16 traps); Louisa Co., 4 mi. S. Cuckoo, V-28-VI-5 to VI-16-25, 1987-1989 (13; 4 to 12 traps). WEST VIRGINIA: Tucker Co., Fernow Experimental Forest, ca. 3 mi. S. Parsons, VI-10-19 to VII-19-28, 1991—1993 (5; 20 traps); Hardy Co., 3 mi. NE Mathias, VIH-5-18, 1994 (1; 2 traps). Vanhornia eucnemidarum is a parasit- oid of beetles of the family Eucnemidae (see Deyrup 1985). Eucnemidae are found in wood that has just begun to decay, mainly in beech and maple. Most of the collections were in traps in woods or wood edges, a habitat where the hosts would be expected. All of the Tucker Co. traps were set within a broadleaf forest. The largest collections, however, were from a trap adjacent to a woodpile in my backyard (Fairfax Co.). I found specimens in this trap each of the 13 years of oper- ation. We have a number of large silver maples (Acer saccharum Marsh.) around the house, and we frequently cut limbs and branches and add them to the wood- pile, thus there is wood in various states of decay. Little did I know that I was cre- ating a habitat suitable for the host beetle and its parasitoid as well. I thank the following for allowing field work on their properties: M. Bowers, C. Sacchi, and E. Connor, University of Vir- ginia Blandy Experimental Farm; J. and B. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Number of Specimens March April May June July Aug. Sept. Oct. Fig. 1. Seasonal flight activity of Vanhornia eucnemidarum in the mid-Atlantic states. Includes records from all sites from 1982-1994. Kloke, Louisa and Essex counties, Virginia; David R. Smith, Systematic Entomology and T. J. Henry and D. R. Miller, Hardy Laboratory, PSI, Agricultural Research Co., West Virginia. E.M. Barrows, George- Service, U.S. Department of Agriculture, Yo town University, Washington, D.C., provid- National Museum of Natural History, NHB ed specimens from the Fernow Experimen- /68, Washington, D.C. 20560. tal Forest, Tucker Co., West Virginia. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, p. 891 NOTE Polistes dominulus (Christ) (Hymenoptera: Vespidae) New to Maryland Polistes dominulus (Christ) is a common and widespread paperwasp in the paleartic region. The species is common in Europe and often goes under the name P. gallicus (L.) (Menke, Spheces 12: 21, 1986). In the Palearctic P. dominulus is known from North Africa, the warmer parts of France, Belgium, and Germany, and eastward to the Pacific Coast of China. Generally, the spe- cies is found in warmer and drier localities within its range. Polistes dominulus was first collected in the United States in 1980 at Cambridge, Massachusetts (Hathaway, Psyche 88: 169— 173, 1981). Since 1980, P. dominulus has been collected in Connecticut, New Jersey, New York, Ohio, and Pennsylvania (R. S. Jacobson, per. comm.). On 3 October 1994, I. B. Smith, Mary- land Department of Agriculture, and R. Ruschell, University of Maryland, exam- ined two colonies of P. dominulus on the College Park campus. One colony was at the top of a door jamb of a seldom used doorway and the second was on the housing of an exhaust fan at approximately ground level in an exterior stairwell. Over 100 adult P. dominulus were collected. This is a new State record. Polistes dominulus is a distinctive spe- cies, looking more like a yellowjacket (Ves- pula spp.) than any of the Polistes native to North America. It forms a typical paper wasp single comb in the open or in en- closed areas such as the open ends of pipes. Voucher specimens have been deposited in the Maryland Department of Agriculture and the U.S. National Museum of Natural History. Maryland Department of Agricul- ture Contribution Number CN86-94. C. L. Staines and I. B. Smith, Jr., Mary- land Department of Agriculture, Plant Pro- tection Section, 50 Harry S. Truman Park- way, Annapolis, Maryland 21401. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, p. 892 NOTE Transferral of Polycentropus timesis (Denning) Comb. Nov. from the Genus Neureclipsis (Trichoptera: Polycentropodidae) Polycentropus timesis (Denning) comb. nov. Neureclipsis timesis Denning 1948, Bull. Brooklyn Ent. Soc. 43: 119, fig. 1 dé. Polycentropus picicornis.—Blickle & Morse 1955, Bull. Brooklyn Ent. Soc. 50: 96-98 [New Hampshire specimens are P. timesis|. Recent collections of caddisfly material collected at Spruce Hole, Durham, New Hampshire, via Malaise traps, May—June 1990, by Donald S. Chandler, yielded spec- imens of Neureclipsis timesis Denning (1948), a species that previously was only known from the holotype collected in Am- herst, Massachusetts. Examination of the type and other specimens of WN. timesis shows that the hindwing of this species has M with only two branches, revealing that this species does not agree with diagnostic characteristics of the genus Neureclipsis McLachlan, and on this basis it is trans- ferred to the genus Polycentropus Curtis. Further examination of specimens from New Hampshire previously identified as P. picicornis by Blickle & Morse (1955), in the insect collection at the University of New Hampshire, shows that these speci- mens are actually P. timesis. This species is rather closely related to P. picicornis Stephens 1836, and differs in the male genitalia having preanal append- age with a long slender ventroposterior api- cal lobe and a short basodorsal lobe in lat- eral view, see figures of P. timesis in (Den- ning 1948) and P. picicornis in (H. Mal- icky, 1983, Atlas of European Trichoptera, The Hague). It is also noted that specimens of P. picicornis from Europe and western Canada exhibit slight differences in the male genitalia; the form from Canada was described by A. P. Nimmo (1986: 240-241, The adult Polycentropodidae of Canada and the adjacent United States, Qaest. Ent. 22). I am grateful to Oliver S. Flint, Jr., Smithsonian Institution, for his helpful sug- gestions, Donald S. Chandler, University of New Hampshire, for providing specimens examined, W. Pulawski, California Acade- my of Sciences, for the loan of the type, and P. Perkins, Museum of Comparative Zoology, and A. P. Nimmo for providing additional material. John S. Weaver III, Department of Plant Biology, University of New Hampshire, Durham, New Hampshire, 03824. