te-tatie 6 “« Pei rm hi One cope 2 : y Snsteiee Pag hye vibe etait Cheat bet BANOS CST f : , 2 Rute ade . é see fe Neath all Mitte hota ; Bs : i te ‘é WO "8 ee : ehake - : ee . eet neta piqued * F 4 Mpa i : ; F 3 Weise aever tees os Stitnces tp onecre shh : ‘ * ‘ 7 - > : E aA EEA NEA gh ating ( : Hs8 4 ; S : A ; z ¢ eta Vege igre dnts res : . ‘ 7 , 7 $7 B 2 ¢ 2 * ’ Arise even Pet AE, UR gs a af cei : ‘ Pevdpahanat eae doe ie INN EM Sane . . ‘ ™ ~ Peas Pe haan bhoen wis a RA nnn RL AD AR ME et eu ear 1 wire j VOL. 102 JANUARY 2000 NO. 1 @ ee (ISSN. 0013-8797) +N ELaX PROCEEDINGS | ea of the j N iad Ri PUBLISHED QUARTERLY CONTENTS ASSIS-PUJOL, CRISTIANE VIEIRA DE and MICHEL LECOQ—Comparative study of sper- mathecae in eleven Rhammatocerus Saussure 1861 grasshopper species (Orthoptera: Achadidae .Gomphnocerinacs Scyllimint) it yee, een eee na at eee ries cite ae ecm Keone ee aac ees 120 BROWN, JOHN W.—Revision of Lobogenesis Razowski and Odonthalitus Razowski (Lepidoptera: Tortricidae: Tortricinae), with comments on their monophyly ....................0eeeeee eee ee 2 BUTLER, LINDA and JOHN STRAZANAC—Macrolepidopteran larvae sampled by tree bands in temperate mesicund sernctorests injeastern, WiMlteGestates| siieeeiec one. sane sees sets neice 188 CICERO, MARY R. and EDWARD M. BARROWS—Milesiine flower flies (Diptera: Syrphidae) in a central Appalachian broadleaf forest: Abundances, flight periods, and diflubenzuron.... 50 CRYAN, JASON R. and LEWIS L. DEITZ—Review of the New World treehopper tribe Stegaspidini (Hemiptera: Membracidae: Stegaspidinae): Ill: Flexocentrus Goding, SAVLOCENIFUSHS tal sand, O71 DEULSCTUSsD CLUZ ancy a ene tele terns ocd t= ee aicielacs pe iste re Aaah tata 82 DARSIE, RICHARD E.,, JR.—Description of the pupae of five species in the subgenus Armigeres, genus Armigeres Theobald, with a key to species of the known pupae of the sub- PenUshUMiptekan GUC Ae amet ste satie en rh Mel vartele = ceuctiie Mie ara cistaevesieis NSE aol iam ae sl yt eueprite 108 GATES, MICHAEL W.—A new species of Cirrospilus Westwood (Hymenoptera: Eulophidae) fLOMMtAe SOULNWeSteEMNUMIted States anu MCxI GO ca), ac elt tee ae cra tele roo 'e ove aleve emis ievoe siassvayeinvere= 58 GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota signifera (Coquillett) (Diptera: Tephritidae) on Hemizonia pungens (Hooker and Arnott) Torrey and ASGray, (Asteraceae) in SOUthenne @alitOnntaet S10 yeie es «lor tente sie ah asia es viele else ona cle slgaiauaresasaler 69 HASTRITER, MICHAEL W.—Jordanopsylla becki (Siphonaptera: Ctenophthalmidae), a new SHECIES OM CanhOud MCU EVAMAMLESE SILC Mth ye my rac cto ercie ay hacteieei-lopiate slotelsiseris erm roe elem nate elerateret 215) HOEBEKE, E. RICHARD, ROBERT A. BYERS, MIGUEL A. ALONSO-ZARAZAGA, and JAMES F. STIMMEL—Jschnopterapion (Chlorapion) virens (Herbst) (Coleoptera: Curculionoidea: Brentidae: Apioninae), a Palearctic clover pest new to North America: Recognition featiressdisintoution:)and DIOMOMMICS Heke aic to clo sy claveciewc J+) eisioe ts eleharstais mie eerste 151 HUNG, AKEY C. F. and HACHIRO SHIMANUKI—Nucleotide sequence and restriction site analy- ses in three isolates of Kashmir bee virus from Apis mellifera L. (Hymenoptera: Apidae) ...... 178 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2000 Davip G. FurtH, President MIcHAEL G. PoGuE, 7reasurer JOHN W. Brown, President-Elect THEODORE R. SCHULTZ, Program Chair Stuart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Ho us B. WILLIAMS, Corresponding Secretary MiIcHAEL E. ScHAuFF, Past President Jon A. Lewis, Custodian Davip R. SmitH, Editor Publications Committee THOMAS J. HENRY Wayne N. MaAtTHIs Gary L. MILLER, Book Review Editor Honorary President Louise M. RuUssELL Honorary Members KARL V. KROMBEIN RONALD W. HoDGEs DoNALD M. ANDERSON WILLIAM E. BICKLEY All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.— Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:30 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $25.00 (U.S. currency). PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub- lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send address changes to the Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomo- logical Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE PP. 699-700 OF THE JULY 1999 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- West, Beltsville, MD 20705. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 10 January 2000 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. 102(1), 2000, pp. 1-20 A REVISION OF THE ENDEMIC HAWAIIAN REDUVIID GENUS SAICELLA USINGER, WITH DESCRIPTIONS OF FOUR NEW SPECIES (HETEROPTERA: REDUVIIDAE: EMESINAE) DAN A. POLHEMUS Department of Entomology, MRC 105 , Smithsonian Institution, Washington, DC 20560-0105, U.S.A. (e-mail: polhemus.dan @ nmnh.si.edu) Abstract.—The Hawaiian genus Saicella i s revised, and four new species are described: S. perkinsi from Oahu, S. lilinoe from Maui, S. kipahulu from Maui, and S. mulli from Hawaii. A key to species is provided, accompanied by distribution maps, habitus figures, and illustrations of key characters. Key Words: Reduviidae, Hawaiian Islands Assassin bugs, in the family Reduviidae, are uncommon and elusive components of the native Hawaiian insect biota. Only two endemic genera, Saicella Usinger and Nes- idiolestes Kirkaldy, both in the subfamily Emesinae, are known from the islands, con- taining a total of five currently described species between them. Even with the de- scription of the additional new taxa now present in museum collections, it seems un- likely that the total native Hawaiian redu- viid fauna will exceed 15 species. This stands in contrast to the native Hawaiian Nabidae, which appear to fill many typical reduviid niches in the islands. The latter group contains 25 described species, with an equal or greater number awaiting de- scription. This is a striking reversal of fam- ily dominance in comparison to continental settings, where species of Reduviidae far outnumber those of Nabidae, and is a pe- culiarity of community organization in the Hawaiian archipelago and other isolated Pa- cific island groups. It is probably indicative of a difference in overwater dispersal abil- ities between these two groups of Heter- optera (Leston 1957), and of their relative capabilities to exploit underutilized ecolog- ical niches. , Saicella, new species, biogeography Here I revise the classification of Saicel- la, providing descriptions of four new spe- cies. The two previous species in this ge- nus, S. smithi Usinger (1958) and S. usin- geri Wygodzinsky (1966), were described from Maui and Kauai respectively. Two of the new taxa described herein are also from Maui, with the other two coming from the islands of Oahu and Hawaii. These new species now extend the known range of the genus throughout the archipelago, and it seems quite likely that further collecting will produce examples of this genus from Molokai and possibly Lanai. Collections of Saicella have been sporad- ic at best over the last 100 years. The first two examples were taken by R. C. L. Per- kins on Maui in 1894 and 1896, during his collecting work for the Fauna Hawaiiensis. They seem to have escaped the notice of Kirkaldy, who authored the treatment of Heteroptera for this series, and were not formally described until 1958, when R. L. Usinger happened across one of the speci- mens in the British Museum, and subse- quently located the second in Honolulu, at the Bishop Museum. Usinger was eventu- ally able to collect a series of a second spe- i) cies on Kauai in 1961, which was described by Wygodzinsky (1966) in his monograph of world Emesinae. A few additional spec- imens were collected in the mid-1980’s by researchers from the Bishop Museum dur- ing biological surveys on Maui and Oahu, and a series of specimens was reared from eggs by hobbyist William Mull on Hawaii. Even so, the sum total of specimens col- lected between 1894 and 1994 amounted to less than 50 individuals, many of them im- matures. This situation has changed markedly in the last several years, due to a better un- derstanding of the microhabitats preferred by Saicella species and improvements in collecting techniques. In particular, the use of small scale pyrethrin fogs targeted at mossy logs and tree trunks in areas of wet native forest has been especially effective. This technique produced 80 specimens from five different localities on Maui and Kauai in only nine months during 1998 and 1999, thereby more than doubling the adult captures from the previous hundred years, and revealing two new species in the pro- cess. Continued use of such local fogging may well prove Saicella to be typical, though cryptic, components of wet, high el- evation native forest communities on all the major Hawaiian volcanoes. The comparatively high diversity of Sai- cella on Maui is an interesting biogeo- graphic anomaly, especially considering that all three Maui species have come from Haleakala, the mountain that forms the east- ern portion of the island, while none are yet known from the West Maui Mountains. The Haleakala species divide the mountain into three discrete areas of endemism, one in the area near upper Waikamoi Gulch, another on the central portion of the northern face in the upper Hanawi Gulch area, and a third in Kipahulu Valley (Fig. 12). These areas of Saicella endemism are roughly congru- ent with portions of the mountain retaining a surface of Kula Volcanic lavas with a mean age of 0.41 ma, which are surrounded by younger flows of the Hana Volcanics PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON that erupted in the late Pleistocene, 0.4 ma later (Clague and Dalrymple 1987, Langen- heim and Clague 1987). It is tempting to hypothesize that the patterns of speciation currently observed in Maui Saicella are the result of allopatry due to vicariance, with the eruption of the Hana Volcanics having isolated sections of the older Kula Volcanic surface, which then functioned as islands of rain forest habitat, but tests of this model must await a cladistic analysis. The Maui pattern is all the more interesting in light of the fact that no similar pattern of intra-is- land endemism is observed on the much older island of Kauai, where a single spe- cies, S. usingeri, occurs on both the Alakai Plateau and in the Makaleha Mountains, de- spite the isolation of these massifs from each other by sheer cliffs, deep valleys, and other topographic barriers easily as signifi- cant as those separating the three allopatric species on Maui. METHODS Synonymies given are nomenclatural only. All measurements are given in milli- meters. Characters typical of the genus that are noted in the generic description are not repeated in the individual species descrip- tions. To avoid confusion, tergite number- ing in the descriptions refers to visible ter- gites; the basal tergites of many Emesinae are reduced or hidden (see Wygodzinsky 1966), so that the location of actual tergite I is often difficult to ascertain. Institutional abbreviation codes used in the Material Examined sections are ex- plained in the Acknowledgments section. In addition, information not found on original specimen labels but helpful in their inter- pretation is given in brackets in the Material Examined sections. Spellings for place names follow those found on the topo- graphic maps of the United States Geolog- ical Survey, and do not include diacritical marks. In the following discussions, the word ‘‘Hawaii”’ refers to the island of Ha- waii itself. If the entire Hawaiian island VOLUME 102, NUMBER 1 chain is under discussion then the phrase ‘*Hawaiian Archipelago” is employed. Genus Saicella Usinger 1958 (Figs. 1-13) Type species.—Saicella smithi Usinger 1958: 440. Description.—Small, micropterous redu- viids, length 4.5—6.0 mm (Figs. 1—2). Body surface shining, portions of head, thorax, and basal abdominal segment often bearing short, appressed, wool-like pile. Ground color varying from pale green to yellowish brown, marked with red, dark brown, or black, legs often conspicuously annulate with dark brown or black. Head short, divided into anteocular and postocular portions by deep transverse sul- cus, both portions strongly convex above, anteocular section with sides subparallel when viewed dorsally, postocular section semi-globular both dorsally and laterally; eyes small, set laterally, lying below dorsal surface of head; rostrum 3 segmented, bent at first intersegmental suture, cylindrical, length of segment I subequal to lengths of segments II and III combined, segment I bearing 2+2 spinelike setae to either side of midline, segment II with 1+1 similar se- tae (Fig. 3); antenniferous tubercles large, antennae inserted near anterior margin of head, lacking long hairs in both sexes; buc- cula with a pair of socketed spine-like setae, gena with 3—4 similar setae. Pronotum covering anterior half of me- sonotum, divided into anterior and posterior sections by deep, forward curving trans- verse sulcus; anterior section subglobular, with a deep longitudinal median sulcus sep- arating two elevated lobes; posterior section transverse; anterior acetabulae opening for- ward and downward. Scutellum (exposed portion of mesoscutum) subequal in length to posterior section of pronotum, bearing a long, erect spine. Metanotum subequal in length to scutellum, also with a long, erect spine. Forewings consisting of short, slen- der pads, reaching at most to base of ab- domen, membrane absent. Abdomen broadly attached to thorax, not constricted at base, connexival margins out- wardly convex, broadly arcuate, occasion- ally sculptured, dorsal tergites often bearing small tumescences posteromedially, ventri- tes broadly convex. Male with seventh vis- ible tergite triangular, elongate, often with a longitudinal medial keel, completely cov- ering genital segments when viewed dor- sally; ventrite VIII fully visible, large; py- gophore large, length equal to % total ab- domen, sclerotized dorsally, posterosuperi- or process narrowing suddenly on distal half, coming to a spine-like point; paramer- es small, club-like. Foreleg stout, coxa with 2 long, stout, socketed spines basally, followed by 2-3 smaller spines distally; trochanter unarmed; femur with 5 long, stout, socketed spines on basal % of ventral surface (Fig. 3), in- termixed with long erect pale setae and set within a row of smaller spinules, length of the longest spines subequal to width of fe- mur, inner dorsal surface of femur bearing about 10 moderately long socketed spines; tibia slender, ventral surface with two lon- gitudinal combs of erect, pale, spine-like setae, small brushes of gold setae also pre- sent at tip on inner face, and on central sec- tion of outer face; tarsus two segmented, basal segment slightly shorter than second, both segments with numerous short setae on ventral surfaces, lacking specialized se- tae; claws curved, inner claw medially in- cised, outer claw with two small subbasal projections. Middle and hind legs slender and elongate, hind femur surpassing apex of abdomen, all segments bearing numerous very short, semi-erect, distally angling se- tae; middle and hind tarsi 3 segmented, these segments subequal in length. Discussion.—Usinger (1958) considered the subfamilial placement of Saicella to be equivocal, due to the uniformly micropter- ous condition, and suggested that the genus might fall in either the Saicinae or Emesi- nae. Wygodzinksy (1966) provided detailed arguments for a placement in the Emesinae, 4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tribe Ploiariolini, an interpretation followed herein. KEY TO THE SPECIES OF SAICELLA 1. Wing pads long; reaching to base of abdomen, surpassing posterior tip of scutellar spine when viewed from above (Fig. 2); Maui WE RHA ELD coeds ane de aT © smithi Usinger — Wing pads shorter, not reaching to base of ab- domen, usually not surpassing posterior tip of scutellar spine (except in S. perkinsi, where this spine is vertical), definitely not exceeding base ofmetanotallispinei(Gicai)i yates 2 2. Antennal segment I distinctly annulate, with al- ternating dark and pale bands (Fig. 1) ..... 3 — Antennal segment I usually unicolorous brown, bearing at most a few faint indications of dark- ErpannuUlatlONSie. wiry js oe eee a one ae 5 3. First visible abdominal tergite with conical tu- mescence anteromedially; ground color pale green overlain with darker markings; Maui Gf tO ler aus Daca DI OPER Ley CECk oan 0, dyons fo S. lilinoe, n. sp. — First visible abdominal tergite lacking a conical tumescence; ground color yellowish brown, overlain with darker brown or reddish mark- Ine sa Walalpandehlawalluewe emery) cars es ete 4 4. Wing pads extending to tip of scutellar spine: anterior lobes of pronotum dull, bearing patch- es of pale appressed hairs; abdomen with red markangsilaterallliyiKaualeyepeseis eats F keagnied «yaad ht Mos shen S. usingeri Wygodzinsky Wing pads tiny, hard to see, not extending past base of scutellar spine; anterior lobes of pron- otum glabrous, shining; abdomen without red markings laterally; Hawali...... S. mulli, n. sp. 5. Pronotum with numerous long, slender, erect, pilose hairs; hind tibia unicolorous medium brown, without well defined annulations; an- terior margin of first visible abdominal tergite strongly angled upward anteromedially; Oahu ed SE GES ict ch CaS res eel OUR Eee fh S. perkinsi, n. sp. — Pronotum lacking long, erect, pilose hairs, bearing only patches of pale, appressed hairs; hind tibia with numerous black annulations; anterior margin of first visible abdominal ter- gite not strongly angled upward anteromedial- liv; Mi alse. bpd Sokpalts ay clceresey = S. kipahulu, n. sp. Saicella smithi Usinger (Figs. 2—3, 9, 12) Saicella smithi Usinger 1958: 440. Type.—Holotype, 2, from Haleakala, Maui, 5,000 ft., in The Natural History Mu- seum, London. Diagnosis.—Recognized by the relative- ly long wing pads that reach beyond the apex of the scutellar spine to the base of the abdomen (Fig. 2); absence of obvious an- nulations on antennal segments I and II; and absence of raised tubercles on the abdomi- nal tergites (Figs. 2-3, 9). Redescription.—Micropterous female: General coloration yellowish brown, with darker brown markings on thorax and base of abdomen; legs multiannulate with dark brown; lateral portions and posterior margin of abdominal tergites sparingly marked with bright red. Length 4.70 mm, maximum width (across abdomen) 1.50 mm. Head \ength/width = 0.68/0.54, covered by a thick layer of short, pale, curling, re- cumbent setae; width of vertex 2.9 dorsal width of an eye (0.32/0.11); length of an- terior lobe of head 2.50 dorsal length of an eye (0.40/0.16); eyes small, consisting of approximately 20 ommatidia each; length of posterior lobe of head 1.68 dorsal length of an eye (0.27/0.16); ocelli absent; length of antennal segments I-IV = 1.98/ 2.07/0.52/missing; rostrum length 0.85, reaching to bases of fore coxae; coloration of head yellowish brown, without contrast- ing dark markings; antennal segment I dark yellowish, distal % medium brown, basal half with very faint suggestions of 1 or 2 brown annulations, antennal segments II and III uniformly light brown. Pronotum length (midline)/width = 0.57/ 0.63, covered by an obscure layer of short, pale, recumbent setae; coloration uniform golden yellowish brown; anterior section with lateral lobes shining, lacking patches of appressed setae; posterior section bearing a small, conical tumescence centrally. Scutellum triangular, coloration uniform- ly golden yellowish brown, lateral angles bearing an obscure layer of short, pale, re- cumbent setae, central section produced into an erect, slender, backward angling spine posteromedially; length/width = 0.27/ Oals: Hemelytra short, micropterous, reaching to base of abdomen, separated medially by an erect, slender, backward angling spine VOLUME 102, NUMBER 1 fe 1 Fig. 1. Saicella lilinoe. Female, dorsal habitus (Young Sohn illustration). 6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-3. Saicella smithi, female (Arthur Smith illustrations). 2, Dorsal habitus. 3, Body and forelegs in lateral view. arising from the underlying mesonotum, gested, covered by an obscure layer of this spine similar in size, shape and orien- short, pale, recumbent setae, posterior mar- tation to that arising from scutellum; wing gins rounded, membrane absent; coloration pads rugulose, with claval vein barely sug- yellowish brown. VOLUME 102, NUMBER 1 Legs elongate, with fore coxa approxi- mately 3.5X as long as thick (0.63/0.18); fore trochanter bearing numerous slender erect gold setae; fore femur fusiform, over 5.7X longer than wide (1.26/0.22); colora- tion of legs yellowish brown with brown markings; fore femur yellowish brown ba- sally, with a broad brown annulation on dis- tal half; fore tibia with 3 brown annulations, knee pale, apex dark; middle and hind fem- ora each with 4 brown annulations, these annulations roughly equal in length to in- tervening pale spaces; middle and hind tib- iae with 5 and 7 brown annulations respec- tively, these annuli concentrated in the bas- al half of the segment; middle and hind tarsi uniform brown; lengths of leg segments as follows: fore femur/tibia/tarsal 1/tarsal 2 = 1.26/0.99/0.06/0.20; middle femur/tibia/tar- sal 1/tarsal 2/tarsal 3 = 2.14/2.87/0.06/0.06/ 0.06; hind femur/tibia/tarsal 1/tarsal 2/tarsal 3 = 2.86/4.16/0.05/0.05/0.05. Abdomen yellowish brown, with lateral and posterior sections of first visible tergite darker brown, lateral sections of first through sixth visible tergites also bearing irregular bright red maculations; connexival margins broadly arcuate, abdominal tergites lacking raised tumescences posteromedial- ly, first through sixth visible segments dor- sally concave, with lateral margins curved upward and narrowly emarginate, posterior margin of sixth visible tergite bisinuate, seventh visible tergite broadly triangular, flat, with a very small indentation at ex- treme posterior apex (Fig. 9); all tergites shining, covered by a sparse, obscure layer of very short, pale, recumbent setae. Ventral surface yellowish brown, cov- ered by an obscure layer of short, pale, re- cumbent setae; abdominal paratergites red- dish. Micropterous male: Unknown. Distribution.—Maui (Haleakala, Waika- moi area) (Fig. 12). Ecological associations.—Unknown. Material examined.—HAWAIIAN IS- LANDS, Maui: 1 92, Haleakala, 5,000 ft 7 P30: mij. EEAEIV. 18943 RoC: I. Perkins (paratype, BPBM). Discussion.—The description above was taken from a female paratype, housed in the Bishop Museum. It is one of the two orig- inal specimens of S. smithi on which Usin- ger (1958) based his description. The Ar- thur Smith figure accompanying Usinger’s (1958) description, reproduced in Fig. 2, is accurate in most respects, but depicts ob- vious dark annulations on antennal segment I that are not present on the Bishop Muse- um specimen examined by the author. Both of the above specimens were taken by Perkins from “Haleakala, 5,000 ft.”’; based on Perkins’ field notes, the type lo- cality must lie somewhere upslope of Olin- da and west of Waikamoi Gulch. Much of the forest in this area has now been cleared for cattle ranching, but a tract is still pro- tected in The Nature Conservancy of Ha- wail’s Waikamoi Preserve. Repeated sur- veys in this remaining forest block, how- ever, have failed to produce further speci- mens of S. smithi. As a result, the male characters of this species remain unknown. Saicella lilinoe Polhemus, new species (Figs. 1, 8, 12) Diagnosis.—Recognized by the short wing pads that do not reach beyond the apex of the scutellar spine (Fig. 1); strongly annulate antennal segments I and II (Fig. 1); and presence of raised tubercles poster- omedially on the fourth through sixth visi- ble abdominal segments (Figs. 1, 8). Description.—Micropterous male: Gen- eral coloration pale green, with darker brown markings on head, thorax and ab- domen; legs multiannulate with dark brown or black (Fig. 1). Length 5.25 mm, maximum width (across abdomen) 1.80 mm. Head \ength/width = 0.80/0.56, covered by a thick layer of pale, appressed setae; width of vertex 4.8 dorsal width of an eye (0.44/0.09); length of anterior lobe of head 2.4 dorsal length of an eye (0.45/0.19); eyes small, consisting of approximately 20 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 4-9. 7, S. kipahulu. 8, S. lilinoe. 9, S. smithi. ommatidia each; length of posterior lobe of head 1.63 dorsal length of an eye (0.31/ 0.19); ocelli absent; length of antennal seg- ments I-[V = 3.10/2.75/0.75/0.55; rostrum length 1.00, reaching to bases of fore coxae; coloration of head uniform medium brown, Saicella species, dorsal view of female terminal abdomen. 4, S. mulli. 5, S. usingeri. 6, S. perkinsi. without contrasting markings; antennal seg- ment I dark yellow, bearing 5 evenly spaced dark brown annulations, tip light; antennal segment II dark yellow, bearing 6 evenly spaced dark annulations, tip dark; antennal segments III and IV uniform medium VOLUME 102, NUMBER 1 brown except narrowly pale at extreme ba- ses and tips. Pronotum length (midline)/width = 0.75/ 0.50; anterior section yellowish green, be- coming embrowned toward pleurae, lateral lobes shining, lacking setae, intervening longitudinal sulcus bearing numerous ap- pressed pale setae; posterior section of pronotum milky greenish white, lacking se- tae, bearing a small, conical tumescence centrally. Scutellum triangular, lateral angles black, central section pale green, produced into an erect, slender, backward angling spine pos- teromedially, this spine strongly curving when viewed laterally; a ring of pale, ap- pressed setae present around base of spine; length (including spine)/width = 0.32/0.27. Hemelytra short, micropterous, reaching only to middle of scutellar spine when viewed from above, separated medially by an erect, slender, pale green, backward an- gling spine arising from the underlying me- sonotum, this spine similar in size, shape and orientation to that arising from scutel- lum; wing pads consisting of tiny, elongate flaps, venation and membrane absent; col- oration black centrally, margins brown. Legs elongate, with fore coxa approxi- mately 4.0 as long as thick (0.80/0.20); fore trochanter bearing numerous slender erect pale setae; fore femur fusiform, 7.0 longer than wide (1.75/0.25); coloration of legs dark yellow with brown or black mark- ings; fore femur yellowish brown basally, with a broad brown annulation on distal half; fore tibia with 3 diffuse brown annu- lations on outer face, knee pale, apex nar- rowly pale; middle and hind femora each with 4 brown annulations, these annulations broad and diffuse basally, becoming smaller than width of intervening pale areas distal- ly; middle and hind tibiae each with 4—6 brown annulations, these annuli regularly spaced on middle tibia, more irregularly spaced on hind tibia, with a prominent black annulation on basal %4, followed by other smaller annulations in distal half of segment; middle and hind tarsi uniform me- dium brown; lengths of leg segments as fol- lows: fore femur/tibia/tarsal 1/tarsal 2 = 1.75/1.37/0.09/0.25; middle femur/tibia/tar- sal 1/tarsal 2/tarsal 3 = 3.50/4.70/0.09/0.09/ 0.09; hind femur/tibia/tarsal 1/tarsal 2/tarsal 3 = 4.80/6.60/0.10/0.10/0.10. Abdomen pale green, yellowish centrally, this ground color overlain with irregular black markings on entirity of first visible tergite and posterolaterally on second through sixth visible tergites, the lateral dark markings on third through sixth visible tergites extending inward along posterior tergite margins to form transverse bands, anterior sections of third through sixth vis- ible tergites with blackish brown marks in the form of posteriorly directed triangles, this combination of markings giving the ab- domen a transversely striped appearance (Fig. 1); connexival margins forming incip- ient knobs at sutures, intervening sections weakly concave; first visible tergite with a small, conical tumescence anteromedially, third through fifth visible tergites with smaller, more rounded raised tumescences posteromedially, sixth visible tergite trian- gular, longitudinally keeled on _ posterior half; all tergites shining, covered by a sparse, obscure layer of very short, pale, re- cumbent setae. Ventral surface of head and thorax black, abdominal venter greenish brown, covered by an obscure layer of short, pale, recum- bent setae; abdominal paratergites narrowly margined with pale green adjoining con- nexival margin. Male genitalia with paramere stout, slightly curving, bearing a small lateral tab at tip; tip of pygophore narrowed and elon- gate, tapering evenly to truncate apex with a slightly expanded terminal cap. Micropterous female: Length 5.70 mm, maximum width (across abdomen) 1.90 mm. Similar to male in general structure and coloration, but with abdomen broader and more strongly expanded; posteromedial tumescences on abdominal tergites more highly developed, particularly on fifth vis- ible tergite, posterior margin of sixth visible 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tergite broadly V-shaped, seventh visible tergite roughly triangular, rounded posteri- orly (Fig. 8). Distribution.—Maui (Haleakala) (Fig. 12): Ecological associations.—All specimens so far captured have been taken by applying a light pyrethrin fog to mossy tree trunks and root masses of ohia trees (Metrosideros polymorpha). These insects appear to prefer the sheltered, almost cave-like situations found in cavities beneath multi-stemmed Metrosideros and fallen logs, and multiple specimens are often taken at a single spot, indicating a tendency toward aggregation. Material examined.—Holotype, microp- terous 6: HAWAIIAN ISLANDS, Maui, Haleakala, Poo Uli Cabin area, near head- waters of Kuhiwa Stream, Hanawi Natural Areay Reserve 5,200), ft VSs85) me]; 20°45.03'N, 156°07.40'W, 5-6 May 1998, CL 8324, D. A. Polhemus (BPBM). Para- types: HAWAIIAN ISLANDS, Maui, Ha- leakala: 7 3d, one 2, same data as holotype (USNM, BPBM); 14 6,9 @, same locality as preceeding, 5 May 1998, lot Ol, pyre- thrin fog on mossy ohia [Metrosideros po- lymorpha], J. K. Liebherr (CUIC); 11 36,5 ?, 5 immatures, State Fence Camp cabin, along headwaters of Heleleikeoha Stream, Hana Forest Reserve, 5,300 ft. [1,615 m.], 20°44.41'N, 156°06.12'’W, 11-12 May 1998, CL 8326, D. A. Polhemus, J. K. Lie- bherr and C. Ewing (USNM, BPBM); 5 6, 4 2, same locality as preceeding, 12 May 1998, lot 01, pyrethrin fog on mossy ohia [Metrosideros polymorpha], J. K. Liebherr (CUIC); 1 36, Haleakala National Park, Ki- pahulu Valley, West Rim below Kuiki, 2,090 m. [6,855 ft.], 14 May 1993, lot O1, sifting moss from ohia [Metrosideros po- lymorpha], J. K. Liebherr and A. C. Me- dieros (CUIC). Etymology.—The name “‘lilinoe’’ refers to the Hawaiian goddess of mists, an ap- propriate name for this species of the cloud forests. Discussion.—In addition to the charac- ters mentioned in the key and diagnosis, Saicella lilinoe may be easily recognized by its green and black coloration in living in- dividuals, the striped appearance of the ab- domen (Fig. 1), and the glabrous anterior pronotal callosities. This species appears to be distributed across the northern face of Haleakala, from the Koolau Gap eastward to Kipahulu Val- ley. This is an extremely wet cloud forest zone, with precipitation exceeding 9,000 mm (360 inches) a year. A single specimen is also at hand from the upper section of Kipahulu Valley, further around the eastern tip of the island near to the type locality of S. kipahulu; this is the only place where two species of Saicella are known to be poten- tially sympatric. Saicella kipahulu Polhemus, new species (igs! 7; 12) Diagnosis.—Recognized by the short wing pads that do not reach beyond the apex of the scutellar spine; unicolorous antennal segments I and II, and absence of raised tu- bercles posteromedially on the fourth through sixth visible abdominal segments. Description.—Micropterous male: Gen- eral coloration medium brown to yellowish brown, with dark brown or reddish mark- ings on abdomen; legs multiannulate with dark brown. Length 5.20 mm, maximum width (across abdomen) 1.25 mm. Head \ength/width = 0.79/0.55, covered by a thick layer of pale, appressed setae; width of vertex 3.7 dorsal width of an eye (0.37/0.10); length of anterior lobe of head 4.2 dorsal length of an eye (0.50/0.12); eyes small, consisting of approximately 20 ommatidia each; length of posterior lobe of head 2.42 dorsal length of an eye (0.29/ 0.12); ocelli absent; length of antennal seg- ments I-IV = 2.25/2.00/0.75/0.40; rostrum length 0.95; coloration of head uniform me- dium brown, without contrasting markings; antennal segment I medium brown, becom- ing slightly paler basally, lacking annula- tions; antennal segments II-IV uniform me- VOLUME 102, NUMBER 1 = o D £ 1) =] a) @) S. perkinsi OC) eri on Kauai. 11, S. perkinsi on Oahu. Distribution of Saicella species. 10, S. using Figs. 10-11. 12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dium to dark brown except narrowly pale at extreme bases and tips. Pronotum length (midline)/width = 0.55/ 0.75; anterior section medium brown, lat- eral lobes bearing scattered, elongate patch- es of appressed gold setae, intervening lon- gitudinal sulcus also bearing appressed gold setae; posterior section of pronotum yellow- ish brown, with a raised longitudinal me- dian carina, set with scattered short, semi- erect Setae. Scutellum triangular, medium brown, central section pale yellowish brown, pro- duced into an erect, slender, backward an- gling spine posteromedially, this spine strongly curving when viewed laterally; a ring of appressed gold setae present around base of spine; length (including spine)/ width = 0.35/0.30. Hemelytra short, micropterous, reaching nearly to tip of scutellar spine when viewed from above, separated medially by an erect, slender, yellowish brown spine arising from the underlying mesonotum, this spine sim- ilar in size and shape to that arising from scutellum, but more vertical; wing pads consisting of narrow, elongate flaps, bear- ing a few short, erect gold setae, venation and membrane absent; coloration medium brown. Legs elongate, with fore coxa approxi- mately 4.0 as long as thick (0.75/0.20); fore trochanter bearing numerous slender erect pale setae; fore femur fusiform, 6.0 longer than wide (1.50/0.25); coloration of legs dark yellow with brown or black markings; fore femur yellowish brown ba- sally, with a broad brown annulation on distal half; fore tibia with 3 diffuse brown annulations, knee pale, apex dark; middle and hind femora each with 4 brown an- nulations, these annulations broad and dif- fuse basally, becoming smaller than width of intervening pale areas distally; middle and hind tibiae each with 6 brown annu- lations, these annuli regularly spaced on middle tibia, more irregularly spaced on hind tibia, with a prominent black annu- lation on basal %4, followed by other small- er annulations in distal half of segment; middle and hind tarsi uniform medium brown; lengths of leg segments as follows: fore femur/tibia/tarsal 1/tarsal 2 = 1.50/ 1.20/0.10/0.25; middle femur/tibia/tarsal 1/ tarsal 2/tarsal 3 = 2.60/3.70/0.08/0.08/ 0.08; hind femur/tibia/tarsal 1/tarsal 2/tar- sal 3 = 3.60/5.10/0.08/0.08/0.08. Abdomen dark yellow, this ground color overlain with irregular black markings on basal % of first visible tergite and antero- medially on second through sixth visible tergites, the dark markings on these latter tergites extending outward along anterior tergite margins to form transverse bands, giving the abdomen a transversely striped appearance, additional irregular dark mark- ings present on connexival segments, su- tures between tergites and connexiva irreg- ularly blotched with red; connexival mar- gins evenly convex and curving, lacking knobs at sutures; first visible tergite with anterior margin raised to form a low, tri- angular tumescence anteromedially, second through fifth visible tergites lacking tumes- cences, sixth visible tergite elongate, trian- gular, tip swollen and rounded, pale; all ter- gites shining, covered by a sparse, obscure layer of very short, pale, recumbent setae. Ventral surface of head and thorax black, abdominal venter greenish brown, covered by an obscure layer of short, pale, recum- bent setae; abdominal paratergites narrowly margined with pale green adjoining con- nexival margin. Male genitalia with paramere stout, bear- ing a small lateral tab at tip; tip of pygo- phore narrowed and elongate, tapering evenly to acuminate apex. Micropterous female: Length 5.75 mm, maximum width (across abdomen) 1.80 mm. Similar to male in general structure and coloration; abdomen broader and more strongly expanded, posterior margin of sixth visible tergite nearly straight, seventh visible tergite triangular (Fig. 7). Distribution.—Maui (Haleakala) (Fig. 12): Ecological associations.—The habitat VOLUME 102, NUMBER 1 © = S. smithi @) =S. lilinoe [m| = S. kipahulu Fig. 12. Distribution of Saicella species on Maui. preferences of this species are very similar to those of S. Jilinoe, with individuals being taken from mossy tree trunks and root mas- ses in wet montane forests. Material examined.—Holotype, microp- terous 6: HAWAIIAN ISLANDS, Maui, Haleakala, West Camp, upper Kipahulu Valley, Haleakala National Park, 6,400 ft. [1,950 m.], 20°43.36’N, 156°07.79'W, 18- 22 May 1998; C833 1D ArrPolhemus, B. H. Gagné and R. Takumi (BPBM). Par- atypes (all micropterous): HAWAIIAN IS- LANDS, Maui, Haleakala: 1 6, 8 2, same data as holotype (USNM, BPBM, CUIC); 1 3,1 &, rim of pit crater downslope from West Camp, upper Kipahulu Valley, Hale- akala National Park, 6,050 ft. [1,845 m.], 20°43.18’N, 156°07.73'’W, 19 May 1998, 13:00—17:00 hrs., CL 8333, D. A. Polhe- mus, B. H. Gagné and R. Takumi (USNM); 1 3,3 2, Kipahulu Valley, West Camp, Ha- leakala National Park, 1,850 m. [6,070 ft.], 28 February 1984, at night, EK G. Howarth (BPBM); | 2, same data as preceding but 12 July 1983, on tree trunk, E G. Howarth (BPBM); 3 2, same data as preceding but 17 July 1983, under wet ohia (Metrosideros polymorpha) bark, EF G. Howarth (BPBM). Etymology.—The name “kipahulu”’ is a noun in apposition and refers to the Kipa- hulu Valley type locality. Discussion.—In addition to the charac- ters given in the key and diagnosis, Saicella kipahulu may be recognized by its brown and black coloration in living individuals, and by the scattered patches of appressed golden hairs on the anterior pronotal cal- losities. On the basis of current collections, this species appears to be confined to Kipahulu Valley, a deep, cliff-bound gap cutting the eastern face of Haleakala. All collections have come from the upper section of the 14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON valley, in the highest forest zone near the transition to the subalpine grasslands. Saicella usingeri Wygodzinsky (Figs. 5, 10) Saicella usingeri Wygodzinsky 1966: 408. Types.—Holotype, d¢ and allotype & from Kawaikoi Ridge, Kokee, Kauai, in Bishop Museum, Honolulu. Diagnosis.—Recognized by the annulate first antennal segment; relatively long wing pads that reach to the apex of the scutellar spine when viewed from above; yellowish brown coloration with red markings on the abdomen; and short length of the seventh visible female abdominal tergite, which is rounded posteriorly rather than broadly an- gulate. Redescription.—Micropterous male: General coloration golden yellowish brown with slight greenish overtones, darker brown or red markings present on head, thorax and abdomen; legs multiannulate with dark brown. Length 6.10 mm, maximum width (across abdomen) 1.45 mm. Head \ength/width = 0.75/0.60, covered by a thick layer of wooly, appressed gold setae; width of vertex 3.1 dorsal width of an eye (0.37/0.12); length of anterior lobe of head 2.2 dorsal length of an eye (0.37/ 0.17); eyes small, consisting of approxi- mately 20 ommatidia each; length of pos- terior lobe of head 2.06 dorsal length of an eye (0.35/0.17); ocelli absent; length of antennal segments I-IV = 4.10/3.50/0.80/ 0.60; rostrum length 1.00, reaching to bases of fore coxae; anterior lobe of head dark yellowish brown, posterior lobe of head dark brown; antennal segment I dark yel- low, bearing 6 evenly spaced dark brown annulations of varying size, tip light; anten- nal segment II dark yellow, bearing 6 even- ly spaced dark annulations, tip dark except at extreme apex; antennal segments III and IV uniform medium brown except narrowly pale at extreme bases and tips. Pronotum length (midline)/width = 0.78/ 0.87; anterior section golden brown cen- trally, becoming embrowned toward pleu- rae, lateral lobes shining, bearing scattered patches of wooly, appressed gold setae, in- tervening longitudinal sulcus bearing nu- merous appressed gold setae; posterior sec- tion of pronotum yellowish brown, lacking evident setae, bearing a low longitudinal ca- rina medially rising to a small, conical tu- mescence posteromedially. Scutellum triangular, golden yellowish brown, central section paler yellowish, pro- duced into an erect, slender, backward an- gling spine posteromedially, this spine an- gulate near middle when viewed laterally; a ring of pale, appressed gold setae present around base of spine; length (including spine)/width = 0.45/0.27. Hemelytra short, micropterous, reaching to tip of scutellar spine when viewed from above, separated medially by an erect, slen- der, pale yellowish, backward angling spine arising from the underlying mesonotum, this spine similar in size, shape and orien- tation to that arising from scutellum; wing pads consisting of tiny, elongate pads, wid- ening somewhat posteriorly, venation and membrane absent; coloration medium brown, margins slightly lighter. Legs elongate, with fore coxa approxi- mately 4.5 as long as thick (0.90/0.20); fore trochanter bearing numerous slender erect pale setae; fore femur fusiform, 7.5 X longer than wide (1.87/0.25); coloration of legs dark yellow with brown or black mark- ings; fore femur yellowish on basal *%, with a broad brown annulation on distal % of outer face; fore tibia with 3 diffuse brown annulations on outer face, knee pale, apex dark; middle and hind femora each with 4 brown annulations, basal annulation broad and diffuse, all annulations shorter than in- tervening pale areas; middle and hind tibiae each with 10—11 brown to black annula- tions on outer faces, these annuli of des- creasing size distally but regularly spaced on middle tibia, more irregularly spaced and concentrated distally on hind tibia; middle and hind tarsi uniform medium VOLUME 102, NUMBER 1 HAWAII Fig. 13. Distribution of Siacella mulli on Hawaii. brown; lengths of leg segments as follows: fore femur/tibia/tarsal 1/tarsal 2 = 1.87/ 1.35/0.12/0.20; middle femur/tibia/tarsal 1/ tarsal 2/tarsal 3 = 3.75/5.40/0.07/0.07/0.07; hind femur/tibia/tarsal 1/tarsal 2/tarsal 3 = 5.00/8.20/0.10/0.10/0.10. in Abdomen golden yellowish brown with greenish overtones, this ground color over- lain with irregular dark brown to black markings anteromedially on all tergites, markings on first, second, third, and sixth visible tergites more extensive, these mark- 16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ings extending outward along anterior ter- gite margins to form transverse bands, giv- ing the abdomen a transversely striped ap- pearance; paratergites irregularly blotched with red on central sections, with dark markings on posterolateral angles; connex- ival margins broadly arcuate, lacking knobs at sutures; first visible tergite broadly raised anteromedially but not bearing a tumes- cence, all other tergites flat, lacking raised tumescences, sixth visible tergite elongate and triangular, longitudinally keeled on posterior half; all tergites shining, covered by a sparse, obscure layer of very short, pale, recumbent setae. Ventral surface yellowish brown mottled with dark brown, covered by an obscure layer of short, pale, recumbent setae; ab- dominal paratergites with spiracles creamy white. Male genitalia with paramere club-like, bearing a small lateral tooth near tip; apex of pygophore strongly and suddenly nar- rowed, tip acuminate (see Wygodzinsky 1966: Figs. 124b, c). Micropterous female: Length 5.50 mm, maximum width (across abdomen) 1.75 mm. Similar to male in general structure and coloration, but with abdomen broader and more strongly expanded; posterior mar- gin of sixth visible tergite VI bearing a V- shaped indentation medially, seventh visi- ble tergite short, broadly rounded (Fig. 5). Distribution.—Kauai (Alakai Plateau) (Fig. 10). Ecological associations.—A series of this species was taken near upper Koaie Stream by applying a light pyrethrin fog to mossy, fallen ohia (Metrosideros polymorpha) trunks, in the same manner used to obtain the Maui specimens discussed previously. Wygodzinsky (1966) states that Usinger ob- tained his type series by beating tree ferns (Cibotium sp.). Label data from another specimen collected by Swezey indicates it came from Tetraplasandra, an arborescent native forest plant with broad leaves. Material examined.—HAWAITIAN IS- LANDS, Kauai: 3 46, rainforest along Koaie Stream at USGS gauging station, near crossing of Mohihi—Waialae Trail, 1;130° om: (3, 700] sili 22-06 47 ANE 159°35'18"W, 7 January 1999, D. A. Pol- hemus (USNM); 1 6, 1 2, Alakai Swamp Trail, Kawaikoi Ridge, Kokee, 18 August 1961, tree ferns, R. L. Usinger (holotype and allotype, BPBM); 2 3, Kualapa, 2,000 ft. [610 m.], 23 October 1979, S. L. Mont- gomery (BPBM); | 2, Kokee, 4-6 August 1961, Maa, Miyatake and Yoshimoto (BPBM); 1 3, 2 immatures, Kokee, 20 Au- gust 1925, on Tetraplasandra, O. H. Swez- ey (BPBM). Discussion.—Saicella usingeri is endem- ic to Kauai; specimens are at hand from both the Alakai Plateau and the Makaleha Mountains, indicating that this species may occur on all the high mountain massifs of the island, including Namolokama and Mt. Kahili. Saicella mulli Polhemus, new species (Figs. 4, 13) Diagnosis.—This species may be recog- nized by the near-absence of wing pads, and by the unusual condition of the seventh vis- ible female abdominal tergite, which is di- vided into two lobes by a longitudinal me- dian sulcus (Fig. 4). Description.—Micropterous male: Gen- eral coloration yellowish brown, with dark- er brown markings on head, thorax and base of abdomen; legs multiannulate with dark brown. Length 5.50 mm, maximum width (across abdomen) 1.15 mm. Head \ength/width = 0.70/0.55, covered by a thick layer of short, pale, recumbent setae; width of vertex 3.5 dorsal width of an eye (0.32/0.09); length of anterior lobe of head 2.36 dorsal length of an eye (0.37/0.11); eyes small, consisting of ap- proximately 20 ommatidia each; length of posterior lobe of head 2.63 dorsal length of an eye (0.29/0.11); ocelli absent; length of antennal segments I-IV = 3.10/2.75/ 0.85/0.60; rostrum length 0.98, reaching to bases of fore coxae; coloration of head yel- VOLUME 102, NUMBER 1 lowish brown, darker along longitudinal midline on frons and vertex; antennal seg- ment I dark yellowish, distal ”; medium brown, basal 7% with 5 or 6 small brown annulations, antennal segments II and IV uniformly light brown. Pronotum length (midline)/width = 0.72/ 0.69; anterior section medium brown cen- trally, lateral lobes shining, bearing scat- tered patches of wooly, appressed gold se- tae only anterobasally, intervening longitu- dinal sulcus bearing numerous appressed gold setae; posterior section of pronotum golden yellowish brown, lacking evident setae, broadly domed centrally but not pro- duced to a sharp tumescence. Scutellum triangular, medium brown, central section pale yellowish, produced into an erect, slender, backward angling spine posteromedially, this spine angulate near middle when viewed laterally; a ring of pale, appressed gold setae present around base of spine; length (including spine)/ width = 0.26/0.27. Hemelytra very short, micropterous, dif- ficult to discern, not reaching to base of scutellar spine, separated medially by an erect, slender, backward angling spine aris- ing from the underlying mesonotum, this spine similar in size, shape and orientation to that arising from scutellum but not as sharply angulate near middle when viewed laterally. Legs elongate, with fore coxa approxi- mately 3.85 as long as thick (0.77/0.20); fore trochanter bearing numerous slender erect gold setae; fore femur fusiform, over 7.4X longer than wide (1.62/0.22); colora- tion of legs golden yellowish brown with dark brown markings; fore femur yellowish brown basally, with a broad, diffuse brown annulation on distal 4%; fore tibia with 3 dif- fuse brown annulations, knee pale, apex dark; middle and hind femora each with 4 brown annulations, most basal of these an- nulations diffuse, all annulations roughly equal in length to intervening pale spaces; middle and hind tibiae with 8 and 10 brown annulations respectively, these annuli con- 17 centrated in the basal *% of these segments; middle and hind tarsi uniform brown; lengths of leg segments as follows: fore fe- mur/tibia/tarsal 1/tarsal 2 = 1.62/1.29/0.07/ 0.25; middle femur/tibia/tarsal 1/tarsal 2/ tarsal 3 = 3.10/4.50/0.10/0.10/10; hind fe- mur/tibia/tarsal 1/tarsal 2/tarsal 3 = 4.50/ 6.60/0.08/0.08/0.10. Abdomen with lateral margins broadly ar- cuate, all tergites lacking raised tumescenc- es posteromedially, first through sixth visi- ble segments dorsally concave, paratergites curved upward and narrowly emarginate, posterior margin of fifth visible tergite broadly V-shaped, sixth visible tergite elon- gate, triangular, domed along longitudinal midline on posterior half; all tergites shin- ing, covered by a sparse, obscure layer of very short, pale, recumbent setae; colora- tion yellowish brown, irregularly mottled with darker brown. Ventral surface medium brown, covered by a sparse layer of short, recumbent gold setae; abdominal paratergites light brown, spiracles whitish. Male genitalia with paramere relatively slender, broadly curving; tip of pygophore narrowed and elongate, apex rounded. Micropterous female: Length 5.30 mm, maximum width (across abdomen) 1.20 mm. Similar to male in general structure and coloration, but with abdomen broader and more strongly expanded; posterior mar- gin of sixth visible tergite broadly V- shaped, rising to a knob-like tumescence posteromedially; seventh visible tergite short, bisected into two lobes along longi- tidinal midline by a deep sulcus, posterior margin broadly rounded (Fig. 4). Distribution.—Hawaii (Volcano area) (Fig. 13). Ecological associations.—Taken on mossy trunks of ohia trees (Metrosideros polymorpha). Material examined.—Holotype, microp- terous 6: HAWAIIAN ISLANDS, Hawaii, reared from eggs laid by female taken at Puu Makaala, 3,700 ft. [1,130 m.], 10 Au- gust 1975, on mossy bark of ohia [Metros- 18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ideros polymorpha], W. P. Mull (BPBM). Paratypes: HAWAIIAN ISLANDS, Ha- waii: 1 2, same data as holotype (BPBM). Etymology.—Named in honor of Wil- liam Mull, who collected the only known specimens and reared numerous individuals through all life stages. Discussion.—Saicella mulli is endemic to Hawaii; the only known series was reared from eggs laid by a single female captured at Puu Makaala, near Volcano. It seems likely that this species will also be found to inhabit windward Mauna Kea and the Kohala Mountains. Saicella perkinsi Polhemus, new species (Figs. 6, 11) Diagnosis.—Recognized by the long, pale, pilose hairs on the pronotum, and the absence of annulations on the middle and hind tibiae, and the enlarged, triangular form of the seventh visible male abdominal tergite (Fig. 6). Description.—Micropterous female: General coloration golden yellowish brown, with darker brown markings on head, thorax and abdomen, red markings laterally on abdominal tergites; legs mul- tiannulate with dark brown. Length 6.10 mm, maximum width (across abdomen) 1.50 mm. Head \ength/width = 0.81/0.55, covered by a thick layer of appressed, wooly golden setae, interspersed with numerous long, pale, erect pilose setae; width of vertex 4.0X dorsal width of an eye (0.40/0.10); length of anterior lobe of head 3.1 dorsal length of an eye (0.44/0.14); eyes small, consisting of approximately 20 ommatidia each; length of posterior lobe of head 2.6 dorsal length of an eye (0.37/0.14); ocelli absent; length of antennal segments I-IV = 4.10/3.75/0.95/0.70; rostrum length 0.97, reaching to bases of fore coxae; coloration of head uniform golden brown, without contrasting markings; antennal segment I uniform golden brown, without annula- tions; antennal segments II-IV medium brown, also lacking annulations. - Pronotum length (midline)/width = 0.81/ 0.90; anterior section golden brown, lateral lobes with thick linear patches of appressed, wooly, golden setae, intermixed with nu- merous long, pale, erect, pilose setae, inter- vening longitudinal sulcus also bearing nu- merous appressed gold setae; posterior sec- tion of pronotum golden brown, bearing nu- merous long, pale, erect, pilose setae, swollen centrally, not produced to a tumes- cence. Scutellum triangular, lateral angles me- dium brown, central section golden brown, produced into an erect, slender, nearly ver- tical spine posteromedially, this spine straight when viewed laterally; numerous long, pale, erect pilose setae present around base of spine; length (including spine)/ width = 0.15/0.25. Hemelytra short, micropterous, reaching to base of erect, slender, golden brown, spine arising nearly vertically from the un- derlying mesonotum, this spine similar in size, shape and orientation to that arising from scutellum; wing pads consisting of tiny flaps, widening slightly on posterior halves, venation and membrane absent; col- oration medium brown, set with a few long, pale, erect, pilose setae. Legs elongate, with fore coxa approxi- mately 4.0 as long as thick (1.0/0.25); fore femur fusiform, 7.4 longer than wide (2.00/0.27); coloration of legs golden brown with dark brown markings; fore fe- mur yellowish brown on basal %, with a broad brown annulation on distal %; fore tibia with 3 diffuse brown annulations, knee pale, apex dark; middle and hind femora each with 4 brown annulations, these an- nulations roughly equal to length of inter- vening pale areas; middle and hind tibiae uniform medium brown, lacking annula- tions; middle and hind tarsi uniform medi- um brown; lengths of leg segments as fol- lows: fore femur/tibia/tarsal 1/tarsal 2 = 2.00/1.00/0.12/0.22; middle femur/tibia/tar- sal 1/tarsal 2/tarsal 3 = 3.90/5.25/0.06/0.06/ 0.07; hind femur/tibia/tarsal 1/tarsal 2/tarsal 3 = 5.70/8.90/0.06/0.06/0.07. VOLUME 102, NUMBER 1 Abdomen yellowish brown, this ground color overlain with irregular dark brown markings centrally on all tergites and pos- terolaterally on paratergites; connexival margins broadly convex, evenly curving; first visible abdominal tergite strongly flexed upward anteromedially, remaining tergites flat, lacking tumescences, poste- rior margin of sixth visible tergite with small V-shaped indentation medially, sev- enth visible tergite triangular (Fig. 6); all tergites shining, covered by a sparse, ob- scure layer of very short, pale, recumbent setae. Ventral surface of head, thorax and ab- domen golden brown, covered by an ob- scure layer of short, pale, recumbent setae, intermixed on mesosternum with numerous long, pale, erect, pilose setae; abdominal paratergites irregularly marked with red bordering connexival margins. Micropterous male: Unknown. Distribution.