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, p. 893 NOTE Replacement of a Homonymic Species Name in Anobiidae (Coleoptera) Up to 1977 the wood-boring beetles of the genus Xyletinus of this hemisphere con- sisted of 13 species from North America north of Mexico, and a single species from Mexico. In 1977 (White, Proceedings of the Entomological Society of Washington 79: 533) described as new ten species of Xyle- tinus, bringing the total of species known from North America north of Mexico to 19, and five from Mexico. The species of Xyletinus that occur in America north of Mexico are readily sepa- rated into two groups: the elongated, large eyed species (body 2.3—2.7 times as long as wide, eyes separated by 1.0—3.7 times fron- tal width of an eye), and the less-elongated, small eyed species (body 1.6—2.0 times as long as wide, eyes separated by 4—7 times frontal width of an eye). All six of the new species from America north of Mexico be- long to the less-elongated, small-eyed spe- cies. It has been called to my attention by Mr. Petr Zahradnik (Jiloviste-Strnady, Praha, Czechoslovakia) that the Mexican species Xyletinus cylindricus White (1977: 533) is a junior, primary homonym preoccupied by Xyletinus cylindricus Kofler (1970. Zeit- schrift der Arbeitsgemeinschaft Osterreich- ischer Entomologen. 22: 143-145). I here- by propose Xyletinus subcylindricus NEW NAME, as a replacement. Richard E. White, Systematic Entomolo- gy Laboratory, PSI, Agricultural Research Service, USDA, % National Museum of Natural History, Washington, D.C. 20560. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 894-895 NOTE Sassafras albidum: A New Host Plant Record for Larval Melanolophia signataria (Lepidoptera: Geometridae) from Central Virginia Larvae of Melanolophia_ signataria (Walker) have been reported to feed on a wide variety of woody plants in eastern North America. Recorded host plants in- clude: Abies spp. (firs), Abies balsamea (L.) Miller (balsam fir), Acer spp. (maples) Al- nus spp. (alders), Betula spp. (birches), Bet- ula alleghaniensis Britton (yellow birch), Larix laricina (DuRo1) K. Koch (tamarack), Picea spp: “(spruces);, “Picea “glauca (Moench) Voss (white spruce), Populus spp. (poplars), Quercus spp. (oaks) and Ul- mus americana L. (American elm) (Mc- Guffin. 1944. Canadian Entomologist 76: 124; Forbes 1948. Lepidoptera of New York and neighboring states. Part I]. Mem- oir 274. Cornell University Agricultural Ex- periment Station; Tietz 1952. The Lepidop- tera of Pennsylvania. Pennsylvania Agri- cultural Experiment Station, Pennsylvania State College, State College, PA; Covell. 1984. A field guide to the moths of eastern North America. Houghton Mifflin Co. Bos- ton, MA). Forbes (op cit) noted past con- fusion between larval host plants of M. sig- nataria and the closely related M. canadar- ia (Guenee) and questioned the validity of some host plant records for both species. In this note I report a new, verified host plant record for larval M. signataria from central Virginia. During early June 1991, I collected larval M. signataria from leaves of sassafras, Sas- safras albidum (Nutt.) Nees, growing in a forest edge near Lovingston, Nelson Coun- ty, Virginia, as part of a survey of the insect herbivore complex of S. albidum. Field-col- lected M. signataria larvae were confined to separate recloseable plastic sandwich bags containing fresh leaves of S. albidum to determine: 1) if larvae actually feed and develop on S. albidum: or 2) if captures were incidental on a non-food host. Captive larvae of M. signataria fed readily on leaves of S. albidum, pupated, and emerged as adults within one month of collection, indicating that S. albidum is an acceptable host plant for larval feeding and develop- ment. Cursory field surveys conducted in the same area during 1992 again noted the occurrence of larvae of M. signataria feed- ing on leaves of S. albidum, confirming the trophic association between insect and plant at this site. Melanolophia signataria was by far the least numerous member of the larval lepi- dopteran fauna on S. albidum at the Vir- ginia site (only three larvae could be col- lected for rearing in 1991), which was dominated by Caloptilia sassafrasella (Chambers) (Lepidoptera: Gracillariidae), Epimecis hortaria (EF) (Lepidoptera: Geo- metridae) and Papilio troilus L. (Lepidop- tera: Papilionidae). It is uncertain if the rel- ative rarity of larval M. signataria on S. albidum at the Virginia study site is due to the infrequence of the species in the area in general, to the presence of more desir- able primary host plants on which adults preferentially oviposit, or because larvae feed on such a broad array of host plants that they are widely dispersed across nu- merous hosts. I thank Dr. D. C. Ferguson and M. A. Solis, of the Systematic Entomology Lab- oratory, Agricultural Research Service, United States Department of Agriculture, Washington, D.C., for identifying reared adults of M. signataria. Dr. M. A. Solis (Systematic Entomology Laboratory) and VOLUME 97, NUMBER 4 one anonymous referee made constructive comments on the manuscript. Voucher specimens are deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. 895 Charles E. Williams, Department of Bi- ology, Clarion University of Pennsylvania, Clarion, Pennsylvania 16214-1232. PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 896-900 SOCIETY MEETINGS 999th Regular Meeting—January 5, 1995 The 999th Regular Meeting of the En- tomological Society of Washington was called to order by President John W. Neal, Jr. in the Naturalists’ Center of the National Museum of Natural History at 8:04 pm on January 5, 1995. The meeting was attended by 20 members and two guests. Minutes for the 998th meeting were read for outgoing Recording Secretary Dr. M. Alma Solis by incoming Recording Secretary Ms Darlene D. Judd. Minutes were accepted as read. President Neal called for officer reports. Reporting for Membership Chair M. Alma Solis, Dr. Nathan M. Schiff announced that no new membership applications had been submitted for December 1994. President Neal called for reports of old business. Committee member, Dr. John M. Heraty reported the 1000th meeting would be held at the Log Lodge in Beltsville on Ground Hog Day and wine and cheese would be served. Dr. Heraty elaborated on the recent trials and tribulations in the search for a guest speaker and announced that the Committee expected a firm com- mitment from a potential individual in the following week. President Neal announced that the 1000th meeting would have two speakers. A visiting speaker, as yet uncon- firmed and a speaker from the membership, Theodore J. Spillman. In the 1984 Proceed- ings of the Entomological Society of Wash- ington 86: 1—10, Mr. Spillman published an account of the first 100 years of Society ac- tivities and his presentation for the 1000th meeting will convey the meaning “‘of So- ciety.”” This concluded the reports of old business. President Neal called for new business, but there was none. President Neal called for the presentation of notes, exhibitions, and accessories. Pres- ident Neal noted for the first time in many meetings that Dr. Nathan M. Schiff had nothing to present to the membership. An ensuing discussion suggested Dr. Schiff’s memory lapse was due to a “‘seasonal-af- fected disorder.”’ Ms Jill Swearingen brought a copy of the “‘Identification Guide to Ant Genera of the World” by Barry Bol- ton (1994, $65.00), published by Harvard University Press, Cambridge, Massachu- setts. President Neal then queried the mem- bership on the possibility of an amended format for 1000th meeting. Dr. Raymond J. Gayné suggested maintaining the format of the Regular meeting for the benefit of vis- itors and guests. President Neal subsequent- ly instructed Program Chair Nathan Schiff to request the membership to bring unusual notes and specimens to this particular meet- ing and questioned whether a special an- nouncement should be sent to members. Dr. Schiff reported that a flyer announcing the speakers and other information would be sent to members, but those present should remind their colleagues to bring notes and exhibitions to the 1000th meeting. It was resolved that the format for the Regular meeting would be maintained for the 1000th meeting. Dr. Nathan M. Schiff, Program Chair, in- troduced the speaker for the evening, Dr. Paul J. Spangler, Department of Entomol- ogy, Smithsonian Institution, whose talk was entitled ““An Expedition to the Moun- tain of the Mist, Venezuela.”’ The presen- tation focused on a biotic survey conducted in 1985 of the Cerro de la Neblina in south- ern Venezuela. One hundred forty-four re- search scientists spent 45 days collecting, illustrating, preserving, and preparing spec- imens from variety of habitats. The slide presentation captured the varied landscape, dramatic views, unusual flora and fauna, various collection methods, and the day to day activities of field biologists. Investiga- tors collected 54,926 specimens and wrote 125 publications from this one expedition. VOLUME 97, NUMBER 4 Among the Insecta, 89 species were new to science, with the orders Coleoptera, Odo- nata, and Trichoptera producing the most new species. In addition to the presentation, Dr. Spangler displayed a drawer of insects, botanicals, and maps of the area. President Neal called for the introduction of new members and visitors. Mrs. Mignon Davis announced that a retirement party had been given by the Department of En- tomology for Dr. Oliver S. Flint at the Nat- ural History Building on 5 January 1995. Dr. Flint specializes on the order Trichop- tera and had been a Research Scientist with the Smithsonian Institution for 37 years. The meeting was adjourned at 9:28 pm. Refreshments were provided by Dr. Har- old Harlan. Darlene D. Judd, Recording Secretary 1000th Regular Meeting—February 2, 1995 The 1000th Regular Meeting of the En- tomological Society of Washington was called to order by President John W. Neal, Jr. in the USDA Auditiorium, Building 003 in Beltsville, Maryland, at just after 8:00 pm on February 2, 1995. The meeting was attended by 60 members and 55 guests. President Neal gave a short welcoming address to the audience. His presentation focused on the organizational meeting of the Entomological Society of Washington held on February 29, 1884, making our So- ciety the 4th oldest entomological society in America. President Neal brought with him a copy of the announcement for that meeting held at the home of C. V. Riley. The first order of business by President Neal was to postpone the reading of the minutes from the 999th Meeting of the En- tomological Society of Washington to the regular March Meeting. There were no ob- jections to the amended by-laws and it was tabled without a motion. President Neal called for reports from officers. Member- ship Chair Dr. M. Alma Solis announced that no new membership applications had been submitted for January 1995. 897 President Neal called for reports of old business. In behalf of the Society, President Neal acknowledged Drs. W. Steven She- phard, Nathan Schiff, and John Heraty for their skillful organization and planning of activities encompassing the 1000th meet- ing. President Neal then called for presenta- tion of notes and the exhibition of speci- mens. The President brought for exhibit two books: the “‘Fourth Annual Report to the State of Missouri,” by C. V. Riley and “A History of Economic Entomology,” by L. O. Howard (1930). As a personal recomen- dation of the latter, Dr. Neal stated “it will absolutely make you forget about what is going on in the real world.” Dr. Nathan M. Schiff, Program Chair, in- troduced the first speaker for the evening. Theodore J. Spilman, a member of our So- ciety spoke on “Vignettes of a Thousand Meetings.”” Mr. Spilman acted as the au- dience’s tour guide for a _ historic stroll through previously published reports in the Society minutes. Highly entertaining and craftily delivered, the personalities of the founders and flavor of the meetings left one with impression that a good time was had by all in attendance at those early meetings. Mr. Spilman closed his presentation by an- nouncing that the 2000th meeting of the Entomological Society of Washington would be held on March 4, 2106. Dr. Pam Henson, Smithsonian Archives, introduced the second speaker for the eve- ning, Dr. David L. Hull from Northwestern University. He spoke on “‘Why Scientists Behave Scientifically.”” The talk focused on aspects of science which he percieves are done well by scientists. President Neal normally would have re- quested at this time that visitors stand and introduce themselves; however, he broke with tradition and requested that the follow- ing individuals stand and be acknowledged: Honorary Members, followed by Past Pres- idents and finally Past and Present Officers and Chairpersons. (By the secretary’s ac- count, only the visitors remained seated.) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fo the Society he made the following sa- lute: You have served science and society for over a century and now you have met through a thousand meetings in ways never dreamed by your founders. May you con- tinue to be a servant of society and be a shared experience and opportunity for those who find insects and their kin to be so com- pelling. To the Entomological Society of Washington, may you always find your members as curious and excitable, and as a Scientific society, be as strong and as vi- brant at your 2000th meeting as you are to- day. Finding no other business the meeting was adjourned at 9:46 pm. A lavish assortment of wines and cheeses were provided courtesy of the Entomolog- ical Society of Washington. Darlene D. Judd, Recording Secretary 1001st Regular Meeting—March 2, 1995 The 100Ist Regular Meeting of the En- tomological Society of Washington was called to order by President John W. Neal, Jr. in the Naturalists’ Center of the National Museum of Natural History at 8:10 pm on March 2, 1995. The meeting was attended by 16 members and seven guests. Minutes for the 999th and 1000th meetings were read by Darlene D. Judd. Minor corrections were suggested for the 999th minutes and both sets of minutes were subsequently ac- cepted. Reports of officers and committees were read. Reporting for Membership Chair M. Alma Solis, Dr. Nathan Schiff announced there were three new members William A. Bruce, Lodewyk Kuenen, and J.R. Vock- eroth. New member, Lodewyk (Bas) Ku- enen, was present and was introduced to the membership. President-elect Dr. Ralph P. Eckerlin re- ported the Entomological Society of Wash- ington’s Spring Banquet will be held Wednesday, 31 May 1995 at the Associates’ Court in the National Museum of Natural History. The Banquet will be attended this year by members of Maryland Entomolog- ical Society and an attempt was made also to invite the Chesapeake Association of Professional Entomologists. Individuals with suggestions for a possible speaker for the Spring Banquet should contact either Dr. Nathan M. Schiff or Dr. Ralph P. Eck- erlin. President Neal called for reports of old business. He thanked Drs. W. Steven Shep- ard, Nathan M. Schiff, and John M. Heraty for their assistance in organizing the 1000th Meeting of the Entomological Society of Washington. A call was made for new business. Pres- ident Neal suggested that the by-laws for the Society be reprinted in the Proceedings of the Entomological Society of Washing- ton. The by-laws were last published in 1987 and appear in need of revision. Vol- unteers were sought to review the current by-laws prior to the April Regular Meeting. Two issues to be discussed at the Executive Meeting are the cost of a life-time mem- bership and inclusion of the June meeting in the list of Regular Meetings of the So- ciety. An Executive Committee meeting will be held 6 April 1995. President Neal received a letter from a member of the Society who suggested that the site of Regular Meetings should be al- ternated between Washington and Mary- land. This prompted a membership discus- sion on possible sites for future meetings. It was decided Dr. Nathan M. Schiff would mail a questionnaire on the subject to area members (approximately 200) between now and next fall. Notes, exhibitions, and specimens were called for. Dr. Paul Spangler brought a book entitled “‘The Lucanid Beetles of the World,’ edited by Hiroshi Fujita (1994), published by Mushi-Sha, Japan. Dr. Wil- liam E. Bickley reported the death of Dr. J. Franklin Yeager, a noted insect physiologist in January of 1995. Dr. David Furth an- nounced a retirement party would be held for Mr. Bill Rowe of the Department of En- tomology, National Museum of Natural History on 29 March 1995. VOLUME 97, NUMBER 4 Dr. Nathan M. Schiff, Program Chair, in- troduced the speaker for the evening, Dr. Martha Weiss, Department of Botany, Uni- versity of Maryland, whose talk was enti- tled “‘Color Learning in Butterflies.” Two guests were present, Roxanne Al- gart, Department of Entomology, Univer- sity of Massachusetts, and Rick Turcotte, Bee Research Laboratory, USDA, Belts- ville. Refreshments were provided by Dr. Wil- liam E. Bickley. The Meeting was adjourned at 9:24 pm. Darlene D. Judd, Recording Secretary 1002nd Regular Meeting—April 6, 1995 The 1002nd Regular Meeting of the En- tomological Society of Washington was called to order by President John W. Neal, Jr. in the Naturalists’ Center of the National Museum of Natural History at 8:04 pm on April 6, 1995. The meeting was attended by 21 members and five guests. Minutes for the 1001st meeting were read by Darlene D. Judd, and subsequently accepted. Reports of officers and committees were read. Membership chair M. Alma Solis an- nounced that there were no new members. President-elect Ralph P. Eckerlin an- nounced that the Chesapeake Association of Professional Entomologists were interested in attending the Spring Banquet. It was re- affirmed that the Spring Banquet will be held Wednesday, 31 May 1995, at the As- sociates Court in the National Museum of Natural History. The speaker will be Dr. Gary Umphrey, University of Western On- tario, who will speak on “‘Analyzing the Literature of Insect Fiction: Themes, Dreams, Schemes and Screams.” The Mas- ter/Mistress of ceremonies will be Dr. Man- ya B. Stoetzel. President Neal announced that the exec- utive committee decided unanimously to elect Drs. J. Richard Gorham and Frank W. Meade as emeritus members. It was also de- cided that a committee would be set up to examine the Society’s by-laws dealing with such items as life membership fees. 899 A call was made for new business. Dr. Manya B. Stoetzel announced that Dr. Rob- ert D. Gordon of the Systematic Entomol- ogy Laboratory would be retiring at the end of April. Notes, exhibitions, and specimens were called for. Warren E. Steiner exhibited some live specimens of the iron clad beetles, Zo- pherus holdemani (Zopheridae), that were recently collected in Texas. Dr. Nathan Schiff introduced the speaker for the evening, Dr. Gary Miller, USDA- ARS Systematic Entomology Laboratory, Beltsville, who gave us an enlightening pre- sentation on “‘Historical Natural History: A Study of Insects in the Civil War.’ Dr. Mil- ler has researched various aspects of the in- volvement of insects in the war ranging from the number of deaths attributed to in- sect-borne disease, well-known entomolo- gists who were enlisted on either side of the conflict (although there did appear to be a preponderance of northerners), games played by the infantry using insects, and the plagues of flies, lice, and fleas that caused more than one member of the audience to scratch in sympathy. Roughly 380,000 sol- diers died from disease during the war; far more than the 200,000 that actually died in battle. Over 25% of the union soldiers con- tracted “‘simple intermittent fever’? or ma- laria. At least one anti-malarial remedy of- fered to southern troops included 30 parts dogwood bark, 30 parts poplar bark, and 40 parts willow bark, which was served with whiskey at 45 degree strength. The willow bark that contains salicylic acid may have offered some relief, but it was probably the whiskey that helped to raise the spirits of the soldiers. Four guests were presented to the mem- bers: Mellissa and Brian Miller, Denise Whitman of the Department of Inverte- brates, Smithsonian Institution; Kim Nesci, Department of Entomology, University of Maryland; and Ian Lowe, Fredrickburg and Sotsylvania National Military Park. Refreshments were provided by Dr. John W. Neal, Jr. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The meeting adjourned at 9:16 pm. John M. Heraty for Darlene D. Judd, Recording Secretary 1003rd Regular Meeting—May 4, 1995 The 1003rd Regular Meeting of the En- tomological Society of Washington was called to order by President John W. Neal, Jr. at the National Visitor Center (Beltsville) at 8:05 pm on May 4, 1995. The meeting was attended by 22 members and 19 guests. Minutes for the 1002nd meeting were read by Dr. John M. Heraty for Recording Sec- retary Darlene D. Judd, and subsequently accepted. Reports of officers and committees were called for. Membership chair M. Alma Solis announced that there were two new mem- bers: Stephen Gamari (University of IIli- nois, Urbana, IL) and Eric L. Johnson (Aphis-PPQ, Seattle, WA). President-elect Ralph P. Eckerlin an- nounced the menu of the Spring Banquet was finalized. The banquet date and speaker were reaffirmed. President Neal announced that it was de- cided that Dr. Norman E. Woodley and Dr. David Smith would review the Society’s by-laws and present their report to the So- ciety in October. No new business was reported. Notes, exhibitions, and specimens were called for. Dr. Manya B. Stoetzel made a presentation of some “dark black things”’ that she identified as conifer-infesting aphids, possibly of the genus Sinara. Dr. Nathan M. Schiff commented on the attrac- tion of yellowjackets to aphids on conifers in the fall near the bee research laboratories. Dr. Schiff raised the question about where the meetings would be held next fall. Dr. Neal commented that they would probably be in the Waldo Schmitt room at the Na- tional Museum of Natural History. Dr. Schiff introduced the speaker for the evening, Dr. James Marden of Pennsylvania State University, who gave a fascinating talk on ‘“‘Surface Skimming Stoneflies: Testing the Evolutionary Hypotheses of In- sect Flight.’” Dr. Marden has been focusing on the functional morphology of Taeniop- terygidae (winter stoneflies). Taenioptery- gids are essentially flightless but use their wings to propel them across the water by wing flapping or by raising the wings into a vertical position and using them as sails. Dr. Marden proposed that skimming is a more plausible intermediate step toward de- veloping functional wings for flight than the use of gill flaps for gliding. His work was supported by various experiments on ob- serving skimming behavior, changing pa- rameters of the wings such as length, and characterizing the morphology of wing ve- nation within a phylogenetic perspective to explain the ancestral origin of this behavior. Dr. Marden proposed that the ability to fly may have developed more than once, but the capability for flight may have developed in a surface-skimming ancestor. Seven guests were presented to the mem- bers. Dr. Stoetzel introduced Dr. Curtis Sa- brosky and Ms Louise Russell. Ms Russell noted that at age 90 she had been a member of the Society for more than 60 years, many more years than the age of most attending members. Refreshments were provided by Dr. Man- ya Stoetzel. The meeting was adjourned at 9:21 pm. Dr. John M. Heraty for Darlene D. Judd, Recording Secretary PROC. ENTOMOL. SOC. WASH. 97(4), 1995, pp. 901-906 PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON Volume 97 OFFICERS FOR THE YEAR 1995 President President-Elect Recording Secretary Corresponding Secretary Treasurer Program Chairman Membership Chairman Custodian Editor Associate Editor Book Review Editor Special Publications Editor Past-President John W. Neal, Jr. Ralph P. Eckerlin Darlene D. Judd Hollis B. Williams Norman E. Woodley Nathan Schiff M. Alma Solis James Pakaluk Thomas J. Henry David R. Smith Gary L. Miller A. G. Wheeler, Jr. Paul J. Spangler Published by The Society WASHINGTON, D.C. 1995 TABLE OF CONTENTS, VOLUME 97 ARTICLES AALBU, ROLF L., THEODORE J. SPILMAN, and KIRBY W. BROWN—The systematic status of Amblycyphrus asperatus, Threnus niger, Pycnomorpha californica, Emmenastus rugosus, and Biomorphus tuberculatus Motschulsky (Coleoptera: Tenebrionidae) ............ 