—Oahu (Waianae Moun- tains) (Fig. 11). Ecological associations.—Taken on mossy tree trunks at night. Material examined.—Holotype, microp- terous 2, HAWAIIAN ISLANDS, Oahu, Waianae Mountains, Kaluaa Gulch, | mi. SE of Puu Hapapa, 700 m. [2,300 ft.], 10 October 1976, on mossy tree trunk, E G. Howarth (BPBM). Paratype: HAWAIIAN ISLANDS, Oahu: 1 2, Waianae Kai Forest Reserve, Kamaileunu Ridge, 720 m. [2,360 ft.], 19 October 1975, north side of ridge on tree trunk at night, W. C. Gagne (BPBM). Etymology.—Named in honor of R. C. L. Perkins, whose pioneering collections of native Hawalian insects have given us an exceptional historical perspective regarding the islands’ insect fauna. Discussion.—Saicella perkinsi is endem- ic to Oahu, and the only specimens so far known have come from the leeward slopes of the Waianae Range. This is one of the drier portions of the island, and it seems curious that this or a related species has not 19 been taken in the wet forests of the Koolau Mountains on eastern Oahu. The long pilose hairs on the pronotum are present in one of the two specimens at hand, but mostly lacking in the other. It is presumed that in the latter case these hairs, being slender and delicate, were rubbed off during the specimen’s collection or subse- quent handling. ACKNOWLEDGMENTS This revision is based primarily on ma- terial collected by the author during field surveys supported by the Smithsonian Insti- tution’s Drake Fund (USNM), or held in the collections of the Bernice P. Bishop Muse- um, Honolulu, Hawaii (BPBM). It could not have been completed without the generous cooperation of the staff in the Department of Entomology at the latter institution, partic- ularly Dr. Scott E. Miller (now with ICIPE, Nairobi), David Preston, Dr. Francis G. Ho- warth, and Gordon Nishida. I also thank Dr. James K. Liebherr of Cornell University (CUIC) and Curtis Ewing, from the Univer- sity of Hawaii, for companionship and as- sistance in the field, and the staff of Pacific Helicopter Tours, Inc., particularly pilots Tom Hauptman and Eric Pacheco, who braved difficult weather over rugged terrain to convey us to sampling sites on the wind- ward slopes of Haleakala. Their skill and de- votion are greatly appreciated. LITERATURE CITED Clague, D. A. and G. B. Dalrymple. 1987. The Ha- waiian-Emporer volcanic chain. Part I. Geologic evolution, pp. 5-54. In Decker, R. W,, T. L. Wright, and P. H. Stauffer, eds., Volcanism in Ha- wail, Vol. 1. USGS Professional Paper 1350. U.S. Government Printing Office, Washington, DC. 839 pp. Langenheim, V. A. M. and D. A. Clague. 1987. The Hawaiian-Emporer volcanic chain. Part II. Strati- graphic framework of volcanic rocks of the Ha- waiian Islands, pp. 55—84. Jn Decker, R. W., T. L. Wright, and P. H. Stauffer, eds. Volcanism in Ha- waii, Vol. 1. USGS Professional Paper 1350. U.S. Government Printing Office, Washington, DC. 839 pp. Leston, D. 1957. Spread potential and the colonisation of islands. Systematic Zoology 6: 41—46. 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Usinger, R. L. 1958. A new genus of Reduviidae from the Hawaiian Islands (Hemiptera). Proceedings of the Hawaiian Entomological Society 16: 437_ 440. Wygodzinsky, PW. 1966. A monograph of the Eme- sinae (Reduviidae, Hemiptera), Bulletin of the American Museum of Natural History 133: ]— 614. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 21-49 REVISION OF LOBOGENESIS RAZOWSKI AND ODONTHALITUS RAZOWSKI (LEPIDOPTERA: TORTRICIDAE: TORTRICINAE), WITH COMMENTS ON THEIR MONOPHYLY JOHN W. BROWN Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, % National Museum of Natural History, Washington, DC 20560- 0168, U.S.A (e-mail: jbrown @sel.barc.usda.gov) Abstract.—The Neotropical tortricid genera Lobogenesis Razowski and Odonthalitus Razowski are revised. Eight species are included in Lobogenesis: L. lobata Razowski (type species) from Costa Rica and Panama; L. penai, new species, from Cochabamba, Bolivia; L. magdalenana, new species, from Colombia and Venezuela; L. larana, new species, from Lara Province, Venezuela; L. contrasta, new species, from Cochabamba, Bolivia; L. antiqua, new species, from Cochabamba, Bolivia; L. peruviana, new species, from Peru; and L. varnicosa, new species, from Argentina. Nine species are included in Odonthalitus: O. lacticus Razowski (type species) from Durango, Mexico; O. bisetanus, new species, from Oaxaca, Mexico; O. improprius, new species, from Oaxaca, Mexico; O. conservanus, new species, from Jalisco, Mexico; O. orinoma (Walsingham), new combination, from Guerrero, Mexico; O. fuscomaculatus, new species, from Michoacan, Mexico; O. poas, new species, from Alajuela Province, Costa Rica; O. viridimontis, new species, from Puntarenas Province, Costa Rica; and O. regilla (Walsingham), new com- bination, from Guatemala. While the monophyly of Lobogenesis is well supported by characters of the male and female genitalia, Odonthalitus, as currently defined, may be paraphyletic with respect to Lobogenesis. Key Words: Phylogeny, systematics, South America, Central America, Anopina, Euliini, new species, genitalia Razowski (1992) proposed Odonthalitus to accommodate the single species O. lac- ticus Razowski, described from two speci- mens (1 6, 1 @) collected in Durango, Mexico. In the same paper, he described the monotypic genus Lobogenesis for the new species L. lobata Razowski, represented by a single male from Costa Rica. Although the male genitalia are moderately divergent, these two monotypic genera share a variety of morphological features and are nearly identical in facies. During the course of continued studies on the systematics of the New World Euliini, I discovered numerous undescribed and two described (misplaced) species that are morphologically and super- ficially similar to these two genera, stimu- lating an assessment of the phylogenetic re- lationship between the two genera and de- scriptions of the new species. A phyloge- netic hypothesis based primarily on structures of the male and female genitalia indicates that while Lobogenesis is mono- phyletic, its separation from Odonthalitus may leave the latter paraphyletic. Although similar in facies to Anopina Obraztsov, the male and female genitalia of Odonthalitus and Lobogenesis are highly 22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON divergent from all other Euliini, with a bi- furcate uncus in the male and greatly re- duced apophyses anteriores in the female. The subbasal attachment of the socius and the free (unjoined) distal arms of the gna- thos of Lobogenesis are reminiscent of some Sparganothini. However, these char- acter states are unlikely to be homologous with those found in Sparganothini, as the two groups have little else in common. Some females of Odonthalitus have bilobed papillae anales reminiscent of those found in some species of Apotoforma Busck (Tor- tricini) (see Razowski 1984, 1993); the greatly reduced apophyses anteriores also are similar in the two. However, because the groups share no other derived character states and male genitalia of the two are highly divergent, these similarities in fe- male genitalia are suspected to represent convergence rather than evidence of com- mon ancestry. If the bilobed papillae anales and greatly reduced apophyses eventually are demonstrated to be homologous _ be- tween Odonthalitus and Apotoforma, it would provide evidence for the placement of Odonthalitus and Lobogenesis in Tortri- cini rather than Euliini. The purposes of this paper are to describe 7 new species of Lobogenesis and 6 new species of Odonthalitus, propose two new combinations in Odonthalitus, present a hy- pothesis of the phylogenetic relationships among the species, and modify the descrip- tions of the genera based on the new taxa. MATERIALS AND METHODS Specimens (n = 90) were obtained from the following institutions: The Natural His- tory Museum (BMNH), London, England; Instituto Nacional de Biodiversidad (IN- Bio), Santo Domingo, Heredia, Costa Rica; San Diego Natural History Museum (SDNHM), San Diego, California, U.S.A.; Essig Museum of Entomology, University of California, Berkeley (UCB), U.S.A; Na- tional Museum of Natural History (USNM), Smithsonian Institution, Wash- ington, D.C., U.S.A. Dissection methodology follows that summarized in Brown and Powell (1991). Illustrations of genitalia were drawn with a camera-lucida attachment on a dissecting microscope. Forewing measurements were made with an ocular micrometer mounted in a dissecting microscope. Terminology for wing venation and genitalic structures fol- lows Horak (1984). Abbreviations and sym- bols are as follows: FW = forewing; HW = hindwing; DC = discal cell; n = number of specimens examined; ca. (circa) = ap- proximately; « = mean. Upper side refers to dorsal surface of wings, under side to ventral surface. Because most species are extremely similar in facies, not all are il- lustrated. In contrast, male and female gen- italia are diagnostic and are illustrated for each species. Dissections of the genitalia are required for accurate species determi- nations, and comparison with the illustra- tions is the best means for identifying spec- imens. PHYLOGENY A phylogenetic analysis was conducted on the 17 taxa recognized as species (plus a putative out-group—Anopina Obraztsov). The analysis was based on 23 morpholog- ical characters, 15 binary and 8 multi-state, that vary among the species of Lobogenesis and Odonthalitus, and exhibit shared, de- rived states at the species level. These in- clude 1 character of the head, 2 characters of the thorax, 15 characters of the male gen- italia, and 5 characters of the female geni- talia. Character state polarity was deter- mined using the out-group method. Because sister group relationships are poorly under- stood within Euliini (e.g., Brown and Pow- ell 1991), selection of an out-group was somewhat arbitrary. The genus Anopina Obraztsov was chosen because of its similar forewing pattern and long antennal cilia in the male. Horak’s (1984) assessment of tax- onomically significant structures in Tortri- cinae was used for determination of char- acter state polarity for characters that could not be determined convincingly using the VOLUME 102, NUMBER 1 out-group (e.g., uncertain homology, both states present in out-group). The characters, along with their putative plesiomorphic and apomorphic conditions, are presented in Ta- ble 1. The distribution of the character states among the 17 taxa and the out-group is presented in Table 2; ‘*?” is used where the character state is unknown (e.g., char- acters for species represented by a single sex). The data set was subjected to parsimony analysis using Hennig86 version 1.5 (Lip- scomb 1994), employing the “‘mhennig*”’ command. This algorithm constructs trees, each by a single pass through the data, by adding the taxa in a different sequence each time, and then applies branch-swapping to each of the trees. The analysis generated 6 equally parsimonious trees with a length of 48, a consistency index of 0.75, and a re- tention index of 0.85. Because over half of the taxa (n = 9) are represented by a single sex, there is a considerable amount of miss- ing data, which resulted in the equivocal placement of several taxa. The cladogram illustrated in Fig. | rep- resents the “‘majority rule’’ consensus tree (see Lipscomb 1994) and is identical to one of the six trees. It is considered the best working hypothesis of the phylogeny of the species. The analysis demonstrates that the monophyly of Lobogenesis is well support- ed (i.e., in all 6 trees), while Odonthalitus, as current defined, may be paraphyletic (i.e., in all 6 trees). The hypothesis of the phylogeny is fairly concordant with the geographic distribution of the two genera: with the exception of L. lobata (from Costa Rica and Panama), Lobogenesis appears to be restricted to South America, while Odonthalitus is entirely Central American (including northern Mexico) in distribution. Superficially adults of Odonthalitus are indistinguishable from those of Lobogene- sis. Synapomorphies for the two include the forewing pattern, loss of the male foreleg hairpencil, elongate cilia of the male anten- na, bifurcate uncus, and reduced apophyses anteriores. The monophlyly of Lobogenesis 23 is supported convincingly by the following: uncus bifurcations long, slender, and atten- uate apically; gnathos arms free distally or only weakly attached by membrane; socius large, subbasally attached, with dorsal lobe; and linear row of tiny spinelike teeth in api- cal region of valva. In contrast, Odonthal- itus Was portrayed as paraphyletic in each of the’6 ‘trees: There are at least three possible ways to resolve the paraphyly. One solution would be to consider the entire clade as a single genus to which either Odonthalitus or Lo- bogensis could be applied (based on the choice of the first revisor). Another would be to include lacticus and regilla with the species considered Lobogenesis in Fig. 1, which would result in the synonymy of Lo- bogenesis with Odonthalitus, and to de- scribe a new genus for the remaining Cen- tral American species. The third would be to maintain Lobogenesis as shown in Fig. 1, restrict Odonthalitus to the species lac- ticus and regilla, and propose a new genus for the remaining Central American species. However, for the present the generic status of both Lobogensis and Odonthalitus are re- tained for a number of reasons. First, the extremely long, slender aedeagus of regilla and the notched papillae anales of lacticus are extremely similar to those of O. viridi- montis and O. poas, suggesting a closer phylogenetic relationship of these species than is portrayed in the cladogram. Second- ly, from a biogeographic perspective, one would expect /acticus and regilla (both from Mexico) to be more closely allied to Odonthalitus than to Lobogenesis. Finally, because the data set for the analysis con- tains so much missing data (i.e., several species are represented by a single sex), any classification based on the results of the parsimony analysis likely would be inher- ently unstable. Hence, a conservative ap- proach is employed that maintains the sta- bility of the generic nomenclature of the group. 24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Characters for phylogenetic analysis of Odonthalitus and Lobogenesis; 0 = plesiomorphic state, 1— 3 = apomorphic states; figure references to apomorphic condition(s), where illustrated. 1. Head O—length of male antennal cilia equal to width of flagellomere 1—length of male antennal cilia 3 times width of flagellomere 2. Thorax O—male foreleg hairpencil present 1—male foreleg hairpencil absent 3. Forewing O—pattern variable 1—pattern with distinctive costal and basal patch 4. Uncus O—unmodified, without bifurcate distal portion 1—bifurcate in distal one-half (Figs. 2—8) 2—bifurcate from base (Figs. 14—18) 3—bifurcations widely separate at base (Figs. 19—20) 5. Uncus O—tips of bifurcation round or blunt 1—tips of bifurcation narrowed and pointed (Figs. 2—5) 2—1ips with narrow triangular enlargement (Figs. 6—8) 6. Gnathos Q—gnathos arms joined distally 1—-gnathos arms weakly joined distally (Figs. 14—16) 2—gnathos arms separate distally (Figs. 2—8, 19-20) 7. Gnathos OQ—arms without lateral process(es) 1—arms with lateral lobes (Figs. 17—18) 8. Socius O—simple, small, slightly digitate 1—large, with moderate dorsal lobe (Figs. 2—5) 2—huge, with large dorsal lobe (Figs. 6-8) 9. Transtilla O—simple bridge 1—lost (Figs. 17—20) 10. Valva Q—simple at base 1—with patch of curved spines at base of costa (Figs. 14-15) 2—with large hook at base of casta (Fig. 16) 3— costa with free hairy lobe near middle (Figs. 17—8) 11. Valva O—costa simple 1—costa with free flange ca. 0.67 distance from base (Figs. 3—5) 2— flange enlarged (Fig. 4) 12. Valva O—simple 1—dense patch of setae in middle of basal one-third (Figs. 4, 6-8) 13. Valva O—simple 1—curved, sclerotized line from patch of setae to apex (Figs. 6-8) 14. Valva Q—apex more or less rounded 1—apex attenuate, somewhat pointed (Figs. 14—18) 15. Valva O—unmodified 1—linear row of tiny spinelike teeth in apical region (Figs. 2—8) 16. Valva O—simple 1—with notch at venter (Figs. 19-20) 17. Aedeagus O—simple, unmodified 1—somewhat undulate (Figs. 3—5) 2— strongly bent, somewhat *‘J°’ shaped (Figs. 14—16) 18. Aedeagus O0—simple, unmodified 1—extremely long, slender (Figs. 17—19) 19. Sterigma O—simple 1—with scobinate lateral patches (Figs. 21—22) 20. Sterigma O—simple 1—rounded or v-shaped notch at ostium (Figs. 9, 12) 21. Sterigma O—simple 1—-with sclerotized arch surrounding ostium (Figs. 10-11, 13) 22. Papillae O—simple, parallel-sided 1—with a mesal notch (Figs. 23—25) 23. Apophyses ant. O—unmodified, moderately long 1—reduced in length (Figs. 21—22) 2— reduced to a short nub (Figs. 9-13, 25—26) 3— lost (Figs. 23-24) VOLUME 102, NUMBER 1 Table 2. Distribution of character states among species; see Table | for explanation of character states (? = missing data). Outgroup 0000 0000 lacticus Tas} 0200 regilla dais 0200 poas Deda 0010 viridimontis RaW Eal 0010 orinoma Deer? 2222 fuscomaculatus Fo plee) BPP conservanus LI 0100 bisetanus ua 0100 improprius 112 0100 contrasta iltaleallslk 2202 peruviana ita ak 22:02 antiqua iBall 2202 penai abaya AL 1201 lobata ibaa 1201 larana alga Lace 1201 magdalenana algal aleACO)atk varnicosa PAM? De De SYSTEMATICS Lobogenesis Razowski 1992 Lobogenesis Razowski 1992: 213; Powell et al. 1995: 144. Type species.—Lobogenesis lobata Ra- zowski 1992, by monotypy. Redescription.—Head: Antennal cilia ca. 3.0 times width of flagellomere in male, ca. 0.5 times width of flagellomere in fe- male. Labial palpus (segments II and III combined) ca. 1.5 times horizontal diameter of compound eye; segment II weakly up- turned, slightly expanded distally by scal- ing; segment III ca. 0.4 as long as II, smooth-scaled, well exposed. Maxillary palpus rudimentary. Frons with overhang- ing tuft of scales. Ocelli small or absent. Chaetosema present. Proboscis present. Thorax: Smooth-scaled. Legs unmodified, male without foreleg hairpencil. Forewing: Length 2.3—2.4 times width; length of DC about 0.55 times FW length; width of DC about 0.20 DC length; CuA, originates about 0.60 along length of DC; all veins separate beyond DC; chorda and M-stem absent. No upraised scale tufts; male with- out costal fold. Hindwing: Sc+R and Rs closely approximate; M, and CuA, closely approximate; M-stem absent; tuft of hair- 0000 0000 0000 000 1000 0101 0000 012 1000 0001 01?? iMeue 1300 0100 0100 Oalbs 1300 0100 0100 013 Dye ONe Dp Be 2200 002 222? 2 Bee 220 O01 0200 0100 2010 001 0100 0100 20/2? Ree 0100 0100 202-2 Due? 0001 1010 00?? Dee, 0001 1010 0000 02 0001 1010 00?? 222 0000 0010 00?? eae 0010 0010 1001 002 0021 0010 1000 NO? 0010 0010 1001 002 apie 2 2D? ??00 102 like scales along 1A+2A, originating near base of wing. Abdomen: Dorsal pits absent; no modified corethrogyne scaling in female. Male genitalia: Uncus bifurcate from a common, slender stalk; each arm slender, attenuate distally or with triangular enlarge- ment at tip. Socius broad and long, with dorsal lobe, not fused to gnathos. Gnathos simple, non-dentate, arms narrow, separate distally. Subscaphium and hami absent. Transtilla a simple band or enlarged, ill-de- fined bridge. Valva moderately short, slight- ly attenuate distally; row of tiny spinelike teeth in middle of apical region. Pulvinus absent. Vinculum complete. Juxta a scler- otized plate. Aedeagus moderately long, usually weakly undulate; phallobase sim- ple; cornuti absent. Female genitalia: Pa- pillae anales narrow. Apophyses posteriores moderately long to short; apophyses anter- iores reduced to a membranous nub. Sterig- ma a simple sclerotized band. Ductus bur- sae moderately long, membranous. Corpus bursae frail, elongate, pear-shaped; spicules and signum lacking. Accessory bursa fre- quently present. Diagnosis.—Superficially, adults of Lo- bogenesis are indistinguishable from those of Odonthalitus. The two can be separated by the following characters of the male gen- 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Odonthalitus Lobogenesis = = aie : =) a = = wn 3 @ = STS, Ss = S > «(68 paees eS Sisige s a S Ss OTs = Siti PSS Sy So Saw eee Bes | = us Be tS te SO OmeS: Se Se Oe a ee eee) ee = 8 Ny ~ iS) Y Sy Sy Os = So 3S = SOS SS oss SS Soe pees Sr oS FG ae See, SS @ Ss S856 85 Ss See oS eo as 16-14 19 ics aoe 10' a ae 11! 5! 18 9 -17 12 23. 20 Tait ual Ball fc; ae eae niet 17 22 16 iS 9 8 le = 1 aa. 10 6! ao 6 q 3 2 1 Fig. 1. Consensus cladogram of phylogeny of Lobogenesis and Odonthalitus. Prime (') and double prime (") refer to successive steps in multi-state characters; minus (—) refers to character state reversal. italia: 1) paired uncus arms (bifurcations) always from a common stalk in Loboge- nesis, either from a common stalk or sepa- rated basally in Odonthalitus; 2) distal tips of paired uncus arms (bifurcations) ex- tremely slender, pointed or triangular dis- tally in Lobogenesis, blunt or rounded in Odonthalitus; 3) arms of gnathos free dis- tally in Lobogenesis, joined distally in Odonthalitus, either as a single process or weakly attached by membrane; and 4) so- cius attached subbasally, with a large dorsal lobe in Lobogenesis, socius attached basally in Odonthalitus. Distribution and biology.—Lobogenesis is known from montane forest habitat from Costa Rica south to Argentina, from 1,100— 3,000 m elevation. Adults have been col- VOLUME 102, NUMBER 1 lected at lights. Nothing is known of the early stages. Three species are known to be sympatric in Bolivia (at Cochabamba) and two in Venezuela (at Rancho Grande). Remarks.—Species of Lobogenesis are rare in collections; the 8 species treated be- low are represented by 37 specimens, 14 of which represent one species. Three species are known only from males and one only from females. Modern systematic revisions of Neotropical tortricids (e.g., Brown and Powell 1991, 2000) typically result in an increase in the number of recognized spe- cies by a factor of 3—4; the present treat- ment results in descriptions of 7 new spe- cies to accompany the single previously de- scribed species. Lobogenesis penai J. Brown, new species (Figss2; 32) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale cream; roughened above, pale cream. La- bial palpus pale cream mesally, mostly brown laterally. Antennal scaling bronze. Thorax: Tan with dark brown tegula. Fore- wing: Length 6.8 mm (n = 1). Upper side pale cream, with irregular, pale brown striae throughout; basal 0.2 dark brown; dark brown, triangular patch bordering costa ca. 0.45—0.60 distance from base to apex. Un- der side nearly uniform dark tan with faint indication of upper side markings. Hind- wing: Upper side white, with faint gray- brown mottling. Under side light gray brown with darker mottling. Genitalia: As in Fig. 2 (drawn from USNM slide 88523; n = 1). Uncus slender at base, bifurcate in distal 0.7, with extremely long, slender spines subapically. Socius large, hairy, with conspicuous portion dorsad of attachment. Gnathos arms extremely slender, elongate, separate distally. Transtilla large, membra- nous, with narrow sclerotized arch in ven- tral portion. Valva broadest at base, rounded apically; basal 0.5 of costa sclerotized. Ae- deagus short, somewhat pistol-shaped, with 27 a short lateral thorn; phallobase with small rounded membranous cap. Female. Unknown. Type.—Holotype, ¢, Bolivia, Incachaca, Cochabamba, tropical cloud forest area, 2,100.1 27-W I/S-1X- 1956 (LL Pena, USNM). Diagnosis.—The male genitalia of L. penadi are most similar to those of L. larana and L. lobata. They can be distinguished from those species by the much longer, more slender gnathos arms and the elon- gate, slender bifurcations of the uncus, with long spines subapically (Fig. 2). Etymology.—The species name is a pa- tronym for Louis Pena, a noted Chilean en- tomologist and the collector of the holo- type. Lobogenesis lobata (Razowsk1) (Figs. 3; 9) Lobogenesis lobata Razowski 1992: 215; Powell et al. 1995: 144. Redescription.—Male. Head: Frons with sparse, smooth scaling below mid- eye, pale light brown; roughened above, pale tan. Labial palpus pale tan mesally, brown mixed with tan laterally. Antennal scaling brown. Thorax: Whitish, with brown tegula. Forewing: Length 5.5 mm (n = 2). Upper side whitish, with faint, sparse tan overscaling; basal 0.2 brown; brown semicircular patch bordering costa ca. 0.45—0.65 distance from base to apex; termen with irregular brown striae. Under side uniform dark tan with faint indication of upper side markings. Hindwing: Upper side whitish gray with slightly darker gray- brown mottling. Under side light gray- brown with darker mottling. Genitalia: As in Fig. 3 (drawn from USNM slide 89442, Panama; n = 2). Uncus narrow at base, bifurcate in distal 0.65, each arm slender, elongate, pointed distally. Socius with free dorsal lobe. Gnathos arms slender, weakly joined near middle, rounded distally, to- gether forming an H-shaped process. Tran- stilla a narrow sclerotized band with a pair 28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-5. Male genitalia of Lobogenesis; valvae spread, aedeagus removed. 2, L. penai. 3, L. lobata. 4, L. larana. 5, L. magdalenana. VOLUME 102, NUMBER 1 of dorsally projecting thorns at middle. Valva nearly rectangular, rounded apically; costa with free flange near middle; narrow sclerotized ridge across face of valva ca. 0.65 distance from base to apex; small lin- ear patch of short, spinelike teeth in apical region. Aedeagus weakly undulate; phal- lobase with large, rounded, membranous cap. Female. FW length 5.0 mm (n = 2). Su- perficially as in male, except thorax dark brown and antennal cilia short. Genitalia: As in Fig. 9 (drawn from USNM< slide 88441, Panama; n = 2). Papillae anales un- modified. Apophyses posteriores moderate; apophyses anteriores extremely short, semi- membranous. Sterigma a sclerotized band, with U-shaped excavation at ostium. Duc- tus bursae relatively long, slender, frail. Corpus bursae rounded. Type.—Holotype, 6, Costa Rica, Puntar- enas Province, 6 km S San Vito, 20/27-IV- 1967 (D. Veirs, UCB). Additional specimens examined.— COSTA RICA: Puntarenas Province: Fea. Cafrosa, Est. Las Mellizas, P. N. Amistad, 1,300 m, 1 2, X-1990 (M. Ramirez & G. Mora, INBio). PANAMA: Cerro Cam- pana, nr. Chica, 1 6, 2/5-IV-1965 (S. S. & W. D. Duckworth, USNM); Cerro Cam- pana, 1 2, 11/14-VII-1967 (O. S. Flint, USNM). Diagnosis.—Lobogenesis lobata is most similar to L. larana and L. magdalenana in several features of the male genitalia. It can be distinguished from those species by the paired thorns from the middle of the transtilla. The sexes were associated by their common occurrence at Cerro Cam- pana. Lobogenesis larana J. Brown, new species (Figse4. 133) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, whitish; roughened above, pale tan. Labial palpus whitish mesally, mostly brown lat- erally. Antennal scaling brown. Thorax: 29 Whitish with dark brown tegula. Forewing: Length 5.2—5.8 mm (* = 5.4; n = 4). Upper side pale whitish, with variable, faint brown striae and reticulations, particularly dense in distal 0.5; basal 0.2 brown; brown semicir- cular patch bordering costa ca. 0.45—0.65 distance from base to apex; termen with ir- regular brown line. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side pale whit- ish gray with slightly darker gray-brown mottling. Under side light gray-brown with darker mottling. Genitalia: As in Fig. 4 (drawn from USNM slide 89444; n = 3). Uncus narrow at base, bifurcate in distal 0.65, each arm slender, pointed distally. So- cius with dorsal lobe. Gnathos arms slender, separate. Transtilla a moderate, evenly sclerotized band. Valva nearly rectangular, rounded apically; an attenuate flange from costa just beyond middle, ending in free, rounded process; sacculus represented by undulate line of sclerotization; a dense patch of setae in basal 0.33 of valva; small linear patch of spinelike teeth in apical re- gion. Aedeagus relatively slender, slightly undulate; phallobase with small membra- nous cap. Female. FW length 6.5—7.0 mm (* = 6.8; n = 6). Superficially as in male, except brownish overscaling of forewing conspic- uously more dense, larger average forewing length, and lacking elongate antennal cilia. Genitalia: As in Fig. 11 (drawn from USNM slide 87880; n = 3). Papillae anales slender, unmodified. Apophyses posteriores slender; apophyses anteriores reduced, slen- der, semi-membranous. Sterigma a simple band with membranous region surrounding ostium, bordered by inverted U-shaped sclerotized region. Ductus bursae moderate- ly long. Corpus bursae oblong. Types.—Holotype, ¢, Venezuela, Lara, Yacambu National Park, 13 km SE Sana- re, cloud forest, 4,800’ [1,500 m], black- light, 4/7-III-1978 (J. B. Heppner, USNM). Paratypes, 3 6, 10 2. VENEZUELA: 3 6, 9 @, same data as holotype. Aragua 30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Province: Rancho Grande, 1,100 m, 1 2, 1/ 7-VIII-1967 (R. W. Poole, USNM). Diagnosis.—The male genitalia of Lo- bogenesis larana are most similar to those of L. magdalenana and L. lobata (Figs. 3— 5) in the overall shape of the valva, the lin- ear patch of spinelike teeth in the apical re- gion of the valva, and the slender bifurca- tions of the uncus. The three can be sepa- rated by the presence of a dense patch of setae in the basal 0.33 of the valva in L. larana and the mesal pair of spines from the transtilla in L. lobata, both features of which are lacking in L. magdalenana. Remarks.—The genitalia of the female from Rancho Grande differ slightly in the shape of the sterigma from those from Ya- cambu. Etymology.—The specific epithet refers to the Venezuelan province of Lara. Lobogenesis magdalenana J. Brown, new species (Bigs 257412,030. 37) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, whitish; roughened above, pale bronze- brown. Labial palpus whitish mesally, brown mixed with tan laterally. Antennal scaling brown. Thorax: Brown. Forewing: Length 5.0 mm (n = 2). Upper side pale whitish, with faint brown striae in distal 0.5; basal 0.2 brown; brown semicircular patch bordering costa ca. 0.45—0.65 dis- tance from base to apex. Under side uni- form dark tan with faint indication of up- perside markings. Hindwing: Upper side whitish with plae gray-brown mottling. Under side light gray-brown with darker mottling. Genitalia: As in Fig. 5 (drawn from BMNH slide 29065, Colombia; n = 2). Uncus narrow at base, bifurcate in dis- tal 0.65, each arm slender, elongate, point- ed distally. Socius with dorsal lobe. Gna- thos arms slender, apparently joined dis- tally by membrane. Transtilla a narrow sclerotized bridge, weakly arched mesally. Valva nearly rectangular, rounded apically; costa with roughened flange in basal 0.5; narrow sclerotized ridge across face of val- va, terminating in free costal flange; linear patch of short, spinelike teeth in apical re- gion. Aedeagus weakly undulate; phallo- base with large, rounded, membranous cap. Female. FW length 5.1—6.5 mm (* = 5.9; n = 3). Superficially as in male, except forewing length larger, forewing with brown overscaling more dense, and lacking elongate antennal cilia. Genitalia: As in Fig. 12 (drawn from BMNH slide 29064; Colombia; n = 3). Sterigma a sclerotized band, with membranous area surrounding ostium; ostium indicated by a sclerotized V- shaped notch. Ductus bursae moderately long. Corpus bursae oblong. Types.—Holotype, 6, Colombia, Mag- dalena, Sierra Nevada de Santa Marta, San Pedro de la Sierra, 1,500 m, 14-VIII- 1973, Oxford Expedition to Colombia (BMNH). Paratypes, 1 ¢6, 2 2. COLOMBIA: Mag- dalena, Sierra Nevada de Santa Marta, San Pedro de la Sierra, 1,500 m, 1 @, 6-VII- 1997 (BMNH); Sierra del Libano, 6,000’ [1,846 mj]; B® 2, V-1899 (HB. “Hin Smith, BMNH). Additional specimens examined.—VEN- EZUELA: Aragua: 1 km south of Rancho Grande; 1d, 5-11-1976 (C2 Ms & Os! Flint, USNM); Rancho Grande, cloud for- est, 1,100 m,; 1 2, 30/31-I11-1978 (J. B: He- ppner, USNM); T. EK Amazonas, Cerro de Neblina, Camp VII, 1,850 m, 0°51'N, 65°58'""W, 1 &, 2/4-XTI-1984 (R. Brown, USNM). Diagnosis.—Lobogenesis magdalenana is most similar to L. larana and L. lobata; the differences in male genitalia are dis- cussed above in the diagnosis of L. larana. The female genitalia can be distinguished by the shape of the mesal notch of the ste- rigma: V-shaped in L. magdalenana and more U-shaped in L. larana. Remarks.—Male and female genitalia of the specimens from Venezuela, deviate slightly from those of specimens from Co- lombia: the linear row of fine spinelike VOLUME 102, NUMBER 1 31 Figs. 6-8. Male genitalia of Lobogenesis: valvae spread, aedeagus removed. 6, L. peruviana. 7, L. antiqua. 8, L. contrasta. 32) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON teeth in the apical region of the valva con- sists of considerably fewer teeth in the male, and the antrum is slightly more rounded in the female. In addition, fore- wing length is conspicuously less in the fe- male from Rancho Grande. However, be- cause the genitalic differences are consid- erably less than that between other species of Lobogenesis, they are interpreted as geo- graphic, infraspecific variation within L. magdalenana. Additional specimens are necessary to determine whether or not this interpretation is correct. Etymology.—The specific epithet refers to the province of Magdalena. Lobogenesis contrasta J. Brown, new species (Figs. 8, 27) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, tan-brown; roughened above, dark cop- per-brown Labial palpus pale tan mesally, dark red-brown laterally. Antennal scaling brown. Thorax: Mostly dark brown. Forewing: Length 6.8—-8.0 mm (x = 7.2; n = 6). Upper side white, with faint, dif- fuse, pale yellow-tan overscaling; basal 0.2 dark brown with some red-brown scales; dark brown, semicircular patch bordering costa ca. 0.45—0.60 from base; two small, brown, triangular dots on costa between large costal patch and apex; ter- men with diffuse brownish striae, with a few pale yellow-green scales; a varibale, ill-defined line from dorsum near tornus. Under side nearly uniform tan-brown. Hindwing: Upper side dingy pale yellow with pale gray-brown mottling. Under side light gray-brown with darker mot- tling. Genitalia: As in Fig. 8 (drawn from USNM slides 68611 and 89440; n = 3). Uncus slender from broad, expanded cap on dorsum of tegumen, bifurcate in distal 0.5, with swollen, foot-shaped processes apically. Socius large, hairy, with con- spicuous portion dorsad of attachment. Gnathos comparatively slender, arms free distally, with shovel-shaped, attenuate tip. Transtilla complete, more strongly scler- toized mesally. Valva broadest at base, slightly attenuate distally, with a pair of triangular lobes at costa, one at base, con- tiguous with transtilla, the second as a free rounded flange near mid-costa; a densely setose region in basal 0.33, con- tinuing toward apex as a long, curved, sclerotized line. Aedeagus relatively slen- der, slightly curved; phallobase with a large, rounded, membranous cap. Female. Unknown. Type.—Holotype, 6, Bolivia, Incachaca, Cochabamba, tropical cloud forest area, 2,100 m, 27-VIII/5-IX-1956 (L. Pena, USNM). Paratypes, 5 ¢, same data as holotype. Diagnosis.—Lobogenesis contrasta 1s most similar to L. peruviana and L. antiqua in size, forewing maculation, and male gen- italia. It be can be distinguished from L. peruviana by its simple aedeagus—that of L. peruviana has a distinct ventral thorn from near the middle. It can be separated from L. antiqua by the shape of the scler- otized ridge across the face of the valva (see Figs. 6-7), which is more strongly curved in the latter. Etymology.—The specific epithet is an adjective, referring to the contrast between the pale ground color and the forewing pat- tern elements. Lobogenesis peruviana J. Brown, new species (Figs. 6, 10, 29) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, whitish and pale tan; roughened above, pale tan. Labial palpus pale tan mesally, brown mixed with yellow-tan laterally. Antennal scaling brown. Thorax: Mostly gray white, with red-brown prothoracic collar and brown tegula. Forewing: Length 7.5 mm (n = 1). Upper side silver white, with faint pale yellowish overscal- ing; basal 0.2 dark red-brown; dark red- brown semicircular patch bordering costa ca. 0.45—0.60 from base, second smaller VOLUME 102, NUMBER 1 33 S— S — O SS “ Figs. 9-11. Female genitalia of Lobogenesis. 9, L. lobata. 10, L. peruviana. 11, L. larana. 34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON patch at ca. 0.75 distance from base; ter- men with diffuse gray-brown striae, with a few pale green and yellow sclaes. Under side nearly uniform tan brown. Hindwing: Upper side whitish with pale grayish mot- tling. Under side light grayish with darker mottling. Genitalia: As in Fig. 6 (drawn from BMNH slide 29061; n = 1). Uncus slender from enlarged cap on dorsum of tegumen, bifurcate in distal 0.5; each tip bearing an enlarged triangular process, with two long, fine, subapical setae. Socius large, hairy, with conspicuous portion dor- sad of attachment. Gnathos arms compar- atively slender, separate distally, each with a shovel-shaped, attentuate tip. Transtilla complete, weakly U-shaped, with small upturned flap from venter near middle. Valva broadest at base, slightly attenuate apically, with a slender, digitate projection from costa at base; densely setose area in basal 0.33, extending toward apex as curved, sclerotized line. Aedeagus undu- late, moderately broad, with distinct thorn from venter near middle; phallobase with large, rounded, membranous cap. Female. FW length 8.0 mm (n = 1). Su- perficially as in male, except lacking elon- gate antennal cilia. Genitalia: As in Fig. 10 (drawn from BMNH slide 29062; n = 1). Papillae anales unmodified. Apophy- ses posteriores moderate in length; apoph- yses anteriores reduced to short, slender, semi-membranous nub. Sterigma a broad, sclerotized band, with inverted U-shaped membranous region surrounding ostium; ostium represented by small U-shaped sclerite. Ductus bursae relatively short, slender, frail. Corpus bursae rounded, pear-shaped. Types.—Holotype, ¢, Peru, Pillahuata, Cuzco, 2,600 m, 14/16-VIII-1982 (M. Mat- thews & M. Packer, BMNH). Paratype, 1 @, same data as holotype (BMNBH). Diagnosis.—Lobogenesis peruviana is the largest species in the genus and is most similar to O. contrasta. It can be distin- guished by the characters cited above in the diagnosis for O. contrasta. Autapomorphies for L. peruviana include the triangular api- cal processes and setae of the distal portion of the uncus, the upturned mesal ventral portion of the transtilla, and the thorn from the venter of the aedeagus. Etymology.—The specific epithet is an adjective referring to the country of its cap- ture. Lobogenesis antiqua J. Brown, new species (Figs. 7, 28) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, tan- brown; roughened above, bronze-brown. Labial palpus pale tan mesally, mostly brown laterally. Antennal scaling brown. Thorax: Dark brown mixed with red brown. Forewing: Length 7.8 mm (n = 1). Upper side dingy white, with faint pale tan-yellow overscaling; basal 0.2 dark brown; dark brown semicircular patch bor- dering costa ca. 0.45—0.60 from base, two additional small costal spots between semi- circular patch and apex; termen with irreg- ular brownish reticulations. Under side uniform tan-brown with faint indication of upper side markings. Hindwing: Upper side whitish with pale grayish mottling. Under side light grayish with darker mot- tling. Genitalia: As in Fig. 7 (drawn from BMNH slide 29063; n = 1). Uncus slender from broad, enlarged cap on dorsum of tegumen; uncus missing distal one-half (assumed to be bifurcate). Socius large, hairy, with conspicuous portion dorsad of attachment; weakly sclerotized along outer edge. Gnathos arms comparatively slender, arms separate distally, each with a trian- gular tip. Transtilla complete, most strong- ly sclerotized mesally. Valva broadest at base, weakly attenuate apically, with two projections from costa, one at base contig- uous with transtilla, the second a free, rounded flange at ca. mid-costa; densely setose area in basal 0.33, extending toward apex as curved, sclerotized line. Aedeagus VOLUME 102, NUMBER 1 Figs. 12-13. nearly straight, relatively slender; phallo- base with broad, rounded, membranous cap. Female. Unknown. Type.—Holotype, 6, Bolivia, Yungas del Espiritu Santo, Cochabamba, 1888-89, Par- avicini Coll., BM 1937-383 (P. Germain, BMNH). Diagnosis.—Lobogenesis antiqua is most similar to L. contrasta; superficially, it has a more yellowish tan forewing ground col- or. The male genitalia of L. antiqua can be Female genitalia of Lobogenesis. 12, L. magdalenana. 13, L. varnicosa. distinguished from those of L. contrasta by the conspicuously longer linear row of spinelike teeth in the apical region of the valva, the larger patch of setae in the basal 0.33 of the valva, and the more undulate sclerotized line extending from the setose patch to the apex of the valva (see Figs. 7— 8). Etymology.—The specific epithet, from the word antique, is an adjective referring to the fact that the type specimen was col- lected more than 100 years ago. 36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Lobogenesis varnicosa J. Brown, new species (Figs= 1333) Description.—Male. Unknown. Female. Head: Frons with sparse, smooth scaling below mid-eye, whitish and pale tan; roughened above, pale tan. Labial palpus pale whitish mesally, light brown laterally. Antennal scaling brown. Thorax: Pale tan. Forewing: Length 6.5—7.2 mm (x = 6.8; n = 2). Upper side dingy white, with faint brownish overscaling and darker brown striae; basal 0.2 dark brown; dark brown triangular patch bordering costa ca. 0.45—0.60 from base, with a dark brown dash immediately below, near apex of DC. Under side nearly uniform tan brown. Hindwing: Upper side dingy whitish with pale gray-brown overscaling and mottling. Under side light grayish with darker mot- tling. Genitalia: As in Fig. 13 (drawn from USNM slide 89267; n = 2). Papillae anales unmodified. Apophyses posteriores moder- ate; apophyses anteriores reduced to short, slender, semi-membranous nub. Sterigma a broad, weakly sclerotized band, with in- verted U-shaped arch dorsad of ostium. Ductus bursae relatively long, undifferen- tiated from corpus. Corpus bursae long, moderately slender. Types.—Holotype, 2, Argentina, Tucu- man, Ciudad Universitaria, 17-H-1959 (J. FE G. Clarke, USNM). Paratype, 1 9, same data as holotype (USNM). Diagnosis.—Superficially, Lobogenesis varnicosa is virtually indistinguishable from most other species in the genus. The genitalia are most like those of L. peruviana among described species. They are easily distinguished by the strongly sclerotized area around the ostium (see Figs. 10, 13). Etymology.—The specific epithet is a manuscript name used by Josef Razowski for this species; its origin is unknown to me. Odonthalitus Razowski 1992 Odonthalitus Razowski 1992: 208; Powell et al. 1995: 145. Type species.—Odonthalitus lacticus Ra- zowski 1992, by monotypy. Redescription.—Head: Antennal cilia ca. 3.0 times width of flagellomere in male, ca. 0.5 times width of flagellomere in female. Labial palpus (segments IT and III combined) ca. 1.5 times horizontal di- ameter of compound eye; segment II weakly upturned, slightly expanded dis- tally by scaling; segment III 0.3—0.4 as long as II, smooth-scaled, exposed. Max- illary palpus rudimentary. Frons with overhanging tuft of scales. Ocelli small or absent. Chaetosema present. Proboscis present, presumably functional. Thorax: Smooth-scaled. Legs unmodified, male without foreleg hairpencil. Forewing: Length 2.3—2.4 times width; length of DC about 0.55 times FW length; width of DC about 0.20 DC length; CuA, originates about 0.60 along length of DC; all veins separate beyond DC; chorda and M-stem absent. No upraised scale tufts; male with- out costal fold. Hindwing: Sc+R and Rs closely approximate; M, and CuA, closely approximate; tuft of hairlike scales along 1A+2A, originating near base of wing. Abdomen: Dorsal pits absent; no modified corethrogyne scaling in female. Male gen- italia: Uncus bifurcate, each arm slender; arms either from a common stalk or wide- ly separated basally. Socius variable, usu- ally narrow, digitate, not fused to gnathos. Gnathos simple, non-dentate, arms nar- row, usually joined distally. Subscaphium and hami absent. Transtilla a simple, slen- der band, frequently reduced or membra- nous. Valva moderately short, somewhat triangular-lanceolate. Pulvinus absent. Vinculum complete. Juxta a sclerotized plate. Aedeagus long, slender, straight or curved; phallobase simple; cornuti absent. Female genitalia: Papillae anales narrow, variably notched mesally in a few species. Apophyses posteriores moderately short, anteriores reduced to a nub in most spe- cies. Sterigma a simple scobinate band. Ductus bursae moderately long, membra- nous. Corpus bursae frail, elongate, pear- VOLUME 102, NUMBER 1 shaped or ovoid; spicules and signum lacking. Accessory bursa frequently pre- sent. Diagnosis.—Adults of Odonthalitus are characterized by a white, cream, or pale tan forewing ground color with a simple pattern that includes a dark, trapezoidal, triangular, or semicircular patch near the middle of the costa and a dark basal patch. In a few spe- cies the pattern is variably obscured by darker overscaling, particularly in females. Males have elongate antennal cilia (ca. 3 times the width of the flagellomere) and lack the characteristic euliine male foreleg hairpencil (Brown 1990). Most species are extremely similar in facies, and examina- tion of the genitalia is required for accurate identification. Superficially, adults of Odonthalitus are indistinguishable from those of Loboge- nesis. Characters that distinguish the two are discussed above in the redescription of Lobogenesis. In addition, a few species of Odonthalitus have a modification of the papillae anales in the female genitalia in which they are variably differentiated into a dorsal and ventral lobe separated by a weak indentation (i.e., viridimontis and paos) or distinct, deep notch (i.e., lacti- cus). Distribution and biology.—QOdonthali- tus is known from montane oak-pine for- est habitat from Durango, Mexico, south to Costa Rica, at 1,000—2,600 m eleva- tion. Adults have been collected at lights. A single individual of O. paos was reared by J. Powell on synthetic diet from eggs deposited by a field-collected female, sug- gesting that the larvae are general-feeders. Two species are known to be sympatric in Costa Rica (at Aquiares, Cartago Prov- ince) and Mexico (at Paradero de Mi Ka, Oaxaca). Remarks.—As with Lobogenesis, most species of Odonthalitus are rare in collec- tions; the 9 species treated below are rep- resented by 53 specimens. Five species are known only from the holotype—3 from males and 2 from females. Approximately 37 62% (n = 33) of the specimens examined represent a single species, O. viridimontis, that appears to be restricted to Costa Rica. The availability of specimens from Costa Rica can be attributed primarily to the ef- forts of parataxonomists associated with INBio, a program that is contributing sig- nificantly to our understanding of the Lep- idoptera fauna of the New World tropics; in addition, the work of Jerry Powell, Dan Janzen, and others have stimulated this ef- fort. The present treatment results in the de- scriptions of 6 new species and the reas- signment of 2 previously described species in Odonthalitus. The new combinations are proposed for 2 species previously consid- ered “‘Unplaced Euliini’ (Powell et al. £995). Odonthalitus bisetanus J. Brown, new species (Figs. 14, 34) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale brown; roughened above, pale yellow. La- bial palpus whitish yellow mesally, brown laterally. Antennal scaling bronze-brown. Thorax: Pale tan. Forewing: Length 7.0 mm (n = 1). Upper side pale tan; basal 0.2 dark brown; dark brown, triangular patch near middle of costa, with vertex of triangle curving outward toward termen; three small irregular spots at costa between patch and apex; apex and termen with irregular brown striae. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side pale cream with pale gray-brown mottling. Under side light gray brown with darker mottling. Genitalia: As in Fig. 14 (drawn from JWB slide 650; n = 1). Uncus bifurcate from base. Gnathos and socius unmodified. Transtilla membra- nous mesally. Valva broadest at base, weak- ly attenuate distally; junction of costal base and transtilla with large hooklike process comprised of 4 strong, weakly curved spines; 2 large setae below hooklike pro- cess along inner edge of valva. Aedeagus moderately slender, nearly j-shaped, with 38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 14-16. Male genitalia of Odonthalitus; valvae spread, aedeagus removed. 14, O. bisetanus. 15, O. improprius. 16, O. conservanus. greatly enlarged membranous cap on phal- Mpio. Yolox, Paradero de Mi Ka, 2,000 m, lobase. 5/14-XI-1980 (E. Welling, UCB). Female. Unknown. Diagnosis.—The group of long curved Type.—Holotype, ¢, Mexico, Oaxaca, spines at the base of the valva in the male VOLUME 102, NUMBER 1 genitalia immediately distinguish Odon- thalitus bisetanus from all other species except O. improprius. These two species can be separated by the following: base of the uncus bifurcations rectangular in O. bisetanus and rounded in O. improprius; tip of the uncus bifurcations sightly flat- tened in O. bisetanus and pointed in O. improprius; patch of long spines from a lobe at the base of the transtilla in O. bis- etanus and from a narrow, attenuate flange in O. improprius; and a pair of large, strong setae just below the basal lobe of the transtilla in O. bisetanus lacking in O. improprius. Etymology.—The specific epithet refers to the two setae at the base of the valva. Odonthalitus improprius J. Brown, new species (Fig. 15) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, whitish; roughened above, whitish. Labial palpus whitish mesally, brown laterally. Antennal scaling pale bronze-brown. Tho- rax: Pale tan. Forewing: Length 5.8 mm (n = 1). Upper side pale tan; basal 0.2 dark brown; dark brown, triangular patch near middle of costa, with vertex of triangle curving outward toward termen; two small triangular spots at costa between patch and apex; terminal region with diffuse brown striae. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side pale cream with pale gray-brown mottling. Under side light gray brown with darker mottling. Genitalia: As in Fig. 15 (drawn from JWB slide 699; n = 1). Uncus bifurcate from base. Gnathos and socius unmodified. Transtilla membra- nous mesally. Valva broadest at base, slightly attenuate distally; junction of costal base and transtilla with flat, claw-shaped flange with 4 strong spines from its venter. Aedeagus moderately slender, nearly j- shaped, with greatly enlarged membranous cap on phallobase. Female. Unknown. 39 Type.—Holotype, 6, Mexico, Oaxaca, Mpio. Yolox, Paradero de Mi Ka, 2,000 m, 5/14-XI-1980 (E. Welling, UCB). Diagnosis.—Odonthalitus improprius is extremely similar to O. bisetanus; the dif- ferences are detailed above in the diagnosis of the latter. Etymology.—The specific epithet is Lat- in for inappropriate or improper. Odonthalitus conservanus J. Brown, new species (Figs. 16, 21) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale tan-brown; roughened above, bronze- brown. Labial palpus pale whitish mesally, brown laterally. Antennal scaling pale bronze-brown. Thorax: Tan brown. Fore- wing: Length 5.0 mm (n = 1). Upper side whitish with irregular, fine, brown reticu- lations throughout; basal 0.2 dark brown; dark brown, triangular patch at middle of costa. Under side uniform dark tan with faint indication of upper side markings. Hindwing: Upper side light gray brown with slightly darker gray brown mottling. Under side light gray-brown with darker mottling. Genitalia: As in Fig. 16 (drawn from JWB slide 606; n = 1). Uncus bifur- cate from base, arising from narrow dorsal nub. Gnathos and socius unmodified. Tran- stilla a broad, weakly sclerotized plate. Val- va broadest at base, attenuate distally, with rounded excavation at costa ca. 0.33 dis- tance from base to apex; inner portion of excavated region bearing a large sclerotized hooklike process, with patch of fine setae from valva just ventrad of hook. Aedeagus moderately slender, nearly j-shaped, with greatly enlarged membranous cap on phal- lobase. Female. FW length 6.5—6.8 mm (x = 6.6; n = 4). Essentially as described for male, except entire FW evenly overscaled with brown, nearly obscuring FW pattern. Gen- italia: As in Fig. 21 (drawn from JWB slide 607; n = 2). Papillae anales unmodi- fied, from bristly base. Papillae anales mod- 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 17-18. poas. erately long, unmodified. Sterigma an irreg- ular, sclerotized band, with a bristly lobe at each postero-lateral margin; mesal portion of sterigma surrounding ostium somewhat diamond-shaped; ostium at anterior vertex of diamond. Ductus bursae membranous, undifferentiated from corpus bursae, slen- der, frail. Types.—Holotype, 3d, Mexico, Jalisco, Male genitalia of Odonthalitus; valvae spread, aedeagus removed. 17, O. viridimontis. 18, O. Sierra de Manantlan, nr. Las Joyas, 1,800 m, 18-VII-1985 (J. Doyen, UCB). Paratypes, 4 2 as follows: MEXICO: Jal- isco: same locality as holotype, 2 2, 16- VII-1985, 1 9%, 18-VII-1985 (J. Doyen, UCB, USNM); 11.6 mi [18.6 km] S of El Chante, 1 2, 3-IV-1987 (N. Bloomfield, SDNHM). Diagnosis.—The male genitalia of Odon- VOLUME 102, NUMBER 1 41 Figs. 19-20. Male genitalia of Odonthalitus; valvae spread. 19, O. regilla. 20, O. lacticus. thalitus conservanus are most similar to those O. bisetanus and O. improprius. They can be distinguished from the latter two by the large hooklike process at the base of the valva, the absence of the patch of hooklike spines at the base of the transtilla, and the broad, shield- shaped transtilla. The female genitalia are most similar to those of O. fuscomaculatus in the presence of a pair of scobinate lobes on the sterigma and the relatively unmodified (1.e., long) apophyses anteriores. Etymology.