481 ADAMSKI, DAVID—Review of the Blastobasidae of the Republic of The Seychelles (Lepi- doptera: Gelechioidea)) 22. .....2.0..0..000. secs tecscseccewacsecscceecesstecstecce co ceccceseesnsesens 489 AMORIM, DALTON De SOUZE—See GRIMALDI, DAVID A. ................0 0. cece eee eee eee 561 ASQUITH, ADAM—Loulucoris, a new genus, and two new species of endemic Hawaiian plant bug (Heteroptera: Miridae: Orthotylinae) .......... SEAR BRED Op co nO Good EUs ORGOaU DE Reno CHOROnbEEaEe 241 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON BARRACLOUGH, D. A. and TERRY A. WHEELER—Three new species of Camillidae (Dip- tera) from the southwestern Nearctic Region, the first species of the family described from the NEW. WOEId soc cescicaasicmmecoe aaseine ds miamscinnecee chicloset etince seesG ee seek eee aeee eG eee 737 BECKER. VITOR ©:—See MILLERS SCOT! Be siccc cease eee eee ee Cee eee eeeeeeeeeeee 117 BOHART, RICHARD M.—A review of New World Entomognathus with descriptions of seven new, species (Hymenoptera, Sphecidaey Crabronm) seen. cee toes eee a nace see secrete 506 BOLDT. PAUL E=See GAGNE, RAYMOND J) 2.52005 le eee 767 BRAILOVSKY, HARRY—New genus and new species of Colpurini (Heteroptera: Coreidae) from, the: Philippine Republic) cog coe jerassec01se)e arorecimins yess e612 ores «sorties aera ise a ee eee 250 BROWN, KIRBY W—See AALEBU, ROLE We % ietccmdecce oem saeecee eet sae at seer 481 BYERS, GEORGE W.—New species of Nearctic snow crane flies of the genus Chionea (Dip- ternal Dipulidac)) pave cretstocin cui otais onic aratae dle ora w ayeis nr sete o 1s wi win (is ols sevavols ayavevsyajalee stale} evetsjnia\siove slo's’ eioievele)sjelostarsysy< 508 CARROLL, J. E—Laboratory evaluation of predatory capabilities of a common wolf spider (Araneae: Lycosidae) against two species of ticks (Acari: Ixodidae) ........................04. 746 CARVALHO, MARCIO ANTONIO de—The making of an entomologist ....................... 236 CARVALHO, MILZA—José Candido de Melo Carvalho—A wife’s view .............--..0005- 231 CASSIS, GERASIMOS—A reclassification and phylogeny of the Termatophylini (Heteroptera: Miridae: Deraeocorinae), with a taxonomic revision of the Australian species, and a review ofethestnbalclassificationtof the: Deraecocorinac! Fe-- eee een ere ce eee ene eee eee nee 258 GCHRISTMESANNE=Seer AT BINS TO HINGED sa coe cas acacia nacee eee ae ee nee eRe en ener 90 CEAYTON. DALE; E:—SeevPRICESROGERID 7) Bama eee eee en aan oan eee eee 659 DIETRICH, CHRISTOPHER H. and STUART H. MCKAMEY—Two new Neotropical tree- hopper genera and investigation of the phylogeny of the subfamily Membracinae (Homoptera: IMembracidae) ssnc yoste cic as cist eee eh Fe aera RE oho Se enol Rete ee Clos oie ctartaierns Bisa een ies 1 EPSTEIN, MARC E.—False-parasitized cocoons and the biology of Aididae (Lepidoptera: ZY SAC INOIMEA| 5512 aah slays sles Oats aiciers Sere ales dla /slote aleva-siy els elalerelatats ciers tn tle areas roeyetaictoe el atste ata oiereeta elena ae 750 EVANS, GREGORY A., MICHAEL E. SCHAUFE MOH LENG KOK-YOKOMI, and RAY- MOND K. YOKOMI—A new species of Aphelinus (Hymenoptera: Aphelinidae) that para- sitizes the spirea aphid, Aphis spiraecola Patch (Homoptera: Aphididae) ...................... 17 EVANS, HOWARD E.—A reconsideration of the cylindricus group of the genus Anoplius Pufours(ymenopteray Pompilidae)) fies -sassceeccee ores oee comes ecinseiecccinermense atte eee eer ee Tad FERREIRA, P. S. F—José C. M. Carvalho: A student’s remembrance ....................0.0000% 240 FERREIRA PSS. ——See: SCHABFNERS I. Gs 2 oSastebeacioce «tele Joldeciste acta iceteiae cn tctete ote cio srereteietars 373 FITZGERALD, SCOTT J. and BORIS C. KONDRATIEFF—A review of the mydid genus Pseudonomoneura Bequaert (Diptera: Mydidae), with the description of two new species .. 22 FLOWERS, R. WILLS—Hermesia Lefevre, a resurrected genus of Neotropical Eumolpinae (Coleoptera Chrysomeli dae) eo mnt .cts steciciee Se see se os oe teestiee oe esi ne Saree treme enters 35 FLOYD, MICHAEL A.—The larva and pupa of the caddisfly species, Helicopsyche paralim- nella Hamilton @irichoptera:Helicopsychidae)! sarc. sce. cee sees sees o sictee ela store seieisetetetsieloitetet= 46 FRANKS, Ji Sees, (Gays hrs as sae cece Saito ae teeter selec em viaisiea ya ewels cisioecetie slow earsose eee 582 FROESCHNER, RICHARD C.—Review of the New World lace bug genera Acanthocheila Stal and Garvalhotingismew-cenus: (Heteroptera: Mimgidac)) 2. ...qesesecues ee eee eects aaose B31 FROESCHNER, RICHARD C.—Nicholas A. Kormilev: A list of his entomological publica- tions*and). proposed! itaxar ce saacccses caci- sane sechinec usm Monaaiecease wee Aetoue snore Pea eee eee eee SyIIS) GAGNE, RAYMOND J. and PAUL E. BOLDT—The gall midges (Diptera: Cecidomyiidae) of Baccharis*spp. (Asteraceae) incthe WnitedsStates® Sst22ss-e ce scnce sce sees cee e ease 768 GOEDEN, RICHARD D., DAVID H. HEADRICK, and JEFFREY A. TEERINK—Life history and description of immature stages of Valentibulla californica (Coquillett) (Diptera: Tephrit- idae) on Chrysothamnus nauseosus (Pallas) Britton in southern California .................... 548 GOEDEN, RICHARD D., DAVID H. HEADRICK, and JEFFREY A. TEERINK—Life history and description of immature stages of Urophora timberlakei Blanc and Foote (Diptera: Te- phritidac)jonsnativeAsteraceaeinysoutherni Californias cea... n..aeeeeeeno eee aeeeeeeen eae eer 779 GOULET, HENRI and DAVID R. SMITH—Four new sawflies from eastern North America, three species of Tenthredo and one of Dolerus (Hymenoptera: Tenthredinidae) .............. 50 GRIMALDI, DAVID A. and DALTON DE SOUZA AMORIM—A basal new species of Ol- biogaster (Diptera: Anisopodidae) in Dominican amber, and its systematic placement ...... 561 GROGAN, WILEIAM(L.. JR:——See-sMcKEEVERS STURGIS. cisa. . oases assis soca des acloe es te 199 HAGAN, DANIEL V.—See MCKEEVER, STURGIS” 3: ccasaceaestrictne see nessa erecta 799 VOLUME 97, NUMBER 4 903 HARRISON, T. and S. PASSOA—Mirabilis-feeding Heliodines (Lepidoptera: Heliodinidae) in centralellinois: wathvdescriptionofea new Species) 2.22 -225---5--ess-- oes eee ao ees so ies os eee 63 HBADRICKSIDAVIDIEL—See|GOEDEN RICHARD ID Maen tsaseee seecen ceeeceriee ec ereee ease: 548 HEADRICK DAVID Hi—See1GOEDEN, RICHARDIDS sconsnseteeet tems deateeecce-mcee sere <-e2 779 HEISS, ERNST—Aractocoris perneri, n. sp., a new apterous Carventinae from Jamaica (Het- eropterawATadidae)) see sca wis eisisiseceinve tice wins ov nays ceie orcas Soe ee me eee erase eloleimiet aa etI assests se 71 HENRY, THOMAS J.