—The specific epithet refers 42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to the “‘conserved”’ status of the type lo- cality of Las Joyas in the Sierra de Man- antlan, a global biosphere reserve in west- ern Mexico. Odonthalitus orinoma (Walsingham 1914), new combination (Fig. 26) Tortrix orinoma Walsingham 1914: 287. Anopina orinoma: Obraztsov 1962: 27; Powell et al. 1995: 142. Redescription.—Male. Unknown. Female. Head: Frons with sparse, smooth scaling below mid-eye, pale tan; roughened above, cream and brown. Labial palpus pale whitish yellow mesally, tan lat- erally. Antennal sclaing light brown. Tho- rax: Brown. Forewing: Length 7.5 mm (n = 1). Upper side pale cream with nearly uniform pale brown overscaling; basal 0.2 dark brown; triangular brown patch border- ing costa ca. 0.45—0.65 distance from base; costa between latter patch and apex with three small brown triangular spots; terminal region with irregular brown striae. Under side uniform dark tan with faint indication of upper side markings. Hindwing: Upper side pale gray-brown. Under side pale gray brown. Genitalia: As in Fig. 26 (drawn from BMNH slide 66598; n = 1). Papillae anales unmodified. Apophyses posteriores unmodified; apophyses anteriores greatly reduced. Sterigma a sclerotized band, with ill-defined scobinate plates on each side of ostium and a sclertoized, tubelike region mesally; ostium covered by rounded, semi- membranous plate. Ductus bursae long, slender, undifferentiated, gradually widen- ing to corpus bursae. Type.—Holotype, 2, Mexico, Guerrero, Omilteme, 8,000’ [2,580 m], VI-1880 (H. H. Smith, BMNH). Diagnosis.—QOdonthalitus orinoma is most similar to O. fuscomaculatus in facies and female genitalia. The two can be dis- tinguished by features of the female geni- talia—O. fuscomaculatus has conspicuous- ly longer apophyses, an undulate anterior edge of the sterigma, and a shorter ductus bursae (see Figs. 22, 26). Remarks.—The forewing pattern is illus- trated in Walsingham (1914) and the geni- talia in Obraztsov (1962). Obraztsov (1962) transferred “‘T.” orinoma to Anopina on the basis of the similarity in forewing pattern, indicating that “‘Until the male of orinoma is known, the systematic position of this species is somewhat doubtful.’”” The com- bination of forewing length and pattern, and the extremely short apophyses clearly in- dicate that it belongs in Odonthalitus. Odonthalitus fuscomaculatus J. Brown, new species (Fig. 22) Description.—Male. Unknown. Female. Head: Frons with sparse, smooth scaling below mid-eye, pale tan; roughened above, pale tan brown. Labial palpus whitish mesally, tan laterally. Anten- nal scaling light brown. Thorax: Brown and pale tan. Forewing: Length 7.5 mm (n = 1). Upper side cream, with nearly uni- form brown overscaling and irregular brown striae; basal 0.2 dark brown; brown triangular patch bordering costa ca 0.45— 0.65 distance from base. Under side uni- form dark tan with faint indication of upper side markings. Hindwing: Upper side pale gray-brown. Under side pale gray brown. Genitalia: As in Fig. 22 (drawn from USNM slide 89449; n = 1). Papillae anales unmodified from bristly base. Apophyses posteriores and anteriores nearly equal in length. Sterigma with a pair of swollen, bristly lobes dorso-laterad of ostium; osti- um cup-shaped, sclerotized. Ductus bursae slender, undifferentiated from corpus bur- sae. Type.—Holotype, 2, Mexico, Michoa- can, San Lorenzo, Rt. 15, km 206, 19-VII- 1966 (O. Flint & A. Ortiz, USNM). Diagnosis.—Odonthalitus fuscomacula- tus is most similar to O. conservanus in structures of the female genitalia. Odon- thalitus fuscomaculatus can be distinguish by its larger, cup-shaped ostium, more slen- VOLUME 102, NUMBER I Figs. 21-23. Female genitalia of Odonthalitus. 21, O. conservanus. 22, O. fuscomaculatus. 23, O. poas. 43 44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 24-26. Female genitalia of Odonthalitus. 24, O. viridimontis. 25, O. lacticus. 26, O. orinoma. VOLUME 102, NUMBER 1 der apophyses posteriores, and the more mesal position of the bristly lobes of the sterigma. Etymology.—The specific epithet is an adjective referring to the dark overscaling of the forewing. Odonthalitus poas J. Brown, new species (Bigs. 18523; 36) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale tan; roughened above, bronzy tan. Labial palpus pale cream mesally, bronzy brown laterally. Antennal scaling brown. Thorax: Dark brown, whitish tan at mid-dorsum. Forewing: Length 5.0—5.5 mm (x = 5.4; n = 4). Upper side white, with irregular brown transverse striae, particularly dense in distal 0.33; basal 0.2 dark brown; dark brown triangular or rhomboidal patch bor- dering costa ca. 0.45—0.60 from base, fre- quently with small, disjunct spot below lower apex of triangle; costa with a few small brown markings between patch and apex. Under side dark tan with faint indi- cation of upper side markings. Hindwing: Upper side dingy white, with pale gray- brown mottling. Under side light gray brown with darker mottling. Genitalia: As in Fig. 18 (drawn from JWB slide 761; n = 2). Uncus moderately broad at base, bi- furcate in distal 0.75, with fine hairs. Gna- thos complex, with various rounded and an- gulate lateral processes and spines. Tran- stilla mostly membranous with setate basal portions. Valva broadest at base, attenuate apically, with excavation near mid-point of costa leaving a digitate, hairy, lobelike flange at termination of basal 0.5 of costa. Aedeagus slender, nearly evenly curved throughout; phallobase with ovoid membra- nous cap. Female. FW length 6.0 mm (n = 2). Su- perficially as in male, except slightly larger average forewing length and lacking elon- gate antennal cilia. Genitalia: As in Fig. 23 (drawn from JWB slide 762, Costa Rica; n = 2). Papillae anales with shallow notch near middle. Apophyses anteriores absent; 45 posteriores short, slender. Sterigma a sim- ple, uniform, slightly spiculate band, with a broad subrectangular flap mesally covering ostium. Ductus bursae moderately broad, with patch of weak sclerotization, gently broadening into frail corpus bursae. Corpus bursae oblong. Types.—Holotype, 6, Costa Rica, Car- tago Province, Rio Aquiares, nr. Santa Cruz, 9 km NW Turrialba, 1,500 m, 15-V- 1985 (J. Powell, UCB). Paratypes, 3 6, 2 2. COSTA RICA: AI- ajuela Province: NE slope of Volcan Poas, 8 km N Vara Blanca, 1,400—1,450 m, 1 6, 6-VI-1988, 2 3, 19-VI-1988 (J. Brown & J. Powell, UCB), 1 2, 25-VII-1990, JAP 90G20 (S. Meredith & J. Powell, UCB), 1 2, 25-III-1992, JAP 92C50 (J. McCarty & J: Powell, UCB): Diagnosis.—Odonthalitus poas is most similar to O. viridimontis. It can be distin- guished superficially from O. viridimontis by the brown, rather than yellow, transverse striae. The base of the uncus is much more narrow in O. poas and the costal lobe of the valva more pronounced (see Figs. 17—18). Biology.—A confined female (JAP 90G20) produced two eggs. The larvae were reared on artificial diet; one success- fully pupated but did not eclose (J. Powell, personal communication). In general, spe- cies that are capable of completing devel- opment on artificial diet are general-feed- ers—the only clue to potential larval host plants of the group. Etymology.—The specific epithet, a noun in opposition, refers to the collecting locality of all but the holotype on the slope of Mount Poas, Costa Rica. Odonthalitus viridimontis J. Brown, new species (Bigs: 177.245 35) Description.—Male. Head: Frons with sparse, smooth scaling below mid-eye, brown and pale yellow; roughened above, bronzy yellow. Labial palpus pale yellow mesally, reddish brown laterally. Antennal scaling brown. Thorax: Dark brown. Fore- 46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 27-37. Adults of Odonthalitus and Lobogenesis. 27, L. contrasta (3). 28, L. antiqua (d). 29, L. peruviana (2). 30, L. magdalenana (@, Bolivia). 31, L. varnicosa (2). 32, L. penai (3). 33, L. larana (2). 34, O. bisetanus (d). 35, O. viridimontis (3). 36, O. poas (3). 37, L. magdalenana (3, Venezuela). VOLUME 102, NUMBER 1 wing: Length 5.0-5.5 mm (* = 5.2; n = 10). Upper side grayish white with faint yellow transverse striae; basal 0.5 dark brown; dark brown semicircular or trian- gular patch bordering costa ca. 0.45—0.60 from base, frequently with small hooklike marking from lower apex of triangle; ter- minal area with brown overscaling. Under side dark tan with faint indication of upper side markings. Hindwing: Upper side light gray-brown with slightly darker gray-brown mottling. Under side light gray brown with darker mottling. Genitalia: As in Fig. 17 (drawn from JWB slide 609, Costa Rica; n = 5). Uncus extremely broad in basal 0.5, arms widely divergent distally. Soci re- duced to tiny pads. Gnathos arms complex, with a slender, curved basal appendage. Valva broad at base, attenuate apically, with rounded excavation along costa in distal 0.5. Aedeagus extremely long, slender, bent at ca. 0.33 distance from base to tip. Female. FW length 5.0—6.5 mm (x = 5.6; n = 8). Superficially as in male, except slightly larger average forewing length, fre- quently with more dense overscaling, and lacking elongate antennal cilia. Genitalia: As in Fig. 24 (drawn from JWB slide 793, Costa Rica; n = 5). Papillae anales flattened laterally, weakly notched near middle. Apophyses anteriores absent. Sterigma a simple band, dilated at middle to accom- modate ostium. Antrum sclerotized. Ductus bursae frail, slender, undifferentiated from corpus bursae. Types.—Holotype, ¢, Costa Rica, Pun- tarenas Province, Monteverde, 1500 m, 11- VI-1988 (J. Brown & J. Powell, UCB). Paratypes; 47 6, 15"2..\COSTA RICA: Cartago Province: Rio Aquiares, nr. Santa Cruz, 9 air km NW Turmialba, 1,500 m, 2 2, 16-V-1985 (J. Powell & P. Opler, UCB), 1 2, 8-VI-1988, 2 5d, 1 2, 10-VI-1988 J. Brown & J. Powell, UCB); Monumento Nacional Guayabo, A. C. Amistad, 1,100 m, 1 3, VI-1994 (G. Fonseca, INBio). Pun- tarenas Province: Monteverde, 2 6, 1 &, 18/19-V-1985 (J. Doyen, UCB), 2 2, 11- VI-1988, 1 3, 13-VI-1988 (J. Brown & J. 47 Powell, UCB), 1 6, 1 2, 22/24-VII-1990 (S. Meredith & J. Powell; UCB), 1 ¢6, 1 &, 29/31-III-1992, UV & MV lights (S. McCarty & J. Powell, UCB); Estacion La Casona, Res. Biol. Monteverde, 1,520 m, 1; fe, XI-1990 (N. Obando), 1 6,1 2, X-1991 (J. Saborio, INBio), 1 6, IX-1992, 1 3, V- 19933) ded 5° Te S-VIlIE1993," 1, = 1994 (all N. Obando, INBio, USNM); San Luis, Monteverde, A. C. Arenal, 1,000—1,350 m, 2 2, VI-1994 (Z. Fuentes, INBio); Buen Amigo, San Luis, Monteverde, 1,000— 15350m, 16) VI-1994) "1°92 XM-1996.(Z: Fuentes, INBio); 2 km E of Monteverde, 2 3d, 12-VI-1988, blacklight (J. Brown & J. Powell, UCB); Estac. Biol. Las Cruces, Rio Jaba, 6 km SE San Vito, secondary forest, 150mm ,e21ss.,. 220/211-1993 y G- “Powell; UCB). Diagnosis.—The male genitalia of Odon- thalitus viridimontis are distinguished easi- ly from those of its congeners by the ex- tremely broad base of the uncus, the shape of the valvae, and the extremely long, slen- der aedeagus (Fig. 17). Although the shape of the aedeagus is similar to that of O. re- gilla (Fig. 18), the two are not similar in other features of the genitalia. Etymology.—The specific epithet is a noun referring to the type locality of Mon- teverde, Costa Rica. Odonthalitus regilla (Walsingham 1914), new combination (Fig. 19) Tortrix regilla Walsingham 1914: 289. ‘‘Eulia”’ regilla: Powell et al. 1995: 146. Redescription.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale tan-brown; roughened above, bronze- brown. Labial palpus pale whitish yellow mesally, bronze-brown laterally. Antennal scaling pale bronze-brown. Thorax: Tan- brown. Forewing: Length 7.2 mm (n = 1). Upper side pale red-brown; basal 0.2 dark brown, basal patch wider at costa than at dorsum; dark brown, irregularly triangular patch at middle of costa; a brown spot near 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON apex of DC at vertex of costal triangular patch; apex and termen with brown band concolorous with costal patch. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side light gray brown. Under side light gray brown. Genitalia: As in Fig. 19 (photo- graph of BMNH slide 5794; n = 1). Uncus with a posterior nub, behind (anterad) which are a pair of extremely slender, elon- gate arms ( = bifurcations of uncus) from dorsum of tegumen. Gnathos swollen dis- tally. Socii short, broad. Transtilla membra- nous, ill-defined. Valva slender, nearly uni- form in width, with weak notch along ven- ter ca. two-thirds distance from base to apex; apex with strong, free, curved hook from weakly rounded base. Aedeagus long, straight, extremely slender. Female. Unknown. Type.—Holotype, ¢, Guatemala, Retal- huleu, Las Mercedes, 3,000’ [970 m], X/XI- 1880, (G. C. Champion, BMNH). Diagnosis.—The male genitalia of Odon- thalitus regilla are moderately divergent from other species in the genus: the nearly parallel-sided valva and the clawlike pro- cess from the apex are unlike any other spe- cies. The long, slender aedeagus is reminis- cent of that of O. viridimontis, and the basal origin of the bifurcation of the uncus is sim- ilar to O. lacticus. Remarks.—The adult is illustrated in Walsingham (1914). The forewing pattern, elongate antennal cilia, bifurcate uncus, and long, slender aedeagus indicate that it be- longs in Odonthalitus. Odonthalitus lacticus Razowski 1992 (Figs. 20, 25) Odonthalitus lacticus Razowski 1992: 208; Powell et al. 1995: 145. Redescription.—Male. Head: Frons with sparse, smooth scaling below mid-eye, pale tan; roughened above, cream. Labial palpus whitish mesally, tan laterally. Antennal scaling light brown. Thorax: Brown. Fore- wing: Length 6.5 mm (n = 1). Upper side cream; basal 0.2 dark brown; triangular brown patch bordering costa ca. 0.45—0.65 distance from base; costa between latter patch and apex with three small brown tri- angular spots; terminal region with irregular brown band. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side pale gray brown. Under side pale gray brown. Genitalia: As in Fig. 20 (drawn from UCB slide 3392; n = 1). Tegumen short, rounded. Uncus com- posed of two distinct, widely separated rods. Socius short. Gnathos long, slender, with accessory process near middle, arms separate distally. Valva relatively short, at- tenuate at apex, venter concave in basal 0.5; costa undifferentiated; sacculus weakly sclerotized. Transtilla a broad rectangular plate. Aedeagus large, curved, broadened apically. Female. FW length 6.0—7.0 mm (x = 6.3; n = 3). Essentially as described for male, except ground color in some specimens evenly dusted with pale brown, reducing definition of forewing markings. Genitalia: As in Fig. 25 (drawn from UCB slide 3312 and JWB slide 603; n = 2). Papillae anales with pronounced notch near middle, result- ing in bilobed appearance. Apophyses an- teriores represented by a short thorn; apophyses posteriores comparatively long. Sterigma a simple, broad, sclerotized band, with a pair of lateral incisions. Ductus bur- sae broad, moderately short, with fine lines of sclerotization. Corpus bursae short, rounded. Accessory bursa from narrow ductus arising near middle of ductus bursae. Types.—Holotype, 2, Mexico, Durango, 4 miles [6.4 km] west of El Palmito, 20- VII-1964, flight trap (J. Powell, UCB). Paratype, | d, same data as holotype. Additional specimens examined.—MEX- ICO: Sinaloa: 15 mi [24 km] west of El Palmito, black and white lights, 1 2, 18- VII-1964 (J. Chemsak & J. Powell, UCB); 8 mi [12.8 km] west of El Palmito, 6,000’, Io, 122X21975:9 at Hight) GycPowell; 3: Chemsak, T. Eichlin & T. Friedlander, UCB). VOLUME 102, NUMBER 1 Diagnosis.—The male genitalia of Odon- thalitus lacticus are fairly divergent from other species in the genus. They can be dis- tinguished by the widely concave distal 0.5 of the venter of the valva, the widely sep- arate bases of the bifurcate uncus, and the relatively robust aedeagus (Fig. 20). The notch in the papillae anales of the female is similar to that of O. viridimontis and O. paos, but is much deeper. ACKNOWLEDGMENTS I thank the following for allowing me to examine material in their care: Jerry Powell (UCB), Eugenie Phillips (INBio), Kevin Tuck (BMNH), and David Faulkner (SDNHM). I thank the following for re- viewing the manuscript: Jerry Powell, Uni- versity of California, Berkeley, California; Richard Brown, Mississippi State Univer- sity, Mississippi State, Mississippi; William Miller, University of Minnesota, St. Paul, Minnesota; and E Christian Thompson and David Smith, Systematic Entomology Lab- oratory, USDA, National Museum of Nat- ural History, Washington, D.C. I thank J. Powell for providing me with the opportu- nity to conduct field work in Costa Rica and for comments on rearing attempts. The fig- ures were skillfully rendered by Susan Escher (Figs. 2—5, 9-18, 20, 24), Front Royal, Virginia, and Linda Lawrence (Figs. 6-8, 21-23), USDA, Systematic Entomol- ogy Laboratory, Washington, D.C. 49 LITERATURE CITED Brown, J. W. 1990. Taxonomic distribution and phy- logenetic significance of the male foreleg hairpen- cil in the Tortricinae (Lepidoptera: Tortricidae). Entomological News 101: 109-116. Brown, J. W. and J. A. Powell. 1991. Systematics of the Chrysoxena group of genera (Lepidoptera: Tortricidae: Euliini). University of California Pub- lications in Entomology 111. 87 pp. 2000. Systematics of Anopina Obraztsov (Lepidoptera: Tortricidae). University of Califor- nia Publications in Entomology. In press. Horak, M. 1984. Assessment of taxonomically signif- icant structures in Tortricinae (Lep., Tortricidae). Mitteilugen der Schweizerischem Entomologisch- en Gesellschaft 57: 3—64. Lipscomb, D. 1994. Cladistic Analysis Using Hen- nig86, version 1.5. George Washington Universi- ty, 122 pp. Obraztsov, N. 1962. Anopina, a new genus in the Cne- phasiini from the New World (Lepidoptera: Tor- tricidae). American Museum Novitates 2082: 1— 39. Powell, J. A., J. Razowski, and J. W. Brown. 1995. Tortricidae: Tortricinae, pp. 138—150. /n Heppner, J. B., ed., Atlas of Neotropical Lepidoptera, Checklist: Part 2, Hyblaeoidea-Pyraloidea-Tortri- coidea. Association for Tropical Lepidoptera. Sci- entific Publishers, Gainesville, FL. Razowski, J. 1984. A list of the known Neotropical Tortricini moths (Lepidoptera, Tortricidae) with description of the first South American species. Revista Brasiliana Entomologica 28: 203-206. . 1992 [1990]. On some peculiar Neotropical tortricine genera (Lepidoptera: Tortricidae). SHI- LAP Revista de Lepidopterologia 18: 209-215. . 1993. Revision of Apotoforma Busck, 1934 (Lepidoptera: Tortricidae), with descriptions of four other Tortricini. Acta Zoologicae Cracovien- sia 36: 183-197. Walsingham, Lord T. de Grey. 1914. Biologia Centrali- Americana. Insecta, Lepidoptera Heterocera, Vol. 6. British Museum (Natural History), London. 482 PP. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 50-57 MILESIINE FLOWER FLIES (DIPTERA: SYRPHIDAE) IN A CENTRAL APPALACHIAN BROADLEAF FOREST: ABUNDANCES, FLIGHT PERIODS, AND DIFLUBENZURON Mary R. CICERO AND EDWARD M. BARROWS Laboratory of Entomology and Animal Behavior, Reiss Building Suite 406, Department of Biology, Georgetown University, Box 571229, Washington, DC 20057-1229, U.S.A (e-mail: barrowse @ gunet.georgetown.edu) Abstract.—We used 20 Malaise traps to collect adult milesiine syrphids in four water- sheds in a central Appalachian broadleaf forest during early April through late September in 1991 through 1993. The traps captured 31 species in 12 genera. As a group, these flies flew throughout our annual sampling periods, their sample sizes varied with species and year, and their abundances peaked in late May or mid-June, depending on the year. For all species whose sample size was 210 individuals, male-female sex ratios varied from 0.1 through 3.0. We performed a split-plot analysis of flies captured from mid-May through early August, a period in each of the 3 years when traps captured at least one fly during a sampling period. This analysis did not find an effect of trap site or diflubenzuron on fly population sizes based on Malaise sampling, because these effects did not exist or our sample size was too small to detect them. Key Words: The broad aims of our study are to learn more about the biology of syrphid flies in the tribe Milesiini and determine whether aerially-applied diflubenzuron affects pop- ulation sizes of these beneficial insects in a temporate broadleaf forest. Diflubenzuron (Dimilin®, Uniroyal) is an insect growth regulator used to control forest and other pest arthropods. We report which milesiine species are present in this forest and their flight periods, seasonal abundances, and sex ratios, based on sampling with Malaise traps. Further, we used a split-plot analysis to test the hypotheses that diflubenzuron and trap site affect the number of individ- uals captured. Milesiine syrphids are beneficial insects in forests and other habitats, where they may be Batesian mimics of some stinging hymenopterans, honeydew consumers, food Appalachian forest, diflubenzuron, Malaise traps, phenology, Syrphidae for other organisms, nectarivores, polleni- vores, pollinators, and scavengers (Gilbert 1981, 1986; Maier 1982; Vockeroth and Thompson 1987; Waldbauer 1988; Owen 1991). We studied this syrphid taxon be- cause its members have diverse habits and are taxonomically well known and easily identified. Syrphids and bees are the major pollinators in many habitats. Federal and state agencies have treated thousands of hectares of forests with diflu- benzuron to control gypsy moths, Lyman- tria dispar (L.), in the eastern United States (USDA Forest Service 1994). This pesti- cide also reduces the numbers of nontarget, beneficial forest insects including honey bees (Egger 1977), macrolepidopterans and nonlepidopteran mandibulate herbivores (Martinat et al. 1988); macrolepidopterans (Sample et al. 1993; Butler ect ale 1997); VOLUME 102, NUMBER 1 stoneflies (Griffith et al. 1996), and yellow- jackets (Barrows et al. 1994). However, there are no published reports regarding the possible effects of aerially-applied difluben- zuron on forest syrphids. MATERIALS AND METHODS We studied syrphids in the 1,902-ha Fer- now Experimental Forest in Tucker County, West Virginia, over a 3-year period (1991— 1993). Detailed accounts of our materials and methods are published in Barrows et al. (1994) and Barrows (1995). The Fernow Experimental Forest is from 533 to 1,112 m in elevation and has slopes up to 60%, a mean of 145 frost-free days, a mean annual precipitation of 147 cm, and mean temperature of 9°C (Anonymous 1987). Acer rubrum L., Betula lenta L., Fa- gus grandifolia Ehrhart, Liriodendron tuli- pifera L., Prunus serotina Ehrhart, and Quercus spp. are common trees in this for- est: We used watersheds 1, 4, 7, and 13 as study areas. Watersheds 1, 7, and 13 are conterminous, and watershed 4 is about 200 m from watershed 7 (Adams et al. 1994, map). Watersheds | and 13, the difluben- zuron-treated plots (test plots) totaled 42 ha, and watersheds 4 and 7, the control plots totaled 63 ha. On 16 May 1992, the USDA Forest Service aerially applied diflubenzu- ron (Dimilin® 4L) with a helicopter at 35.1 g (AI) ha"! to watersheds 1 and 13. During April through September, all 3 study years were drier than a 4l-year av- erage of 76 cm for the Fernow Experimen- tal Forest, 1991 and 1993 were warmer than average (15.9°C), and 1992 was cooler than average. During these months, mean tem- perature was 17.6, 14.9, and 17.1°C, and the mean precipitation was 56.9, 59.9 and 64.3 cm, in 1991, 1992, and 1993, respectively. We captured flies from 29 April through 27 September 1991-1993 using five Townes-style Malaise traps (Townes 1972; manufactured by Golden Owl Publishers, Lexington Park, MD) placed 20—35 m apart on the ground in a transect in each of the 51 watersheds as illustrated in Barrows et al. (1994). Each transect ran across a valley and approximately perpendicularly to a sec- ond-order stream. From the vantage point of looking upstream, two traps were on the left side and two were on the right side of a central trap. Valley left sides had north- erly exposures, and right sides had south- erly exposures. The central trap of each transect was from 0 to 6 m from the edge of a stream. We placed all traps on the for- est floor, not on logging roads or in other artificial openings. A trap was made of 1-mm-mesh, nylon gauze, a supporting frame, and a collecting head. A 0.95-liter jar, which was part of the head and contained 95% ethanol, collected the flies as well as thousands of species of other arthropods. Each trap was 1.2 m wide, 1.7 m long, 1.0 m high at its back, and 2.0 m high at its front (head end). Trap roofs were made of white gauze and baffles and sides were made of black gauze. We emp- tied our 20 traps every 10 d on the same Julian day in April through September in 3 yr, totaling 895 samples. Falling tree limbs, mice, bears, and pos- sibly other animals occasionally damaged our traps; we repaired, or replaced, them as soon as we found damage. Five samples were incomplete because of trap damage and could not be used in our statistical an- alyses. We identified syrphids to genus us- ing keys in Vockeroth and Thompson (1987) and to species with the reference collection at the National Museum of Nat- ural History, Smithsonian Institution, and deposited a voucher collection of these flies in that Museum. For species with large enough sample sizes for analysis, we used x’ goodness of fit tests to determine wheth- er they had sex ratios that significantly dif- fered from male-female sex ratios of 1:1, 1: 2, and 2:1. To look for a possible difference in the number of syrphid individuals among years, we used analysis of variance. Finally, we used a split-plot analysis to look for possible diflubenzuron and trap-site effects on the number of captured flies (PROC oy PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON _— JE O = total = 661 1992 total = 349 total = 732 O WY) LLJ < Tp Figure 1 LL = = LL O O = 110 140 170 200 230 260 110 140 170 200 230 260 110 140 170 200 230 260 Apr May Jun Jul Aug Sep Apr May Jun Jul Aug Sep Apr May Jun Jul Aug Sep JULIAN DAY Fig. 1. Numbers of milesiines captured in each 10-day sample in the four watersheds, Fernow Experimental Forest, 1991-1993. MIXED, SAS Institute 1992). The treat- ments were pesticide (with the two levels application and nonapplication) and trap site (five levels or locations). We calculated the grand mean of the trap means of flies for eight sampling periods (mid-May through early August) per watershed per year. This was a period when most milesi- ines flew and there was at least 1 fly per sampling period in each of the 3 sampling years. In 1991, the preapplication year, mi- lesiine populations were higher in the ap- plication watersheds than in the nonappli- cation watersheds, possibly because the for- mer were better syrphid habitats. To adjust for this difference, we analyzed the differ- ences between the 1991 and 1992 and be- tween the 1991 and 1993 grand means, test- ing the null hypothesis that the difference between the application-watershed means of fly number equals the difference between the nonapplication-watershed means of fly number. To look for a possible trap-site ef- fect, we tested the null hypothesis that the mean numbers of flies from the five trap sites are equal. RESULTS AND DISCUSSION Species, sex ratios, flight periods, and abundances.—Traps collected a total of 1,742 milesiines belonging to 31 species in 12 genera (Fig. 1, Table 1—2). The Fernow Experimental Forest is within the previous- ly known ranges of all of these species; however, this is the first published species list of Milesiini from a central Appalachian broadleaf forest. By the ends of 1991, 1992, and 1993, our traps obtained 84%, 94%, and 100% of the 31 milesiine species, respectively. Some species were rare in samples, and not all species were in samples every year. The six most common species comprised 76% of the 1,742 collected specimens. We were unable to run our traps for more than 3 years, but Owen’s (1991) study sug- gests that our traps may have obtained the majority of the milesiine species in our study site during our three-season study pe- riod. She caught 43,749 individuals in 91 species and 41 genera, by using one Townes-style Malaise trap (Townes 1972) in her British garden for 15 successive years. The trap obtained the vast majority of its total number of syrphid species by the end of its fifth year of operation, rarely caught some species, caught markedly varying numbers of some species from year to year, and obtained a previously uncaught species during most years from the 5th through 15th years of operation. Bankows- ka (1980) found 25 species in seven mile- VOLUME 102, NUMBER 1 Table 1. Abundances of milesiine syrphids and percentages of males collected in Malaise traps from late April through late September, in all four water- sheds in the Fernow Experimental Forest, West Vir- ginia, 1991-1993. Number Species Captured % Males* Blera analis (Macquart) 30 47° B. badia (Walker) 263 65° B. nigra Williston | 100 B. pictipes (Bigot) IS 53° B. umbratilis (Williston) l 100 Brachypalpus oarus Walker 102 47° Chalcosyrphus inarmatus (Hunter) 20 55? C. libo (Walker) 17 298 C. nemorum (F.) 6 7, C. plesius Curran 2 100 C. vecors Osten Sachen 3 33 Criorhina nigriventris Walton 9 22 C. verbosa Walker 36 Le Lejota aerea (Loew) 9 56 Prerallastes thoracicus Loew 123 Sk: Somula decora Macquart 25 24 Specomyia vittata Wiedemann 30 23 Spilomyia alcimus (Walker) 4 100 S. fusca Loew 9 22 Temnostoma alternans Loew 111 She T. balyras Walker 573 522 T. barberi Curran 21 10 T. trifasciatum Robertson | 27 T. venustum Williston 8 25 T. vespiforme (L.) 167 69: Teuchocnemis lituratus (Loew) 66 50° Xylota 78-1 Thompson 4 25 X. 78-3 Thompson 17 294 X. bicolor Loew DZD, 36! X. nebulosa Johnson | 100 X. quadrimaculata Loew 36 17 Total 1,742 “The proportion of males for all species combined is 47%. > This sex ratio is not different than 1.0 or 50% males (df = 1, P < 0.05, x? goodness of fit test). © This sex ratio is not different than 2.0 or 66% males (df = 1, P < 0.05, x* goodness of fit test). 4 This sex ratio is not different than 0.5 or 33% males (df = 1, P < 0.05, x? goodness of fit test). sline genera in Polish broadleaf forests, compared to the 31 species in 12 genera reported in our study (Table 1). In our study, male-female milesiine sex ratios varied from 0.1 through 3.0. Four species had male-female sex ratios that were not different from 1:2; seven species, Bye) 1:1; and two species 2:1 (Table 1). In all species in all 3 years, 47% of the syrphids were males which is not different than a 1: 1 sex ratio (50% males) (df = 1, x? = 2.431, P = 0.119). Deviations from a 1:1 sex ratio in certain species may be due to factors including having an adult sex ratio that is actually different from 1:1 and one sex’s being more likely to be trapped than the other. Milesiines flew throughout most of our monitoring period; however, the traps col- lected the vast majority (95%) of them from early May through late July (Fig. 1, Table 2). Traps acquired individual milesiine spe- cies for an mean of 59.4 + 5.6 SEM days (range 10—130 days, 31 species), obtaining all species except for Chalcosyrphus ne- morum (FE.) and Pterallastes thoracicus Loew, for 100, or fewer, days. This mean and ranges are approximations because our sampling intervals were 10 days. Of the species with sample sizes of 10 or more individuals, three species peaked in abundance in early May; three in mid-May; one in late May; one in early June; eight in mid-June; and one in late June (Table 2). The main annual peaks were in late May 1991 and mid-June in 1992 and 1993 (Fig. 1). The troughs between the two large peaks in 1991 and 1993 coincide with rainy periods. Visnyovszky (1988) also found rain-related drops in Malaise-trap sample sizes in these flies in Hungarian apple or- chards. Among flies in the traps, Temnostoma balyras Walker was the most common, fol- lowed by Blera badia (Walker), T. vespi- forme L., Pterallastes thoracicus, and T. al- ternans Loew (Table 1). The only published study of North American Milesiine flight periods is based on the faunas of Piatt and Mason Counties in central Illinois, both ap- proximately 40° north, and Emmet County in northern lower Michigan, approximately 45° north (Waldbauer 1988). This study in- volved high-fidelity mimics of stinging hy- menopterans which were sampled by hand netting. Our study site (approximately 39° 54 Mablen2: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Abundances of milesiine syrphids and percentages of males collected in Malaise traps from late April through late September, in all four watersheds, Fernow Experimental Forest, West Virginia, 1991-1993 Month and Sampling Period? April May June July August September Species il La ee es eee ReGen ee oe SOL aoe Tinion sian Bren Blera analis 0) 0) | 3 4 20 1 0) 1 Oo 0 O= © @ © @ B. badia 2 2 5 35 78 93 Sy) IS} l 2 0 O OY © @ O B. nigra 0) l 0) 0) 0) 0) 0) oO © O © © O © O B. pictipes O 0) 0) Th 1 4 1 oO © © © OO © O B. umbratilis 10) 0) 0) 0) 0) i 0) oO © © OO @ © O © O Brachypalpus oarus 12 59 17 11 0) 0) 1 oO OF @ O O- OD © O O Chalcosyrphus inarmatus 0) l 8 =) 4 1 oO © O O OO O O C C. libo l 4 4 4 0) 2 l l Oo OO © OF O&O © © C. nemorum O 0) l 1 0) ] | l Yo O © OL. O-O © C. plesius 0) 0) 0) | 0) 1 0) oO O © O © O @ O O C. vecors O 0) 0) 0) 0) 2 | oO Oo O O GO O O © © Criorhina nigriventris D ] 4 2 0) 0) 0) oO © © @ OO OY WO O C. verbosa 6 16 3) 0) 0) 0) 0) oO OO © GO © © O © Lejota aerea 0) 6 3} 0) O 0) 10) Oo Oo © & © O OF OD O Pterallastes thoracicus® 6) 0) l 9 NS) 41 sy alse ay a Del aril 0) Somula decora‘ 0 0) 0) 6 9 5 4 1 Oo OY O OG © © @ Specomyia vittata‘ 6 0) 7 16 2 1 0) tO @ © O O @ © @ Spilomyia alcimus* 0) 0) 0) 0) O 0) | oy Oo 2 OO OO O O @ S. fusca‘ 0) 0) 0) 0) 0) 0) 0) ye Or Be Oe O Temnostoma alternans® 0) 0) 6 29 13 60 16 3} 3 Oe th Oe OY. @ Temnostoma balyras* 2 17 28) 186" 136) 154" 7 437 192 lL © OO O © © © T. barberis 0 0) 1 5 6 6 2 YY 0 CO O @ © @ © O T. trifasciatum’ 0) 0) 0) 0) 5 4 | l YO OO OO O GO © @© O T. venustums 0) 0) l l 1 | 1 | x YO © © © O O @ T. vespiforme® 0) 0) 9 36 19 82 14 7 © © OO O OD @ ©@ Teuchocnemis lituratus 0) 3 25 21 6 6 3 OO 2 0) © O Oo O © @ Xylota bicolor 0) 0) 0) 0) 3 6 9 | BO O 1 @ ©. @ X. nebulosa 0) O 6) ] 0) 0 0 OF Os 0) OO -O) Or OF @ X. quadrimaculata 0) 0) 2 4 8 9 6 4 l fi Oa © © C X. n. sp. | 0) O 0) 0) 0) 1 2 0) l oO OY O O © NG id Coy 2 0) 4 3 4 2 3 0) 0) l OO 0 O O @ © Totals 31 NS 129 340) 31255507 SiG9 a7 OM SO A Gyr “Sample peaks in species with =10 specimens are in bold face. > | = April: Julian days 110-120; May: 2 = 230; 13° = 120-130; 3 = 130-140; 4 = 140-150; June: 5 = 150-160; 6 = 160-170; 7 = 170-180; July: 8 = 180-190; 9 = 190-200; 10 = 200-210; August: 11 = 230-240; September: 14 = 240-250; 15 = 250-260; 16 = 260-270. 210-220; 12 = 220- © High-fidelity mimics of stinging hymenopterans (Waldbauer 1988). north) shares three milesiine species with each of the Illinois sites and seven species with the Michigan site. Approximate mile- siine flight periods in West Virginia are similar to those in the Illinois sites. Their flight periods in West Virginia were earlier and longer than those in Michigan. The mean numbers of milesiines cap- tured in nonapplication watersheds per trap per sampling period was 2.7 + 0.7, 1.7 + 0.7, and 3.4 + 0.7 SEM in 1991, 1992, and 1993, respectively, showing a difference in number of individuals among years (AN- OVA, df = 10, F = 5.94; P = 0.0199). Malaise traps are an excellent means for monitoring population sizes of insect taxa that are readily captured by these traps. However, they probably never collect target taxa in the exact proportions that they occur in their communities (Barrows 1986; Dar- VOLUME 102, NUMBER 1 Nn Nn 150 e test plots O control plots 120 Figure 2 110 140 170 200 230 260 110 140 170 200 230 260 110 140 170 200 230 260 Apr May Jun Jul Aug Sep Apr May Jun Jul Aug Sep Apr May Jun Jul Aug Sep JULIAN DAY Fig. 2. Numbers of milesiines captured in each 10-day sampling period in application and nonapplication watersheds, Fernow Experimental Forest. The arrow indicates the time of pesticide spraying on application plots. NO. OF MILESIINES CAUGHT ling and Packer 1988; Archer 1990). Visual Trap-site and diflubenzuron.—Our split- counts of some syrphid species suggested plot analysis detected no diflubenzuron, or that a Malaise trap did not collect them in _ trap-site, effect on milesiines (P > 0.05, for the proportions that they occurred near the both factors) (Fig. 2, Table 3). However, trap (Owen 1991). this finding does not indicate that aerially- Table 3. Split-plot analysis of possible diflubenzuron and trap-site effects on numbers of milesiine syrphids. Treatments Year Mean + | SEM? Pe No pesticide (1 and 13) 199] 42 = 0.90 not applicable No pesticide (4 and 7) 2.7 220.53 Pesticide (1 and 13) 1991 minus 1992 2.2 + 0.64a 0.2016 No pesticide (4 and 7) 0.9 + 0.64a Pesticide (1 and 13) 1991 minus 1993 —0.3 + 0.98a 0.4642 No pesticide (4 and 7) 0.7 + 0.98a Trap | 199] 2 6ie= (O:8i1a 02297 Trap 2 19+ 0.8la Trap 3 2.0 = '0:8ia Trap 4 3.1 == 10:8ila Trap 5 3.4 + 0.8la Trap | 1992 1.8 = 0.55a 0.3438 Trap 2 1.3, '0.55a Trap 3 IpOy==O55a Trap 4 IESe=="0i55a Trap 5 2.9 = 0:55a Trap | 1993 4.6. cb lda 0.8302 Trap 2 Sos) Labla Trap 3 302 Telia Trap 4 B32 ee ela Trap 5 39° Talla « This mean is the average number of syphids per 10-day sample per year per watershed. Within a year, means followed by the same letter are not different from each other (P = 0.05, least-squares-means test). These means are based on two control, or test, plots per year, or four traps per subplot per year. ® The null hypothesis is that the means within groups (pesticide treatment or trap-site treatment) are equal. 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON applied diflubenzuron would never decrease milesiine populations in forests. We might not have detected a possible diflubenzuron effect because our sample size was small, milesiines might have entered our test wa- tersheds from untreated ones and masked a pesticide effect, or both. No apparent effect of trap site on milesiine sample size sug- gests that these flies were relatively evenly distributed in our sample area. In conclusion, we report that 31 milesiine species occurred in a central Appalachian broadleaf forest and they were of markedly different abundances and sex ratios in Mal- aise-trap samples. Although, our study found no evidence that diflubenzuron af- fected milesiine population sizes, a more comprehensive study might find such an ef- fect. This pesticide remains on _ treated leaves for many months on plants and in leaf litter (Wimmer 1995), and larval mi- lesiines are scavengers which could be poi- soned by them. If forest managers continue to use diflubenzuron to control gypsy moths and other forest pests, more studies of this pesticide and syrphids are warranted. ACKNOWLEDGMENTS We are pleased to thank many people who helped with our study: Mary Beth Ad- ams, Pamela J. Edwards, and Fredericka Wood (USDA Forest Service, Parsons, WV); Zorayda C. Alviar, Holi D. Ieng, Jo- seph H. Neale, and James V. Tedesco (Georgetown University); Joseph E. Bar- rows, Julie N. Barrows, Melissa L. Bar- rows, and Robert E. Harrigan (Bethesda, MD); Donald A. M. Mackay (Pleasantville, NY), David R. Smith (USDA, Washington, DC); Amy H. Onken and Richard C. Rear- don (USDA Forest Service, Morgantown, WV); Mary J. Camp and William E. Potts (Statistical Consulting Services, USDA, Beltsville, MD); Janet D. Rupp (Golden Owl Publishers, Lexington Park, MD); and Deborah J. Smith (Falls Church, VA). Lin- da Butler (West Virginia University, Mor- gantown, WV), Chris T. Maier (Connecticut Agriculture Experiment Station, New Ha- ven, CT), and an anonymous reviewer made insightful comments that improved this paper. EK Christian Thompson (System- atic Entomology Laboratory, USDA, Wash- ington, DC) kindly gave us invaluable help with syrphid identification and other mat- ters. This study was funded by Georgetown University and the Appalachian Integrated Pest Management Gypsy Moth Project of the Northeastern Area, State and Private Forestry, USDA Forest Service. LITERATURE CITED Adams, M. B., J. N. Kochenderfer, E Wood, T. R. An- gradi, and P. Edwards. 1994. Forty years of hy- drometeorological data from the Fernow Experi- mental Forest, West Virginia. USDA Forest Ser- vice, Northeastern Forest Experiment Station. General Technical Report NE-184. 24 pp. Anonymous. 1987. Forest Research: Fernow Experi- mental Forest. U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station NE-INF-75-87. 12 pp. Archer, M. E. 1990. The solitary aculeate wasps and bees (Hymenoptera: Aculeata) of an English sub- urban garden. Entomologist’s Gazettee 41: 129— 142. Bankowska, R. 1980. Fly communities of the family Syrphidae in natural and anthropogenic habitats of Poland. Memorabilia Zoologica 33: 3—93. Barrows, E. M. 1986. A hornet, paper wasps, and yel- lowjackets (Hymenoptera: Vespidae) in suburban habitats of the Washington, D.C., Area. Proceed- ings of the Entomological Society of Washington 88: 237-243. . 1995. Chapter 8. Pollinating insects—Native species, pp. 66—76. Jn Reardon, R. C., ed., 1995, Effects of Diflubenzuron on Non-target Organ- isms in Broadleaf Forested Watersheds in the Northeast. USDA Forest Service FHM-NC-0595. 174 pp. Barrows, E. M., S. S. Wolf, & D. M. Lynch. 1994. Diflubenzuron effect on yellowjacket (Hymenop- tera: Vespidae) worker numbers in a central Ap- palachian broadleaf forest. Journal of Economic Entomology 87: 1488-1493. Butler, L., G. A. Chrislip, V. A. Kondo, and E. C. Townsend. 1997. Effect of diflubenzuron on non- target canopy arthropods in closed, deciduous wa- tersheds in a central Appalachian forest. Journal of Economic Entomology 90: 784—794. Darling, D. C. and L. Packer. 1988. Effectiveness of Malaise traps in collecting Hymenoptera: The in- fluence of trap design, mesh size, and location. Canadian Entomologist 120: 787-796. Egger, A. 1977. Die Nebenwirkungen von DIMILIN VOLUME 102, NUMBER 1 auf Bienen—eine Untersuchung anlasslich einer Flugzeugapplikation im Forst. Centralblatt fiir das Gesamte Forstwesen 94: 65-72. 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Canadian Entomologist 125: 173-179. SAS Institute. 1992. SAS/STAT software: Changes and enhancements, Release 6.07. Technical Report P-229. SAS Institute, Cary, NC, 620 pp. Townes, H. 1972. A light-weight Malaise trap. Ento- mological News 83: 239-247. United States Department of Agriculture Forest Ser- vice. 1994. Gypsy moth news 36. Visnyovsky, E. 1988. Phenological study of the syr- phid flies caught by a Malaise trap, pp. 123-127. In Niemcezyk, E. and A. FE G. Dixon, eds., Ecology and Effectiveness of Aphidophaga. SPB Academ- ic Publishing, The Hague, The Netherlands. Vockeroth, J. R. and F C. Thompson. 1987. Syrphidae, pp. 713-743. In McAlpine, J. F, ed., Manual of Nearctic Diptera, Vol. 2. Biosystematics Research Centre, Ottawa, Ontario. Waldbauer, G. P. 1988. Asynchrony between Batesian mimics and their models. The American Naturalist 131: S$103-S121. Wimmer, M. J. 1995. Chapter 16. Terrestrial Environ- ment, pages 23-30. Jn Reardon, R. C., ed., 1995, Effects of Diflubenzuron on Non-target Organ- isms in Broadleaf Forested Watersheds in the Northeast. USDA Forest Service FHM-NC-0595. 174 pp. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 58-61 A NEW SPECIES OF CIRROSPILUS WESTWOOD (HYMENOPTERA: EULOPHIDAE) FROM THE SOUTHWESTERN UNITED STATES AND MEXICO MICHAEL W. GATES Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: gates @citrus.ucr.edu) Abstract.—Cirrospilus coachellae, n. sp. (Hymenoptera: Eulophidae), is described from the southwestern United States and Mexico. Comparisons are made with described species of Cirrospilus and differences between Cirrospilus and the closely related Za- grammosoma are discussed. Cirrospilus coachellae is the primary parasitoid providing control of the citrus peelminer, Marmara sp. (Lepidoptera: Gracillariidae), a cyclical pest of citrus in the southwestern United States. Key Words: Eulophidae in the genus Cirrospilus Westwood are cosmopolitan in distribution and parasitize insects with cryptic feeding habits. Cirrospilus parasitizes leaf-miners, leaf-rollers and galling insects in the orders Lepidoptera, Coleoptera, Diptera and Hy- menoptera (Schauff et al. 1997, Boutek 1988). Species of Cirrospilus typically are larval ectoparasitoids, but also have been documented to utilize primary ichneumo- noid parasitoids attacking concealed insects and to directly attack several life stages of the primary host (Boucek and Askew 1968). Currently, there are over 300 species of Cirrospilus described worldwide, of which the majority occur in the Holarctic region. Cirrospilus belongs to the subfamily Eu- lophinae, as characterized by the submar- ginal vein smoothly joining the parastigma, a postmarginal vein often longer than the stigmal vein, and a minimum of three setae on the dorsal surface of the submarginal vein (Schauff et al. 1997). Cirrospilus is characterized by having the funicle two- segmented, notauli complete, and the post- Hymenoptera, Eulophidae, Cirrospilus, taxonomy, Chalcidoidea marginal vein shorter than or equal in length to the stigmal vein (Schauff et al. 1997). Gordh (1978) recognized another closely related eulophine genus, Zagram- mosoma Ashmead, as distinct from Cirros- pilus on the basis of two characters: head with the vertex distinctly vaulted so as to extend above the compound eye in Zagram- mosoma, and the propodeum with a well developed median carina in Cirrospilus but only a weak carina in Zagrammosoma. LaSalle (1989) further separated these gen- era by noting in Zagrammosoma the notau- lus curves to meet the axilla anteriorly, thus not approaching the scutoscutellar suture, while in Cirrospilus the notaulus extends straight to the scutoscutellar suture. LaSalle discounted the propodeal carina character because of its inconsistency across taxa. Yefremova (1996) cited additional char- acters which serve to distinguish Zagram- mosoma from Cirrospilus. In Zagrammo- soma, the axilla is situated anterior to the posterior margin of the mesoscutum and typically is elongated. Cirrospilus has the axilla located posterior to the posterior mar- VOLUME 102, NUMBER 1 gin of the mesoscutum, and the axilla is more triangular than the axilla of Zagram- mosoma. Based upon material examined by the author, Yefremova’s character appears sufficient to separate the two genera. How- ever, the question remains as to whether recognizing Zagrammosoma as distinct will eventually render Cirrospilus paraphyletic. The species described here belongs to Cirrospilus, however it splits the diagnostic characters pertaining to the vaulted vertex and notaulus configuration. Cirrospilus coachellae has the vertex distinctly vaulted and the notaulus straight, while other mem- bers of Cirrospilus do not possess the vault- ed vertex (as seen in Zagrammosoma). Cir- rospilus coachellae has the axillar character used by Yefremova (1996) to differentiate Cirrospilus and Zagrammosoma. Specimens used in this study were bor- rowed from or are deposited in the institu- tions referred to in the type material section. The acronyms are as follows: BMNH: The Natural History Museum, London, U.K. UCR. University of California, River- side, Entomology Research Mu- seum, Riverside, California, U.S.A. USNM: National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A. Cirrospilus coachellae Gates, new species (Figs. 1—4.) Type Material.—Holotype °@, card mounted, ““USA: CA: River. Co., Coachella Valley, NW Salton Sea, 5.VII.96, M. Guil- len, collector/Ex Marmara sp. peelmines on grapefruit,’ deposited in USNM. Paratypes: same data as holotype, card mounted, (Total paratypes: 23 2, 12 6; de- posited in UCR, 8 2, 5 6; deposited in USNM, 8 2, 4 6; deposited in BMNH, 7 2, 3 3). Additional pointed material: *“Mex. Baja Cal. Sur, Las Barracas, 14-IV 59 1985/Coll. P. Debach, Pantrap’’, (4 2, de- posited in BMNH). Diagnosis.—Cirrospilus coachellae can be distinguished from other members of the genus by the unique possession of a vaulted vertex, which is not seen in other species, and the notaulus not curving to intercept the axilla anteriorly. Female.—Length 2.16—2.75 mm. Head yellow except for following, which are brown: three sets of stripes originating from occipital foramen, first set diverging to in- tercept lateral ocelli, second proceeding lat- erally to contact posterodorsal margin of eye, and third set proceeding ventrad and curving anterodorsally to meet ventral eye margin (Fig. 1); lower face with line ex- tending from between toruli to clypeal mar- gin; two stripes originating lateral to toruli and intercepting ventral half of malar sul- cus; frons with two tapering stripes located between anterodorsal eye margin and ex- tending half distance to lateral scrobal mar- gin; two spots between scrobal depression and medial eye margin extending to contact scrobe; scrobal depression; scape and ped- icel dorsally (Fig. 2). Mesosoma and me- tasoma yellow except for following which are brown: median stripe on pronotum, me- soscutum and scutellum; two pairs of stripes dorsolaterally and laterally on pron- otum, these fused anteriorly and diverging posteriorly; lateral lobe with stripe medial- ly; axilla with antero-medial spot; broad submedial stripes on scutellum; propodeum medially from one-third posterior margin to about four-fifths anterior margin, except transverse, teardrop-shaped yellow spots lateral of dorsellum; gastral tergite 1 (Gt1) with triangular spot surrounding petiolar in- sertion, distal point of spot approaching posterior edge of tergite; Gt2—4 with lateral tergal spots separate from medial spots on Gt2 and increasing fusion caudally to form a solid stripe on Gt4; Gt3—7 with broad, longitudinal stripe connecting transverse bands at intersegmental junctions (Fig. 3). Wings hyaline and venation yellowish ex- 60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Scutoscutellar suture Figs. 1—4. tasoma, dorsal view. 4, Wing, dorsal view. Figs. 1, 4, scale bar = 1 mm. Fig. 2, scale bar = 50pm. cept brown at junction of stigmal and post- marginal veins (Fig. 4). Head: Quadrate, 1.1X higher than broad; distance between posterior ocelli 1.5-2.0X greater than distance between posterior ocellus and eye margin. Face fine- ly reticulate and moderately setiferous; oc- cipital carina absent; eyes reddish, eye height nearly 2 greater than malar space; antenna 8-segmented; scape and pedicel with fine longitudinal carinae medially; Cirrospilus coachellae. 1, Head, anterior view. 2, Antenna, lateral view. 3, Mesosoma plus me- pedicel longer than broad; one anellus; two funicular segments setose and lacking sur- face sculpture; Fl 1.0—1.1X as long as broad; F2 0.83—1.0X as long as broad; cla- va 1.7X as long as broad, tapering distally. Mesosoma: Dorsum finely reticulate; mesoscutal midlobe about 1.2 as long as broad; scutellum quadrate to 0.9X as long as broad; propodeum reticulate with smooth median area; callus with few erect setae; coxae with fine longitudinal striations and VOLUME 102, NUMBER 1 with a few setae; hind femur 3.0—5.0X as long as broad. Forewings reaching beyond tip of gaster, with submarginal vein 0.8— 0.9X length of marginal vein; stigmal vein 2.4—4.0X longer than postmarginal vein; all veins setose and stigmal vein more darkly infuscate than remaining veination; wing evenly setose distad of parastigma, except for small asetose area at apex of stigmal vein. Metasoma: Gaster 1.1 longer than me- sosoma; petiole short; tips of ovipositor sheath visible dorsally. Male.—Length 1.67—2.32 mm. Smaller than female, body striping often paler and less extensive than in female. Variation.—Aside from differences in specimen size, the primary variation is in the coloration and striping patterns. Longi- tudinal striping on the mesosoma ranges from contacting the anterior and posterior margins of a tergite (e.g., pronotum, scu- tellum), to being represented only as a cen- tral dash or absent altogether (e.g., parap- sidal stripe). The gastral coloration varies primarily in thickness and extent of stripes. Biology.—This species has been recov- ered in large numbers from the citrus peel- miner, a new species of Marmara (Lepi- doptera: Gracillariidae) (Guillen, Davis and Heraty, in prep.), in the Coachella Valley of Riverside County, CA. It is a gregarious ec- toparasitoid of the peelminer larva and up to 12 C. coachellae have been documented from a single individual (Guillen, personal communication). This same miner attacks the stems and leaves of Nerium oleander L. (Apocynaceae) and Gossypium sp. (Sola- naceae) in Arizona (M. Guillen, personal communication), which also support C. coachellae. Cirrospilus coachellae has also been recovered from another similar undes- cribed species of Marmara mining stems of tree tobacco, Nicotiana glauca Grah. (So- lanaceae). 61 Distribution.—This insect has been re- covered from Riverside County, California, Yuma County, Arizona, and Baja Califor- nia, Mexico. Etymology.—This species is named for the Coachella Valley in the Colorado Desert of southern California where the type series was recovered. ACKNOWLEDGMENTS This work was partially supported by a UC-IPM grant to John Heraty (UCR). Ad- ditional support is provided by a grant from the Citrus Research Board (CRB) to John Heraty to survey native leafminer parasit- oids. Thanks also to John LaSalle (BMNH) for providing additional specimens of this species collected in Baja California Sur and to Arthur Fong of the California State Park System for permission to collect on state park system lands. Thanks to John Heraty who provided critical comments upon ear- lier drafts of this manuscript. LITERATURE CITED Bouéek, Z. 1988. Australasian Chalcidoidea (Hyme- noptera): A biosystematic revision of fourteen families with a reclassification of species. CAB International, Wallingford, England, 832 pp. Bouéek, Z. and R. Askew. 1968. Palaearctic Eulophi- dae (excl. Tetrastichinae). (Hym. Chalcidoidea). Index of Entomophagous Insects. Le Francois, Paris, 260 pp. Gordh, G. 1978. Taxonomic notes on Zagrammosoma, a key to the Nearctic species and descriptions of new species from California (Hymenoptera: Eu- lophidae). Proceeding of the Entomological So- ciety Washington 80: 344-359. LaSalle, J. 1989. Notes on the genus Zagrammosoma (Hymenoptera: Eulophidae) with description of a new species. Proceeding of the Entomological So- ciety Washington 91: 230-236. Schauff, M., J. N. LaSalle, and L. Coote. 