—Proboscidotylus carvalhoi, a new genus and species of sexually di- morphic plant bug from Mexico (Heteroptera: Miridae: Orthotylinae) ......................... 340 HENRY, THOMAS J. and A. G. WHEELER, JR.—Forward ‘commemorative Carvalho USSU peers chee siete eg aR eur TS RPS Panicle re lege Seceiciatcre wis: wera cevesciay Nag tera ec eieleia Setar omrers teeta eee ererereeeies 229 HEYDON, S. L.—The North American species of Systasis Walker (Hymenoptera: Pteromali- (GRYE)) “oScis tan aC OOOO ae Be Go Gnn co dos COT ET UC AC OG REC CHET ne SRP HE a REO REEs ARE. Boome ant De SOBA GG nes 569 HOEBEKE,E: RICGHARD—SeeswHEERER A] GIRS {i...-0-nececeeceeeee ace cere eee eee 701 HU, G. Y. and J. H. FRANK—Structural comparison of the chorion surface of five Philontus Sspeciesn(ColeopterasStaphylimidae) ere. secre tech onesie seins oie sels ecsintie els ae see seme erro blew teeter 582 IRWIN MICHAEL, E:—See\ WEBB DONALD W., iis.-s..c2- + ce. scocceeneescnedescenoeseeeeneens 197 IVES MICHABIEA:—=S ce MIL EERSKERIEY By rnc jects crices ccc see neenecninee ste teecee eens 833 KERZHINERS I Me—See SGHUBAIRAND AW We) toa. warn ce aneenacee ses cee ss Socesereeneeteeenont 379 KIMSEY, LYNN S.—New amisegine wasps from southeast Asia (Hymenoptera: Chry- SIGHS) reese eerste OITA aI Pe ac Tote Seer Nelovns aia sis atesavetebeitistud shots als leave lolannevaa areas cle Sete mea ieee 590 KOK-YOKOME MOH LENG— Seer EVANS: GREGORY Al 27.52 22255-th eee eeeesnee oeeee eee ee 17 KONDRATIEFESB ORIS{C—=SeelEMIZGERAEDS SCO J. ii osc-2ee eo sense -a- cece 22 KONDRATIEFE, BORIS C. and CHARLES H. NELSON—A review of the genus Remenus Ricker (Plecoptera: Perlodidae), with the description of two new species. ..................+.. 596 KOVAC] DAMIR==SeeSTONEDAHEN GARYOM sao xscaosseis ss cise siaiote winiteoeia ee sais eee emcees 427 KROMBEIN, KARL V.—Systematic notes on some Sri Lankan Scoliidae (Hymenoptera: Acu- [EEKED) 3 53s Cap ahd cen Sie bbs ot HaC IRCA OOS Co CeLic EIR IOS GONG RC Ere er er Beer fa carae tty on SAN) Mer ek Mere even rea at Ay HY KROMBEIN, KARL V.—See SCARBROUGH, AUBREY G.. .................... 0 eceeeee seen 689 KROMBEIN, KARL V. and BETH B. NORDEN—Notes on the behavior and taxonomy of Megachile (Xeromegachile) brimleyi Mitchell and its probable cleptoparasite, Coelioxys (Xer- ocoelioxys) galactiae Mitchell (Hymenoptera: Megachilidae) ....................... cece eee eee 86 LANDRY, JEAN-FRANCOIS and DAVID L. WAGNER—Taxonomic review of apple-feeding species of Phyllonorycter Hiibner (Lepidoptera, Gracillariidae) in North America ........... 603 LATTIN, JOHN D., ANNE CHRISTIE, and MICHAEL D. SCHWARTZ—Native black grass bugs (Urbisia—Labops) on introduced wheatgrasses: Commentary and annotated bibliography @icmiptera:srleteropteras Miridac) ian -ecccm sec eancis cece aera saetctsroe a ele azatotnie sisteltonisis sieieiataiaieooreete 90 EFINDSKOGSIPER—SeeSCHUE RAND AT ey areecci oe coe eine aisls tales stellate ielateiais eisisreialsioisiotels 379 MALDONADO CAPRILES, J.—Notes about the Old World genus Hexamerocerus Reuter (Heteropteras Reduyidaes Ectrichodimae) yee ccc ceee econ = nace cis eerie eeiteis ane eeeie cee 112 MALDONADO CAPRILES, J.—Notes about Vescinae, a key to the world genera, and descrip- tioniof two new, Chopardita’ (Heteroptera: Reduviidae)| 2.522 s5-ce= seen oe sce Pome ee tee ee eee 626 MALDONADO CAPRILES, J.—New Nearctic species of Oncocephalus Klug (Heteroptera: Reduviidae:4Stenopodaim ae) ieee yasieiseteters solos nie oreia ote slo) ake os eteacis otss o nich cyeratalste racials Sarerstinseysbstarsiyatos ePete 791 MALDONADO CAPRILES, J. and GEORGE O. POINAR, JR.—Reuteroscopus carvalhoi n. Spa newtossil plant bug: (Heteropteras Minidae: -Phylinac)) yi ects. cnc ecietain e's sels oie olelototorsrelas 346 MATEISS WAYINE) N:=SeewRAOMNAVIIND fsetet cece ssesnacic .ocsenbe semen «se ctsicscbrs cincaetieetlseeet 666 MeEKAMEN.SDUART, P—See DIETRICH, CHRISTOPHERVHE s:-tinescss-e:-eecccestee ee sect er 1 McKEEVER, STURGIS, DANIEL V. HAGAN, and WILLIAM L. GROGAN, JR.—Compar- ative study of mouthparts of predaceous midges of the tribe Palpomyiini (Diptera: Cerato- pogsonidae)ErombpEastemnWnitedsStates: Weare caer selects cleleterarstetslslelele ote stoleraiatosorsislaseteisteracieiesete sia tetars 799 McCPHERS ONE Js E:—See WOOD) DIANE Me (ieee ae ceitiatote sce ate sforataete crcletesiotafote cteteleiele ote terale e scctti sale TMT, MILLER, KELLY B. and MICHAEL A. IVIE—Enallagma optimolocus, a new species of damselfly from Montana (Odonata: Coenagrionidae) ................. cece eee cee e eee eee e ee eee 833 MILLER, SCOTT E., VITOR O. BECKER, and RAUL VELEZ-ANGEL—Podalia bolivari (Lepidoptera: Megalopygidae): A highly sexually dimorphic Neotropical pest ................ 117 MORAN, MATTHEW D.—Intraguild predation between sympatric species of mantids (Man- (KOs SAIS IM beta (ite RYO) eae oserc cela aos en BORO e GEE OPER EhE OD ELDGES aH adoos ab cad ce cb oUs aoe Unda n ed SE ae sar 634 NELSONS CHAREES, H:——SeesKONDRATIEFF BORIS! €3 eo. seer ctte cleeco celts eciciisiie cielo wnt 596 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON NEUNZIG, H. H.—New genera and species of Peruvian Phycitinae (Lepidoptera: ea 21000 (=) en ene Son OO ARTIC Mac oA cage oon ocatnd sccdpaS danpaanicacosuoocaraosnieaon 639 NORDEN; BETH B.—See: SCARBROUGHFAUBRENG Gi ian sceee on teeta ee eee een eee 689 NORDEN, BETH B:=SeesKROMBEIN KARI ya-ce esac nee ee eee ee eee eee eee eee EEe 86 PASSOA;(S:=SeesHARRIS ON dao ee rs Soc s aces ace Seon eee ene Ee ene 63 POINAR, GEORGE ©O;, JR:—See MALDONADO!CAPRIELES, J. sc cccec cece sscsenss eee > eee 346 POLHEMUSSDANEA=—sec POLHEMUSSIOHIN #ies eee none ne ee eee ene ene 350 POLHEMUS, JOHN T:—The identity and synonymy of Nepa fusca Linnaeus, 1758 (Heter- Opterar N|pid ac)! Hesse Selassie aca Sarate oicle clete's locate ainratatalelovstataleyoetatelece ote aS uls laa ote tales ees ee RRL cerare 123 POLHEMUS, JOHN T.—Nomenclatural and synonymical notes on the genera Diplonychus Laporte and Appasus Amyot and Serville (Heteroptera: Belostomatidae) ...................... 