1997. Chap- ter 10. Eulophidae, pp. 327—429. In Annotated keys to the Genera of Nearctic Chalcidoidea (Hy- menoptera), NRC Research Press, Ottawa, Ontar- 10, Canada. Yefremova, Z. 1996. Notes on some Palaearctic and Afrotropical species of the genus Zagrammosoma (Hymenoptera: Eulophidae). Entomological Re- view 75(5): 163-171. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 62-68 NEW RECORDS OF TWO XYLEBORUS (COLEOPTERA: SCOLYTIDAE) IN NORTH AMERICA NATALIA J. VANDENBERG, ROBERT J. RABAGLIA, AND DONALD E. BRIGHT (NJV) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, MRC 168, Wash- ington, DC 20560-0168, U.S.A. (e-mail: nvandenb @sel.barc.usda.gov); (RJR) Maryland Department of Agriculture, 50 Harry S. Truman Parkway, Annapolis, Maryland 21401, U.S.A. (e-mail: rabaglrj@mda.state.md.us); (DEB) Research Branch, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada K1A OC6 (e-mail: brightd@em.agr.ca) Abstract.—New American locality records are given for two exotic ambrosia beetles (Coleoptera: Scolytidae) in the genus Xyleborus. Xyleborus pfeili (Ratzeburg), a widely distributed Old World species, is reported for the first time in North America, from three counties in Maryland. Xyleborus californicus Wood, of northern Palearctic origin, but previously established in the western United States, is reported for the first time from Maryland, Delaware, South Carolina and Arkansas. Diagnoses and descriptions are given for the two species, along with scanning electron micrographs of key characters. Modi- fications are made to a previous key to include these new additions to the eastern North American ambrosia beetle fauna. Key Words: The introduction through commerce of exotic bark and ambrosia beetles (Coleop- tera: Scolytidae) poses a threat to our native forests and urban plantings. The adult bee- tles damage trees by tunneling and feeding in the cambium region just beneath the bark’s surface (true bark beetles), or by dril- ling into the sapwood and feeding on intro- duced fungal symbiotes (ambrosia beetles). Although scolytid beetles may promote the general health of a stand by culling overly mature or damaged trees (Atkinson et al. 1990), the effect of foreign species, released from controlling factors of their native en- vironment, can be unpredictable or even devastating (U.S. Congress, Office of Tech- nology Assessment 1993). Within the past decade, scolytid intro- ductions and interceptions have increased concern over the effects of exotic species in Xyleborus, Scolytidae, eastern North America, introductions North America. Recently, several econom- ically important bark beetles have become established on this continent (e.g., Tomicus piniperda (L.) (Haack and Kucera 1993) and Hylastes opacus Erichson (Hoebeke 1994, Rabaglia and Cavey 1994)), or been collected at or near United States and Ca- nadian ports-of-entry (e.g, Ips typographus (L.), Cavey and Passoa 1993). Eight species of exotic ambrosia beetles have been estab- lished in eastern North America within this century (Wood 1977, 1982, Atkinson et al. 1990, Hoebeke 1991), and some of these have become significant pests. We report the occurrence of two additional exotic am- brosia beetles new to eastern North Amer- ica: Xyleborus pfeili (Ratzeburg) and X. cal- ifornicus Wood. Xyleborus pfeili was first detected in North America in a mixed sample of Xy- VOLUME 102, NUMBER 1 leborine ambrosia beetles submitted to one of us (NJV) at the USDA Systematic En- tomology Laboratory, Washington, D.C. The specimens were included along with a sample of frass collected from branch/ stump of paw paw, Asimina triloba (An- nonaceae), at the Wye Research and Edu- cation Center, Carmichael, Maryland, and were reported to have emerged from the host in June and July 1992. The lot con- sisted of 2 specimens of Xyleborinus saxe- seni (Ratzeburg), 3 specimens of Xyleborus affinis Eichhoff, and 5 specimens of another species which could not be identified using the keys to exotic and native North Amer- ican fauna in Wood (1982) or Atkinson et al. (1990). The 5 unknowns were recog- nized (DEB) as a new record for the Old World species, X. pfeili. Additional speci- mens were found (RJR) during bark beetle trapping surveys in Maryland from 1994— 1997, where yet another exotic species, Xy- leborus californicus, was detected for the first time. During manuscript preparation, the latter species was also found in Arkan- sas and South Carolina. Although it was first described from a population in Cali- fornia (Wood 1982), it is now known to be a northern Palearctic species (Siberia (S.L.Wood, pers. comm.), China, new in- terception in British Columbia (DEB)). De- scriptions, diagnoses, revised keys and scanning electron micrographs are provided for each of the newly reported exotic spe- cies to aid in identification. Material from the trapping survey has been deposited in the National Museum of Natural History, Washington, D.C. Xyleborus pfeili (Ratzeburg) (Figs. 2, 4—6) Distribution.—This species is native to Europe (Austria, Belgium, Bulgaria, former Czechoslovakia, France, Germany, Greece, Hungary, Italy, Poland, Romania, Spain, Switzerland, and former western USSR), Asia (China, Japan, Korea and Turkey) and Africa (Algeria and Morocco), and has been 63 introduced into New Zealand (Wood and Bright 1992). Diagnosis.—Among_ eastern North American Xyleborus, specimens of X. pfeili with well-developed declivital tubercles are most likely to be confused with X. celsus Eichhoff and X. ferrugineus (FE). Xyleborus pfeili (Fig. 2) differs from X. celsus (Fig. 1) in its more gradual declivity with greater consistancy in the size of interstrial tuber- cles, and the straighter declivital striae 1. It differs from X. ferrugineus by the presence of tubercles on declivital interstriae 1. Spec- imens of X. pfeili with less developed de- clivital tubercles are most likely to be con- fused with X. volvulus (FE). They can be dis- tinguished by the larger size (X. volvulus females are generally less than 2.8 mm), the weakly sulcate elytra with distinctly raised interstriae 1 (Fig. 2), and the more abruptly formed declivity (Figs. 4—5). Description.—Female: Length 3—3.6 mm, cylindrical, reddish brown, with legs paler yellowish. Frons minutely reticulate, weakly shin- ing, coarsely shallowly punctate; longitu- dinal carina weak to obsolete. Pronotum 1.2 times as long as wide, sides roughly parallel, summit near middle; anterior margin of pronotum arcuate, un- armed; anterior and posterolateral areas weakly reticulate and faintly shining, with moderate to weakly developed asperities and moderately long setae; median area posterior to summit devoid of asperities, more polished, with fewer setae and sparse scattered punctures. Elytra about 1.8 times as long as wide; strial punctures on disc moderately large, each with very short recumbent seta, sepa- rated within row by approximate diameter of a puncture; interstrial discal punctures minute, more widely and less regularly spaced, each with stouter, longer, erect to semirecumbant seta; interstriae | with gran- ules sometimes developed in posterior half before declivity. Elytral declivity steep, oblique, linear in profile, occupying ap- proximately posterior 30% of elytra, weak- 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. ly sulcate, highly polished except for duller patches in depressed areas; rows of strial punctures less regular than on disc, deviat- ing around larger tubercles; 2 or 3 larger conical tubercles often present on interstriae 1 and 3; scattered smaller tubercles or gran- ules often occurring on some or all inter- striae; interstriae | slightly elevated. Male: Not seen. Males of Xyleborus spp. are “‘exceedingly rare and flightless”’ (Wood 1982). Described and illustrated by Balachowsky (1949). Apparently similar to female, except for smaller size (2.1—2.6 mm long) and pronotal shape which is ogival in outline and concave in anterior %. Variation.—Specimens vary in develop- ment and position of declivital armature, al- though specimens from the introduced pop- ulation (Figs. 2, 4—5) thus far show less variation than some of the Old World spec- imens examined. Specimens with reduced declivital tubercles show less variation in the relative size of these structures, and some specimens in a series from Brout-Ver- net, France, entirely lack declivital tuber- cles except at base. Specimens examined (all 2, introduced population).—UNITED STATES: MARY- LAND: Cecil Co.: Elk Neck State Forest, 4 June 1997, ex Chalcoprax-baited funnel trap, R.J. Rabaglia. Kent Co.: Coleman, 4 June 1997, ex Chalcoprax-baited funnel trap, R.J. Rabaglia. Queen Anne’s Co.: Car- michael, Wye Research and Education Cen- ter, June-July 1992, R.N.Peterson, ex branch/stump of paw paw, Asimina triloba; Wye, 5 May 1995, ex ethanol-baited funnel trap, R.J. Rabaglia; Wye, 26 September Posterolateral views of elytral declivity. 1, Xyleborus celsus. 2, X. pfeili. 3, X. californicus. 1996, ex ethanol-baited funnel trap, R.J. Rabaglia. Xyleborus californicus Wood (Fig. 3, 7—9) Distribution.—According to Wood (pers. comm.) this species belongs to a complex indigenous to Siberia and neighboring parts of northern Asia. Recently a specimen from China was intercepted in Vancouver, B.C. The species was originally described from a population established in California and Oregon, but its exotic status was never se- riously in dispute (Wood 1982). The sudden occurrence of X. californicus in the Mid- Atlantic region and southeastern United States further supports the exotic origin of this species. Each year its numbers and dis- tribution have increased. Diagnosis.—Wood (1982) states that this species might be confused with pubescens Zimmermann. It can be distinguished by the more abundant pubescence, the smaller size and the lighter coloration. We found X. cal- ifornicus to be most similar to X. pellicu- losus Eichhoff except for the much smaller size and paler color. Both of the latter spe- cies have the same structure of the antennal club and similar body proportions, puncta- tion and vestiture. Xyleborus pelliculosus was not included in Wood’s key because its presence in the United States was reported subsequently (Atkinson et al. 1990). Description.—The following description is from Wood (1982) with the addition of a phrase (in bold face) to describe the anten- nal club: VOLUME 102, NUMBER 1 Figs. 4-9. Female: Length 2.0—2.2 mm, 2.9 times as long as wide; yellowish brown. Frons rather strongly convex; surface strongly reticulate, a few small granules from epistoma to upper level of eyes. Ves- titure of fine, sparse hair. Antennal club (Fig. 9) flattened, subcircular, posterior face solid or with a subapical suture and rows of setae visible on apical third. Pronotum 1.2 times as long as wide; sides almost straight and parallel on basal two-thirds, rather broadly rounded in front; anterior margin unarmed; summit in front of middle; anterior slope steep, rather coarsely asperate; posterior areas strongly 4—6, Xyleborus pfeili. 4, Dorsal view. 5, Lateral view. 6, Detail of head showing left antenna. 7— 9, X. californicus. 7, Dorsal view. 8, Lateral view. 9, Detail of head showing left antenna. reticulate, punctures small, shallow, rather close. Vestiture of fine, short rather abun- dant hair. Elytra 1.7 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal two-thirds, broadly rounded behind; disc occupying basal three-fourths; striae not impressed, punctures small, shallow, distinct, in rows, spaced by diameter of a puncture; inter- striae three to four times as wide as striae, almost smooth, shining, punctures fine, in indefinite rows in some specimens, distinct- ly confused on basal half in others. Decliv- ity steep, convex, general contours as in pu- 66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bescens; strial punctures large, shallow, dis- tinct, their interior surfaces reticulate-gran- ulate; interstriae only slightly wider than striae, their punctures mostly replaced by minute granules on all interstriae, a few granules on 1, 3 and lateral areas; postero- lateral margin rounded, with an indefinite row of scattered granules. Vestiture of rath- er abundant, short, fine hair, distinctly lon- ger on margins of declivity. Specimens examined (all 2 ).—UNITED STATES: DELAWARE: New Castle Co.: Wilmington, 21 August 1997, Delaware Dept of Agric. Coll. Sussex Co.: Redden State Forest HQ Tract, 18 May 1997, Mi- chael A. Valenti; Redden State Forest Ap- penzellar Tract, 23 May 1997, Michael A. Valenti. MARYLAND: Anne Arundel Co.: Odenton, 18 April 1994, R. J. Rabaglia; Somerset Co.: Wellington, 27 April 1994, R. J. Rabaglia. Calvert Co.: Lusby, 11 April 1995, ex Ipslure-baited funnel trap, R. J. Rabaglia. Caroline Co.: Idylwilde Wildlife Management Area, 12 May 1994, R. J. Ra- baglia. Cecil Co.: Elk Neck State Forest, 9 April 1997, ex Lineatin-baited funnel trap, R. J. Rabaglia. Charles Co.: Indian Head, 11 April 1995, ex Ipslure baited-funnel trap, R. J. Rabaglia. Harford Co.: Upper Cross- roads, 3 May 1994, R. J. Rabaglia. Kent Co.: Sandy Bottom, 9 April 1997, ex etha- nol-baited funnel trap, R. J. Rabaglia. Queen Anne’s Co.: Matapeake, 8 April 1997, ex Chalcoprax-baited funnel trap, R. J. Rabaglia; Wye Mills, 23 April 1997, ex ethanol-baited funnel trap, R. J. Rabaglia; Romancoke, 5 May 1997, ex Ipslure-baited funnel trap, R. J. Rabaglia. St.Mary’s Co.: Mechanicsville, 11 April 1995, ex Ipslure baited-funnel trap, R. J. Rabaglia. Talbot Co.: Longwoods, 12 May 1997, ex ethanol- baited funnel trap, R. J. Rabaglia. Worces- ter Co.: Pocomoke State Forest, 27 April 1994, R. J. Rabaglia. SOUTH CAROLINA: Aiken Co.: Savannah River Site, 7 July 1993, ex turpentine-baited bumper trap, R. D. Klaper Coll. Stephen Co.: Chattahoo- chee National Forest, 10 July 1996, ex lob- lolly-baited tent trap, C. A. H. Flechtmann Coll. ARKANSAS: Pulaski Co.: Little Rock, 21 March 1997, ex oak stump, B. Baldwin Coll. Discussion.—Atkinson et al. (1990) pub- lished a key to females of the eastern North American Xyleborus. The key is modified below in order to accomodate the newly re- corded species. Figure captions in italics re- fer to figures in the original publication. Note that X. validus Eichhoff in Atkinson et al (1990) has been removed from Xyle- borus; it is now placed in Euwallacea (see Wood & Bright 1992) and is not included in the modified key. 1. Anterior margin of pronotum usually armed by several coarse serrations (Fig. 8), however serrations absent or reduced in atratus (Fig. 2); body stout, less than 2.2 times as long as wide; mature color black 6423.5 fes.ci 3. gttracon os Shen ceeeietencne 2 — Anterior margin of pronotum unarmed by large serrations (Fig. 10); body slender, greater than 2.5 times as long as wide; mature color usually yellowish or reddish brown Posterolateral costa on declivity armed by 3-5 distinct tubercles. North-central Unit- ed States and Canada, south to Virginia. 2A8=3 SSMS ee eee ease: obesus LeConte — Posterolateral costa on declivity of uni- form height, may appear slightly undulat- ing, but without denticles (Fig. 9) ..... 3 Anterior margin of pronotum armed by numerous small subequal serrations (Fig. 8); declivital striae impressed, declivital interstriae less than 2 times as wide as striae (Figs: 45 9) ss nn oe) ao eee 4 — Anterior margin of pronotum armed by 2— 6 serrations, median pair conspicuously larger than others; declivital interstriae at least twice as wide as striae. North-central United States and Canada. 2.3—2.6 mm REL, She eMC: Eee ae es ee sayi (Hopkins) Anterior margin of pronotum armed by 6— 8 subequal serrations (Fig. 8); declivity flattened, interstrial setae subequal in length to width of interstriae (Fig. 9). North-central United States and Canada, Pacific Northwest of United States and Canaday228=3 25mm ween dispar (EF) — Anterior margin of pronotum with weakly developed serrations (Fig. 2); declivital interstriae 2 impressed, declivity slightly bisulcate; interstrial setae twice as long as width of interstriae (Fig. 4). Asian exotic, 2(1) 4(3) VOLUME 102, NUMBER 1 (1) 6(5) 7(6) 8(7) 9(6) 10(9) 11(10) Tennessee, Georgia, Maryland, West Vir- einias.3 Osmium: Ws ts ae atratus Eichhoff Declivity strongly concave with obtusely elevated margins on posterolateral areas (Fig. 14); sutural interstriae of declivity armed by 2—4 stout tubercles. Southern Texas to Central America. 3.8—4.2 mm Ato tee 35 ioral Ce A oA eR horridus Eichhoff Declivity convex, flattened, or somewhat impressed near midline, but not concave Denticles on some interstriae much larger than on others (Fig. 1) (Figs. /6-/7) ... = 7 Denticles on all interstriae (where present) more uniform in size (Fig. 2, 3) (Figs. 18— 22 ON Aaa apap soisy sci aus Seekers, Suc iar(acew Nee ANS 9 Declivity steep, flat, surface dull; stria | on declivity strongly curved away from midline, with 2 large, pointed, widely spaced tubercles almost on striae; smaller granules on all interstriae only at base or lateral areas of declivity, not on face, forming a circumdeclivital ring (Fig. 1) (Fig. 16). Eastern North America. 3.6—4.5 1100-0 Wigs oe RPC OREO cho tone celsus Eichhoff Declivity less steep, slightly impressed along midline, surface shining; interstriae 1 and 2 armed only at base by small tu- bercles; interstria 3 with 3 widely spaced denticles, the middle one conspicuously larserthantothers, (Hig. D7) csc nt = 8 Anterior portion of pronotum of female impressed, weakly sulcate. Southeastern United States 2:05275 mm ......... Anterior portion of pronotum of female convex, normal (Figs. /O—//), impressed and sulcate in males (Figs. //, /3). East- ern North America, Neotropics. 2.0—2.3 MIM ee acectay ar ret tee ae SE ne ferrugineus (F.) Surface of declivity opaque, dull (Figs. Ole PAD) PAID) Vecccamstt oe ace ae eR me ae 10 Surface of declivity shining (Figs. 2—3) (Figs. 18, 22) except for duller patches in depressed! areas ).quj eles = faa bel) 12 Declivity broadly sloping, occupying pos- terior 30—40% of elytra (Fig. 23), decliv- ity slightly tapered posteriorly; tubercles of interstriae | and 3 small but conspicu- ous (Fig. 20). Eastern North America, Neotropics. 2.0-2.7 mm ... affinis Eichhoff Declivity steep, occupying posterior 15% of elytra, apex blunt, not tapered; tuber- cles of interstriae 1 and 3 very small (Fig. aN ese tse 5, cy tee Stee cee Fane el Pater aries ee 11 Anterior portion of pronotum of female impressed, weakly sulcate. Eastern North 67 America} 2:3=2-4) mm? 0.3. fed. 28! Fi bre event te ono planicollis Zimmermann — Anterior portion of pronotum of female convex, normal (impressed and sulcate in males). Eastern North America. 2.3—2.7 T2010 See tear gihetns orients xylographus (Say) 12(9) Interstrial setae on elytra and declivity placed in a single median row (Figs. 2, 4) ape ac eae eh ak cide arabe chs fact kaa SP 13 — Interstrial setae on elytra and declivity in 2 or 3 randomly placed rows (Figs. 3, 7) Ne aes. Can Cea oe een care Ree ae 16 13(12) Elytral declivity convex, posterolateral ar- eas not subacutely elevated (Figs. 22, 24) Leche Seri bole Ae a Saran ete ae se ces 14 — Elytral declivity flattened, sloping, pos- terolateral areas subacutely margined (Eaig)RCHUSS FILS os whe Scapa Gs eee aes 15 14(13) Discal interstriae 2 times as wide as striae; some declivital tubercles with height and basal width greater than the diameter of strial punctures. Western North America to Honduras, New York to Georgia. 2.2— Pe i237 1) 01 aga eo ae intrusus Blandford — Discal interstriae less than 1.5 times as wide as striae; some declivital tubercles with height and basal width less than the diameter of strial punctures (Fig. 22). Eastern North America. 2.3—2.7 mm SASL, yd Oe eee eNO pubescens Zimmermann 15(13) Length 2.1—2.8 mm; eye with upper part above emargination wider than antennal club; southern Florida, Texas, widespread in Neotropical region .. volvulus (Fabricius) — Length 3.0—3.6 mm; eye with upper part above emargination narrower than anten- nal club; introduced, Maryland ..... veh tee Biseo Sia ao aie oh pfeili (Ratzeburg) 16(12) Length 2.0-2.2 mm; color yellowish- brown; introduced, California, Oregon, Arkansas, Delaware, Maryland, and South G@arolina ou. a as ae californicus Wood — Length 3.2 mm; color dark brown; intro- duced, Maryland, Pennsylvania ..... Ar Maio 5 bo IAB Sees pelliculosus Eichhotf ACKNOWLEDGMENTS We thank S.L. Wood, Brigham Young University, Provo, Utah for confirming the identification of X. californicus and provid- ing information about its possible origin; Lisa Roberts, Systematic Entomology Lab- oratory, for producing the scanning electron microscope images; C. L. Staines, Edge- water, Maryland, J. W. Brown and D. M. Anderson, Systematic Entomology Labo- 68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ratory, ARS, USDA, Washington, D.C. for their review of earlier versions of this man- uscript. LITERATURE CITED Atkinson, T. H., R. J. Rabaglia, and D. E. Bright. 1990. Newly detected exotic species of Xyleborus (Co- leoptera:Scolytidae) with a revised key to species in eastern North America. Canadian Entomologist 122: 92-104. Balachowsky, A. S. 1949. Coléopteres: Scolytides. Faune de France 50. 320 pp. Cavey, J. and S. Passoa. 1993. Possible new introduc- tion European spruce bark beetle. USDA Forest Service, Northeastern Area, Pest Alert. NA-TP- 18-93 Haack, R. and D. Kucera. 1993. Common pine shoot beetle. USDA Forest Service, Northeastern Area, Pest Alert. NA-TP-05-93. Hoebeke, E. R. 1991. An asian ambrosia beetle, Am- brosiodmus lewisi, new to North America (Cole- optera: Scolytidae). Proceedings of the Entomo- logical Society of Washington. 93(2): 420—424. . 1994. New records of immigrant bark beetles (Coleoptera: Scolytidae) in New York: attraction of conifer-feeding species to ethanol-baited trap logs. Entomological News 105(5): 267-276. Rabaglia, R. J. and J. EF Cavey 1994. Note on the dis- tribution of the immigrant bark beetle, Hylastes opacus Erichson, in North America (Coleoptera: Scolytidae). Entomological News 105(5): 277— 279 U.S. Congress, Office of Technology Assessment. 1993. Harmful non-indigenous species in the United States, OTA-F-565 Washington, DC, U.S. Government Printing Office. Wood, S. L. 1977. Introduced and exported American Scolytidae (Coleoptera). Great Basin Naturalist 37: 67-74. . 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist Memoirs 6. 1356 pp. Wood, S. L. and D. E. Bright. 1992. A catalog of Scol- ytidae and Platypodidae (Coleoptera), Part 2: Tax- onomic Index, Vol. A. Great Basin Naturalist Memotrs 13: 1-833. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 69-81 LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF NEASPILOTA SIGNIFERA (COQUILLETT) (DIPTERA: TEPHRITIDAE) ON HEMIZONIA PUNGENS (HOOKER AND ARNOTT) TORREY AND A. GRAY (ASTERACEAE) IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: rgoeden @ucracl.ucr.edu) Abstract.—Neaspilota signifera (Coquillett) is a bivoltine, monophagous fruit fly (Diptera: Tephritidae) apparently developing solely in the flower heads of Hemizonia pungens (Hooker and Arnott) Torrey and Gray in southern California. The egg, first-, second-, and third-instar larvae, and puparium are described and figured. The mouth- hooks of the first and second instars are bidentate, but tridentate in the third instar. The pair of flattened integumental petals are fused laterally with the well developed, stomal sense organs in the first instar; whereas, the integumental petals are papilliform and progressively more numerous in the second and third instars. The dorsal sense organ is well defined in all three instars. The six oral ridges in the second and third instars are dentate ventrally. The life cycle is of the aggregative type. Most eggs are laid singly between the chaff and ovules of preblossom flower heads and perpendicular to the receptacle. First instars feed on ovules, as do the second instars, which also feed on soft achenes in open flower heads, like all third instars. A third of the third instars examined also pitted the receptacles and apparently supplemented their diet with sap. Pupariation occurs inside the mature flower heads, but no protective cell is formed, as with congeners that overwinter as prepuparia. Instead, F, adults emerge from their cells in early summer (June) and either produce a partial second generation in late- blooming flower heads or pass the summer, fall, and winter in riparian habitats as long- lived adults. Surviving, overwintered adults aggregate the next year in early spring (March—April) on preblossom host plants to mate and subsequently oviposit. A Pter- omalus sp. (Hymenoptera: Pteromalidae) was reared from puparia as a solitary, larval- pupal endoparasitoid. Key Words: Insecta, Neaspilota, Hemizonia, Asteraceae, nonfrugivorous Tephritidae, bi- ology, taxonomy of immature stages, flower-head feeding, monophagy, seed predation, parasitoid Revision of the genus Neaspilota (Dip- tera: Tephritidae) by Freidberg and Mathis (1986) facilitated determination of speci- mens reared from California Asteraceae (Goeden 1989) and stimulated several life- history studies, including those on N. viri- descens Quisenberry (Goeden and Headrick 1992) and N. wilsoni Blanc and Foote (Goeden and Headrick 1999). This paper describes the immature stages and life his- tory of a third species from California, Neaspilota signifera (Coquillett). 70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MATERIALS AND METHODS The present study was based in large part on dissections of subsamples of flower heads of Hemizonia pungens (Hooker and Arnott) Torrey and Gray collected during March—May, 1995-1997 from the San Ja- cinto Wildlife Area at 390-m_ elevation, Lakeview, Riverside Co., administrated by the State of California, Department of Fish and Wildlife. One-liter samples of excised, immature and mature flower heads contain- ing eggs, larvae, and puparia were trans- ported in cold-chests in an air-conditioned vehicle to the laboratory and stored under refrigeration for subsequent dissection, pho- tography, description, and measurement. Five first-, 11 second-, and 15 third-instar larvae, and 11 puparia dissected from flow- er heads were preserved in 70% EtOH for scanning electron microscopy (SEM). Ad- ditional puparia were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temperature for adult and parasitoid emergence. Specimens for SEM were hy- drated to distilled water in a decreasing se- ries of acidulated EtOH. They were osmi- cated for 24 h, dehydrated through an in- creasing series of acidulated EtOH and two, 1-h immersions in hexamethyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, and studied with a Philips XL-30 scanning electron mi- croscope in the Institute of Geophysics and Planetary Physics, University of California, Riverside. Most adults reared from isolated puparia were individually caged in 850-ml, clear- plastic, screened-top cages with a cotton wick and basal water reservoir and provi- sioned with a strip of paper toweling im- pregnated with yeast hydrolyzate and su- crose. These cages were used for studies of longevity and sexual maturation in the in- sectary of the Department of Entomology, University of California, Riverside, at 25 + 1°C, and 14/10 (L/D) photoperiod. Single pairs of virgin males and females obtained from emergence cages also were held in each of six, clear-plastic, petri dishes pro- visioned with a flattened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1994) for observa- tions of their courtship and copulation be- havior. Plant names used in this paper follow Hickman (1993) and Bremer (1994); te- phritid names and adult terminology follow Foote et al. (1993). Terminology and tele- graphic format used to describe the imma- ture stages follow Goeden et al. (1998a, b), Goeden and Headrick (1992, 1999), Goe- den and Teerink (1997, 1998, 1999a, b, c), Teerink and Goeden (1998, 1999), and our earlier works cited therein. Means + SE are used throughout this paper. Voucher speci- mens of N. signifera immature stages, adults, and parasitoids reside in my research collections. RESULTS AND DISCUSSION Taxonomy Adult.