649 POLHEMUS, JOHN T. and DAN A. POLHEMUS—A phylogenetic review of the Potamobates fauna of Colombia (Heteroptera: Gerridae), with descriptions of three new species .......... 350 POLHEMUS, JOHN T. and ROBERT W. SITES—The identity of Pelocoris biimpressus Mon- tandon and synonymy of Pelocoris species in the southwestern United States (Heteroptera: INaticoridae)) cps sic Ss inersyatacctstsiarelew slace ciets QOmiore eters Memeretote ao eloroses ale otatanave tetera el cia aleve ata trols MVS STSre 654 POLHEMUS, JOHN T. and PAUL J. SPANGLER—A review of the genus Stridulivelia Hun- gerford and two new species (Heteroptera, Veliidae) from South America .................... 128 PRICE, ROGER D. and DALE H. CLAYTON—A new genus and three new species of chewing lice (Phthiraptera: Philopteridae) from Peruvian ovenbirds (Passeriformes: Furnariidae) ..... 839 PRICE, ROGER D. and ROBERT M. TIMM—The chewing louse genus Aoftiella (Phthiraptera: Gyropidae) from South American night monkeys, Aotus (Primates: Cebidae) ................ 659 RAO, NAVIN and WAYNE N. MATHIS—A revision of the shore-fly genus Rhysophora Cres- sond(DipterasE phy. dridae) tacks fs ol eos eee hae atelier arta sae roreeta ee erates ae eee ee 666 RIDER, D. A. and L. H. ROLSTON—Nomenclatural changes in the Pentatomidae (Hemiptera: 1 (10100) 011) 02) ener ar Anon Caer R AT HOE ARES HAC BOAR OnE CoE ADC Annis Senet a Pe ea UAT aaDEGACe pra aoccice MAD iodo 845 ROESTONMESH=—See RIDE RMDPAS ¢ danceacsnan tina telecasts aes cira ane Eee e TA ECR EERE aT 845 SCARBROUGH, AUBREY G., BETH B. NORDEN, and KARL V. KROMBEIN—A new species of Townsendia Williston (Diptera: Asilidae) from Florida with notes on its association with Perdita graenicheri Timberlake (Hymenoptera: Andrenidae) ...................2222220000- 689 SCHAFENER, J. C. and P. S. EK FERREIRA—Carvalhoisca, a new genus of Orthotylini from Mexicoa(Miridaecieteroptera) maanaaroccmeeeece ce etachi cece aaa inceeacits Satan enone eee eee Si7/8) SCHAUEE MICHAEL FE Sce EVANS GREGORYCAS oie ne ee dae oe Seen eee easel eae i7/ SCHUH, RANDALL T., PER LINDSKOG, and I. M. KERZHNER—Europiella Reuter (Het- eroptera: Miridae): Recognition as a Holarctic group, notes on synonymy, and description of almnew species, ‘Europiella carvathoi, from NorthvAmernica | s7.2422-. 6 2 sdeeee eee acne 379 SCHWARTZ; MIGHABE;D——See, MATIEIN:, JOHIN D2: Fak. cta toric tee eee neon seen eee 90 SCHWARTZ, MICHAEL D.—Metasequoiamiris carvalhoi, a new genus and species of conifer- inhabiting Mirini from China (Heteroptera: Miridae: Mirinae) .................2...ccseeeeeeeees 401 SCUDDER, G. G. E.—The first record for Bothynotus pilosus (Boheman) (Hemiptera: Miridae) inbthes NEAarchic URE St ON 5.25 satis ssiod ata lee Seen ree aes SR ee eee 396 SHAW, SCOTT RICHARD—A new species of Centistes from Brazil (Hymenoptera: Braconidae: Euphorinae) parasitizing adults of Diabrotica (Coleoptera: Chrysomelidae), with a key to New Worlds PECIES Heys ae SEAMS A RH racirass ctara ce As eT tes ec ee raP a tee ae ere eRe ES 153 SITES TROBERT \W.—see-POLHEMUSIOHINUR ites ae nen ascease see ecee ten eee eens 654 SLATER, JAMES A.—New genera and species of Rhyparochrominae from West Africa (He- miptera; ley eacidae) described iin honoriof JiC7 Me Carvalhon 25: casccceen- sete keane cee oee 409 SMITH] DAVID «i= See; GOULET HENRI) Wiest oeet tat detea caer tence sen cis seen eeee ea e eae ee 50 SPANGEER: PAUL Jc—See; POLHEMUS STORING 2235 iacacnee moss aee acetone dae ae eeeee eee 128 SPIEMAN; THEODORE J.—See AALBURROLE Ls cf ceaccetnnee sane neces a teeae aoe ae serene 481 STAINES, C. L.—Francisco de Asis Monrés: A perspective ..............ccecccccecceseeecceeeees 856 STONEDAHL, GARY M. and DAMIR KOVAC—Carvalhofulvius gigantochloae, a new genus and species of bamboo-inhabiting Fulviini from West Malaysia (Heteroptera: Miridae: Cy- lapinae)) exe sides soe Poe hea kis sleeves wisi sed os eee Ra oee Seater RES SC CE OSE A a hecaeee 427 SURDICK, REBECCA F—New western Nearctic Sweltsa (Plecoptera: Chloroperlidae) ...... 161 TEERINKG JEFFREY ‘A‘—See: GOEDEN; RIGHARD DS eestetan ne eeee ea teen eae 548 TEERINKS, JEFFREYCA.— See GOEDEN;, RICHARD Dt a. Sijener tee ar nieenienice see nen eet er Ce) THORPE, KEVINCW:==See: WEBB; RAL PHU ire. thee ten ot ean haa emtiad eee ee eee 695 VOLUME 97, NUMBER 4 TIMMEROBERTIM:—SeerROGERSD BPRIGCE pre eee dae isis este eileeteicts oe sie tee aw ine seer ee TOGASHI, ICHIJI—Description of Neocolochelyna hakusana, sp. noy., and its larva (Hyme- nopteraydenthredinidae) stromiPapant Eejee ete selene crates. eerecte let ieiocetetstal clereisictelers loreie cto ssietes citer VALENTI, MICHAEL A. and RICHARD S. ZACK—Lepidoptera associated with greenleaf manzanita, Arctostaphylos patula E. Greene (Ericaceae), in Shasta County, California ...... VEEEZ-ANGELWRAUE—-SeerMIBEE R= S CO Minbar ate sp saci cicleertilecradelee si lelee otic acct a clears VOEGTLIN, DAVID—Notes on the Mindarus spp. (Homoptera: Aphididae) of North America waithidescriptions1OfitwomewssSpeCleSmmrmeacae ce eteecm sti nae tencie cisternae sitsmiieesrarcre asbyeieteier eh isetersie WAGNERS DAVID) 1—=See WANDRYAJEANHBPRANCOIS | ciacic cee cis ctl es elelateislojete = ei Ee = e a mage 3 7 — ” m pl m ” : m Zz ” er = 7) = 7) LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS S3iyuvUugII A = ” = *« w z %7) P < = < = : < = z “ a ra S = = o 5 NYSE S We A § : ms SY [e) ae - Yor © a o EF Woy 2 E Nw 2, = = = N > = gas : A = > UTION NOILNLILSNI_ NVINOSHLINS S3I1YVYUSIT SMITHSONIAN INSTITUTION LIBRARIES fagdil LIBRARIES ow A > a m 1D: wn UTION NOILALILSNI ae = “ — \> = oe S 2 a) Z =z fddit LIBRARIES NOILNLILSNI UTION NOILNLILSNI INSTITUTION y¥gi7_ LIBRARIES Pa ‘ < . - ae 2 NSS = WS Ww “ JTION NOILNLILSNI BRARIES SMITHSONIAN TLALILSNI If BRARIES INSTITUTION NOILOLILSNI NVINOSHLINS S3IYVYUSITLIBRARIES Ss NOILNLILSNI NOILNLILSNI SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IY¥Vugit Li NVINOSHLINS S3I1¥YVYaIT LIBRARIES SMITHSONIAN INSTITUTION JNOIL w uJ uJ o o -< < ce oe ree] £0 3 a Ei Zz is ra > > - ~ : as oe “fl m 2 m D m z m 2 | NVINOSHLINS S3IYVYEIT LIBRARIES INSTITUTION NC o” = ag ” = Ne. 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