—Neaspilota signifera was de- scribed by Coquillett (1894). Freidberg and Mathis (1986) designated a male lectotype from Los Angeles, California, and pictured the unpatterned wing [as did Foote et al. (1993)] along with drawings of the lateral aspect of the head, male right foretarsus, epandrium and cerci, aculeus and its apex enlarged, and spermatheca. Immature stages.—The egg, larvae, and puparium heretofore have not been de- scribed nor illustrated. Egg: Twenty-five ova dissected from four, field-collected, mature females of WN. signifera plus nine eggs measured in situ in dissected, field-collected flower heads (Fig. 5A) were white, opaque, smooth, elongate- ellipsoidal, 0.72 + 0.01 (range, 0.54—0.85) mm long, 0.20 + 0.003 (range, 0.16—0.24) mm wide, smoothly rounded at tapered bas- al end; pedicel also tapered, 0.025 + 0.001 (range, 0.02—0.03) mm long. All eggs found in flower heads were damaged during removal and unsuitable for VOLUME 102, NUMBER 1 scanning electron microscopy. The egg of N. signifera is similar in shape to those of N. viridescens and N. wilsoni but longer and wider on average than that of N. viridescens (Goeden and Headrick 1992) and shorter and narrower than that of N. wilsoni (Goe- den and Headrick 1999). First instar: White, elongate-cylindrical, bluntly rounded anteriorly and posteriorly (Fig. 1A), minute acanthae circumscribe in- tersegmental lines; gnathocephalon with pair of prominent, flattened, integumental petals dorsad to mouthhooks (Fig. 1B-1); mouthhook bidentate (Fig. 1B-2); median oral lobe laterally flattened, apically pointed (Fig. 1B-3); each integumental petal fused laterally with prominent, stomal sense or- gan (Fig. 1B-4). Posterior spiracular plate bears two ovoid rimae, ca. 0.01 mm in length (Fig. 1C-1), and four interspiracular processes, each with two to four branches, longest measuring ca. 0.01 mm (Fig. 1C-2). The poor condition of my specimens of first instar N. signifera allowed few com- parisons with those of N. viridescens (Goe- den and Headrick 1992) and N. wilsoni (Goeden and Headrick 1999); however, their general habitus is similar and their mouthhooks also are bidentate. Like N. wil- soni, but not N. viridescens, the integumen- tal petal is fused laterally with the stomal sense organ in first instar N. signifera, a condition first reported in Trupanea vicina (Wulp) by Goeden and Teerink (1999b). Second instar: White, elongate-cylindri- cal, rounded anteriorly, truncated postero- dorsally (Fig 2A), minute acanthae circum- scribe intersegmental lines (Fig. 2B-1); gnathocephalon conical (Fig. 2B); dorsal sensory organ a well-defined, dome-shaped papilla with a basal pore sensillum on each side (Fig. 2C-1); anterior sensory lobe (Fig. 2C-2, D-1) bears the terminal sensory organ (Fig. 2C-3, D-2), lateral sensory organ (Fig. 2D-3), supralateral sensory organ (Fig. 2D- 4), and pit sensory organ (Fig. 2D-5); sto- mal sense organ prominent, ventrolaterad of anterior sensory lobe (Fig. 2D-6); mouth- hook bidentate (Fig. 2C-4); median oral 71 AccV SpotMagn WD /————4 1004m 10.0kV 3.0 266x 395 Acc.V Spot Magn 100 kV.3.0 6 Fig. 1. habitus, anterior to left; (B) gnathocephalon, antero- lateral view, |— integumental petal, 2— mouth hook, First instar of Neaspilota signifera: (A) 3— median oral lobe, 4— stomal sense organ; (C) posterior spiracular plate, 1— rima, 2— interspiracular process. lobe laterally flattened (Fig. 2C-5); single row of four, papilliform, integumental pet- als dorsal to each mouth hook (Fig. 2C-6, D-7); six oral ridges with long axes parallel in row lateral to mouth hooks, oral ridges 72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON AccV. Spot Magn WD /-——— |. 200 um Acc Spot Magn 10.0. kV. 3.0 _122x 37.2 : 10.0 kV 3:0 09x Spot Magn AccV Spot Magn WD Budo tha Atc:V 100kV 3.0 1484x 368 ‘ Wo 10.0 kV pAcc.V Spot Magn WO kV 3.0 . 2478x Fig. 2. Second instar of Neaspilota signifera: (A) habitus, anterior to left; (B) gnathocephalon, anterolateral view, |— minute acanthae, 2— oral ridge, 3— pit sensillum; (C) gnathocephalon, 1— dorsal sensory organ, 2— anterior sensory lobe, 3— terminal sensory organ, 4— mouthhook, 5— median oral lobe, 6— integumental petal, 7— oral ridge, 8— pit sensillum; (D) gnathocephalon, 1— anterior sensory lobe, 2— terminal sensory organ, 3— lateral sensory organ, 4— supralateral sensory organ, 5— pit sensory organ, 6— stomal sense organ, 7— integumental petal; (E) anterior thoracic spiracle, 1— minute acanthae; (F) posterior spiracular plate, 1— rima, 2— interspiracular process. dentate along ventral margins (Fig. 2B-2, oblong papillae (Fig. 2E); minute acanthae C-7); pit sensilla circumscribe gnathoce- circumscribe anterior margin of prothorax phalon posteriorad of oral ridges (Fig. 2B- (Fig. 2B-1, B-1); lateral spiracular com- 3, C-8); anterior thoracic spiracle with three plexes not seen; posterior spiracular plate VOLUME 102, NUMBER 1 bears three ovoid rimae, ca. 0.01 mm long (Fig. 2F-1), and four interspiracular pro- cesses, each with two to four branches, lon- gest measuring 0.009 mm (Fig. 2F-2). The habitus of the second instar of WN. signifera is more like N. wilsoni (Goeden and Headrick 1999) than the barrel-shaped second instar of N. viridescens (Goeden and Headrick 1992). Unlike both N. viridescens (Goeden and Headrick 1992) and N. wilsoni (Goeden and Headrick 1999), the dorsal sensory organ of N. signifera is well de- fined in the second instar. However, one dis- tinction between the first and second instar common to all three species of Neaspilota is the presence of oral ridges with dentate margins in the latter instar (Goeden and Headrick 1992, 1999). The stomal sense or- gans of the second instar of all three species are well developed and bear sensory struc- tures, variously described as conical in N. viridescens (Goeden and Headrick 1992), papillose in N. wilsoni (Goeden and Head- rick 1999), but verruciform in N. signifera (Fig. 2D-6). Also, the integumental petals of the second instars of all three species are papilliform and about four in number above each mouthhook in N. signifera (Fig. 2C-6, D-7); whereas, in the first instars these structures are broad, flattened, and paired (Goeden and Headrick 1992, 1999). The mouthhooks of the second instar of N. sig- nifera are bidentate, like those of N. wilsoni (Goeden and Headrick 1999): whereas, those of N. viridescens are tridentate (Goe- den and Headrick 1992). Similar, apparent interspecific differences in dentation were noted among mouthhooks of second instar Trupanea spp. (Goeden and Teerink 1999b and references therein). Finally, the inter- spiracular processes each bear two to four branches (Fig. 2F-2), not five to nine branches like those of N. viridescens (Goe- den and Headrick 1992), nor two to six branches like those of N. wilsoni (Goeden and Headrick 1999). Third instar: White to pale yellow, with posterior spiracular plate dark brown to black, elongate-cylindrical, tapering anteri- 713 orly; posterior spiracular plate on caudal segment flattened and upturned dorsally ca. 60° (Fig. 3A), minute acanthae circum- scribe thoracic and abdominal segments an- teriorly, acanthae more numerous on pos- terior segments; gnathocephalon conical (Fig. 3B); dorsal sensory organ a well-de- fined, dome-shaped papilla (Fig. 3D-1), pit sensillum on each side at base of dorsal sensory organ (Fig. 3D-2); anterior sensory lobe (Fig. 3C-1) bears the terminal sensory organ (Fig. 3C-2), pit sensory organ (Fig. 3C-3), lateral sensory organ (Fig. 3C-4), and supralateral sensory organ (Fig. 3C-5); eight to 10 papilliform, integumental petals in double row above each mouth hook (Fig. 3C-6, D-3); six oral ridges laterad of mouthhook, oral ridges dentate along ven- tral margins (Fig. 3B-1, C-7); stomal sense organ prominent ventrolaterad of anterior sensory lobe (Fig. 3C-8, D-4); mouth hook tridentate (Fig. 3B-2, C-9, D-5); median oral lobe laterally flattened, apically pointed (Fig. 3C-10, D-6); prothorax circumscribed anteriorly with minute acanthae (Fig. 3B- 3); anterior thoracic spiracle on posterior margin of prothorax bears three to four ob- long papillae (Fig. 3E); spiracle of metatho- racic lateral spiracular complex absent or not seen, this complex otherwise consisting of vertical row of four verruciform sensilla and verruciform sensillum posterior to up- per verruciform sensillum (not shown, but otherwise like Fig. 3F); abdominal lateral spiracular complex consists of a spiracle (Fig. 3F-1), vertical row of four verruci- form sensilla (Fig. 3F-2), and a single ver- ruciform sensillum posterior to upper ver- ruciform sensillum in vertical series (Fig. 3F-3); caudal segment broadly circum- scribed by minute acanthae (Fig. 3G-1); ste- lex sensilla dorsolaterad and ventrolaterad of posterior spiracular plates (Fig. 3G-2), lateral stelex sensilla not seen; posterior spi- racular plate bears three ovoid rimae, ca. 0.029 mm in length (Fig. 3G-3), and four interspiracular processes (Fig. 3G-4), each with one to three, simple, pointed branches, longest branch measuring 0.009 mm; inter- 74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON | Acc: 10. 2 4 0 a bee 4 « we + ee Acc Spot Magn . WD. H-————-J 10.0kV 4.0 -2038x 36.3 ~— OM e o O80 [G9 VIG 9, Pa So ww wy Fig. 3. Third instar of Neaspilota signifera: (A) habitus, anterior to left; (B) gnathocephalon, anterolateral view, 1— oral ridge, 2— mouthhook, 3— minute acanthae; (C) 1— anterior sensory lobe, 2— terminal sensory organ, 3— pit sensory organ, 4— lateral sensory organ, 5— supralateral sensory organ, 6— integumental petal, 7— oral ridge, 8— stomal sense organ, 9— mouth hook, 10— median oral lobe; (D) gnathocephalon, anterior view, 1— dorsal sensory organ, 2— pit senillum, 3— integumental petal, 4— stomal sense organ, 5— mouth- hook, 6— median oral lobe; (E) anterior thoracic spiracle; (F) first abdominal spiracular complex, 1— spiracle, VOLUME 102, NUMBER 1 mediate sensory complex with a medusoid sensillum (Fig. 3H-1) and a stelex sensillum (Fig. 3H-2) ringed by minute acanthae (Fig. 3H-3). The habitus of the third instar of N. sig- nifera generally is like that reported for N. viridescens Goeden and Headrick (1992) and N. wilsoni (Goeden and Headrick 1999), except that the minute acanthae cir- cumscribe the body segments differently; i.e., in N. signifera, the anterior part of each segment is circumscribed; in N. wilsoni, all intersegmental areas and all abdominal seg- ments except the pleura are circumscribed; and in N. viridescens, the intersegmental ar- eas are free of acanthae. Unlike these two congeners, the dorsal sensory organ is well defined in the third instar of N. signifera (Fig. D-1) as well as in the second instar; however, I was unable to determine whether this also held for the first instar. The dorsal sensory organ is well defined in the first instar of both congeners. Thus, speculation by Goeden and Headrick (1992, 1999) that this intraspecific, differential degree of def- inition of the dorsal sensory organ may be a consistent generic character has now been invalidated by comparison with a third spe- cies of Neaspilota. Additional bases for differention between instars were noted. The integumental petals in the third instar N. signifera are papilli- form and arranged in a double row above each mouthhook, but are less numerous and form a single row in the second instar. The integumental petals in the second and third instars of N. viridescens also are papilliform and situated similarly (Goeden and Head- rick 1992), but in N. wilsoni are fewer in the third instar (Goeden and Headrick 1999). The stomal sense organs of the third instars of all three species are well devel- oped and bear different sensory structures, TS described as several cone-shaped sensilla in N. viridescens (Goeden and Headrick 1992): as papilliform and pit-type in N. wil- soni (Goeden and Headrick 1999); and as verruciform (Fig. 3C—8), or what might be termed ‘“‘compound verruciform’’, as shown in Figure 3D—4. Like the second instar of N. signifera, the third instar has oral ridges with dentate margins, also reported in the second and third instars of N. viridescens (Goeden and Headrick 1992) and N. wilsoni (Goeden and Headrick 1999). Counting only those oral ridges with dentate margins in a single row ventral to the stomal sensory organ, these apparently number six in the second and third instars of all three species examined to date; therefore, further com- parison with additional species may show this to be a consistent generic character. The third instars of Trupanea imperfecta, T. jo- nesi, T. nigricornis, T. pseudovicina, T. sig- nata, and T. wheeleri also bear serrated oral ridges (Goeden and Teerink 1997, 1998, 1999b, Goeden et al. 1998a, Knio et al. 1996, Teerink and Goeden 1998), but these oral ridges appear to be fewer in number, and not arranged in a more or less regular, vertical row laterad to the oral cavity, as in Neaspilota. Also, the mouth hooks of the third instars of N. signifera and N. virides- cens are tridentate (Goeden and Headrick 1992); whereas, those of N. wilsoni are bi- dentate (Goeden and Headrick 1999). Such interspecific differences in dentation are supported by our findings that the mouth hooks of third-instar Trupanea vicina are bidentate; whereas, those of 12 other con- geners examined from California are triden- tate (Goeden and Teerink 1999b and cita- tions therein). Puparium: Mostly white to yellow, with posterior 2—3 segments grayish to black- ened posteriorly, broadly ellipsoidal and <_ 2— verruciform sensillum, 3— verruciform sensillum; (G) caudal segment, 1— minute acanthae, 2— stelex sensillum, 3— rima, 4— interspiracular process; (H) intermediate sensory complex, 1— medusoid sensillum, 2— stelex sensillum, 3-minute acanthae. 76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Acc.V. ‘Spot Magn WD -—————{_ 500 um 10:0 kV 3.0 46x Spot Magn - WD Tf 0. 306x535 6 Fig. 4. itus, anterior to left; (B) anterior end, 1— invagination scar, 2— anterior thoracic spiracle; (C) caudal seg- ment, |— rima, 2— interspiracular process. Puparium of Neaspilota signifera: (A) hab- smoothly rounded at both ends (Fig. 4A), minute acanthae circumscribe body seg- ments anteriorly; anterior end bears the in- vagination scar (Fig. 4B-1) and anterior thoracic spiracles (Fig. 4B-2); caudal seg- ment circumscribed by minute acanthae, two stelex sensilla, dorsad and ventrad of posterior spiracular plates; posterior spirac- ular plate bears three broadly elliptical ri- mae (Fig. 4C-1), and four interspiracular processes, each with two to three branches (Fig. 4C-2); intermediate sensory complex consists of a medusoid sensillum and a ste- lex sensillum. Twenty-five puparia aver- aged 2.43 + 0.03 (range, 2.20—2.70) mm in length; 1.10 + 0.02 (range, 0.92—1.28) mm in width. DISTRIBUTION AND Hosts The distribution of N. signifera as mapped by Foote et al. (1993) restricted this tephritid to Arizona, California, Oregon, and Washington in the western United States North of Mexico. The distri- bution of its main, probably sole, host plant, H. pungens, apparently does not extend far into Mexico (Shreve and Wiggens 1964, Hickman 1993), nor, perhaps, does N. sig- nifera. Accordingly, Freidberg and Mathis (1986) listed only two collection records for N. signifera from northern Mexico, one from Baja California Norte, and one from Sonora. Hemizonia pungens belongs to the sub- tribe Madiinae of the tribe Helenieae of the family Asteraceae (= Compositae) (Bremer 1994). I have not reared N. signifera from six other species of Hemizonia from Cali- fornia, involving a total of 11, 1-liter sam- ples of mature flower heads. Freidberg and Mathis (1986) also listed Lasthenia (as Baeria) fremontii (A. Gray) E. Greene as another host, apparently based on two sep- arate series of specimens labelled “‘ex’’ or “on” this plant and collected over 5- and 18-day periods, respectively, from Califor- nia. Lasthenia fremontii belongs to a dif- ferent subtribe Baeriinae of the Helenieae. I have not reared N. signifera, nor any other species of Neaspilota (only Trupanea spp.), from 18 samples of mature flower heads from three species of Lasthenia, but not in- cluding L. fremontii (a northern California species, Hickman 1993). Furthermore, 11 of the 12 described species of Neaspilota in California for which hosts are now known VOLUME 102, NUMBER 1 (Goeden 1989 and unpublished data) have hosts belonging solely to the tribe Astereae, with only N. sp. prob. punctistigma Benja- min or near, besides N. signifera, having been reared by me from flower heads of a host in the tribe Helenieae (subtribe Pecti- dinae). In addition, the generalist species, N. viridescens, has now been reared from 21 host species in the tribe Astereae and a single host in the Tribe Senecioneae (Goe- den, 1989, Goeden and Headrick 1992, and unpublished data). Therefore, the above in- formation suggests that the host record for L. fremontii in Freidberg and Mathis (1986) is suspect, possibly based on sweep records for adults, which are problematic predictors of reproductive hosts (Headrick and Goe- den 1998), and that N. signifera is a true monophage on H. pungens. BIOLOGY Egg.—In 10, closed, preblossom, imma- ture flower heads of H. pungens, 12 eggs were inserted pedicel-last, usually between the chaff and ovules, and perpendicular to the receptacle (Fig. 5A). No flower head contained any floret damaged by oviposi- tion. The diameters of the receptacles of these 10 flower heads containing eggs av- eraged 3.64 + 0.13 (range, 2.42—4.56) mm, and these heads held an average of 1.2 + 0.1 (range, 1-2) eggs. Larva.—Upon eclosion, the first instars usually tunneled into an ovule, or into a co- rolla before entering an ovule (Fig. 5B). One first instar was found feeding within each of 18, closed, preblossom flower heads, the receptacles of which averaged 3.63 + 0.18 (range, 2.0—5.13) mm in di- ameter. An average of only 1.3 + 0.3 (range, O—3) ovules was damaged in these 18 heads, or based on 93 + 4 (range, 52-— 150) as the average number of ovules/ achenes counted in 51 preblossom to post- blossom flower heads, about 1.4% (range, O-—3.2%) of the ovules were damaged. No receptacles within these 18 infested flower heads were abraded or pitted by feeding of first instars. a Second instars continued feeding on ovules in preblossom flower heads or in soft achenes in open flower heads (Fig. 5C). All fed with their bodies horizontal to and their mouthparts directed towards the receptacles within a series of adjacent ovules/soft achenes, but always well above the recep- tacles. Receptacles of 23 flower heads con- taining second instars were not fed upon and averaged 4.15 + 0.12 (range, 3.13- 5.15) mm in diameter. These flower heads contained an average of 1.3 + 0.1 (range, 1—3) second instars that had destroyed an average of 6.7 + 0.9 (range, 2—18) ovules/ soft achenes, as calculated for the preceding instar, about 7% (range, 2—19%) of the av- erage total of 93 ovules/soft achenes per flower head. Third instars fed mainly on soft achenes in the centers of open flower heads, or in one case, a postblossom flower head. Thir- ty-four flower heads that averaged 4.09 + 0.14 (range, 2.85—5.70) mm in diameter contained an average of 1.3 + 0.14 (range, 1—4) third instars. An average of 25 + 2.4 (range, 7-80) of the soft achenes therein were damaged, or about 27% (range, 8-— 86%) of the average total of 93 ovules/soft achenes per flower head. Third instars fed with their long axes oriented horizontal or perpendicular to, and mouthparts directed towards the receptacles (Fig. 5D). One-third of the third instars in the 34 infested heads examined pitted the receptacles, and thus supplemented their diet with sap that pre- sumably collected in these depressions. Goeden and Headrick (1992, 1999) de- scribed and discussed this type of feeding by N. viridescens and N. wilsoni. However, unlike the third instars of these congeners, the third instar of N. signifera did not sur- round itself by a protective cell within which most individuals pupariated and overwintered. Instead, upon completing their feeding, the larvae of N. signifera sim- ply oriented with their anterior ends away from the receptacles, retracted their mouth- parts, and pupariated, as described for other florivorous tephritids that do not overwinter 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON emma G Fig. 5. Life stages of Neaspilota signifera in Hemizonia pungens: (A) egg laid between florets in preblossom flower head, (B) first instar tunneling into corolla, (C) second instar feeding on ovule, (D) third instar feeding on soft achenes in center of open flower head, (E) puparium in postblossom flower head, (F) male on open flower head, (G) female on open flower head, (H) mating pair, dorsal view; (I) mating pair, lateral view. Lines =) mime VOLUME 102, NUMBER 1 in flower heads, e.g., Trupanea spp. (Head- rick and Goeden 1998). Pupa.—The receptacles of 24 flower heads containing puparia (Fig. 5E) aver- aged 4.51 + 0.11 (range, 3.42—5.70) mm in diameter and bore an average total of 21.2 + 2.0 (range, 8-38) soft achenes, or cal- culated as above, 23% (range, 7—78%) that were damaged by larval feeding. The re- ceptacles of seven (29%) of the 24 flower heads were pitted. These 24 heads con- tained an average of 1.1 + 0.07 (range, 1— 2) puparia. All puparia of N. signifera were found in the center of the flower heads with their anterior ends facing away from the re- ceptacles and their long axes perpendicular to the receptacles (Fig. 5E). Adult.—Adults emerged from mature flower heads, and were long-lived under in- sectary conditions, as six unmated males (Fig. 5F) averaged 67 + 13 (range, 40-88) days, and five virgin females (Fig. 5G) av- eraged 129 + 15 (range, 91-179) days. Such lengthy lifespans are commensurate with the aggregative type of life cycle de- scribed below for this tephritid, and com- pare favorably with average adult longevi- ties reported for adults of N. viridescens (Goeden and Headrick 1992) and N. wilsoni (Goeden and Headrick 1999). The premating and mating behaviors of N. signifera were not studied in the field, but were observed in petri dish arenas found to be so useful with many other non- frugivorous, tephritid species (Headrick and Goeden 1994). Premating behaviors ob- served with N. signifera were abdominal pleural distension, side-stepping, and sway- ing by males (Headrick and Goeden 1994), and wing hamation, sometimes combined with lofting 10—20°, by both sexes (Head- rick and Goeden 1994). However, no troph- allaxis or nuptial gift presentation was not- ed as reported with N. viridescens (Goeden and Headrick 1992). Eighteen matings (Fig. 5H, I) were observed that usually began during the early to late afternoon. These matings involved 12 different pairs of flies and lasted an average of 238 min, very he) close to the average duration of 235 min. reported for N. wilsoni (Goeden and Head- rick 1999), and similarly long to mating du- rations reported with N. viridescens (Goe- den and Headrick 1992). A post-copulatory behavior reminescent of the mate guarding observed with Dioxyna picciola (Bigot) (Headrick et al. 1996) and Euaresta stig- matica Coquillett (Headrick et al. 1995) was observed repeatedly. Six males re- mained atop females after copulation for a few minutes to as long as 50 min, during which time females bent their ovipositors upward nearly 90° while extruding their aculei. Sometimes this behavior was fol- lowed by additional copulations by the same pair; other times, the males did not engage the extended aculei. Seasonal history.—The life cycle of N. signifera in southern California follows an aggregative pattern (Headrick and Goeden 1994, 1998) in which the long-lived adults are the principal overwintering stage. Come spring (March—April), surviving, overwin- tered adults aggregate on preblossom host plants for mating and oviposition. By early summer (June), F, adults emerge from flower heads. A partial second generation is produced on late-flowering plants. Like oth- er monophagous, nonfrugivorous tephritids in southern California, the adults probably pass the dry, hot summer and fall in riparian habitats, including mountain streams and meadows, then overwinter in riparian hab- itats at lower elevations (Headrick and Goe- den 1994, 1998, Goeden and Teerink 1998a, b). Natural enemies.—Four specimens of Pteromalus sp. (Hymenoptera: Pteromali- dae) were reared from puparia of N. signi- fera as solitary, larval-pupal endoparasi- toids. ACKNOWLEDGMENTS I thank Andrew C. Sanders, Curator of the Herbarium, Department of Botany and Plant Sciences, University of California, Riverside, for identifications of plants men- tioned in this paper. Krassimer Bozhilov in 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the Institute of Geophysics & Planetary Physics, University of California, River- side, greatly facilitated my scanning elec- tron microscopy. I thank the California De- partment of Fish and Game, Region 5, Wildlife Management Division, for selected use of the San Jacinto Wildlife Area for this study. The parasitoid was identified by Mi- chael Gates, Department of Entomology, University of California, Riverside, and Harry E. Andersen, Huntington Beach, Cal- ifornia. I also thank Kristine Gilbert and Jeffrey Teerink for their technical assistance and Louie Blanc, Jeffrey Teerink, and Da- vid H. Headrick for their helpful comments on earlier drafts of this paper. LITERATURE CITED Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc. Portland, Oregon. Coquillett, D. W. 1894. New North American Trype- tidae. Canadian Entomologist 26: 71-75. Foote, R. H., E L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York. Freidberg, A and W. N. Mathis. 1986. Studies of Ter- elliinae (Diptera: Tephritidae): A revision of the genus Neaspilota Osten Sacken. Smithsonian Contributions to Zoology 439: 1-75. Goeden, R. D. 1989. Host plants of Neaspilota in Cal- ifornia (Diptera: Tephritidae). Proceedings of the Entomological Society of Washington 91: 164— 168. Goeden, R. D. and D. H. Headrick. 1992. Life history and descriptions of immature stages of Neaspilota viridescens Quisenberry (Diptera: Tephritidae) on native Asteraceae in southern California. Proceed- ings of the Entomological Society of Washington 94: 59-77. . 1999. Life history and description of imma- ture stages of Neaspilota wilsoni Blanc and Foote (Diptera: Tephritidae) on Hazardia squarrosa (Hooker and Arnott) E. Greene (Asteraceae). Pro- ceedings of the Entomological Society of Wash- ington 101: 897-909. Goeden, R. D. and J. A. Teerink 1997. Life history and description of immature stages of Trupanea signata Foote (Diptera: Tephritidae) on Gnaphal- ium luteo-album L. in southern California. Pro- ceedings of the Entomological Society of Wash- ington 99: 747-754. . 1998. Life history and description of imma- ture stages of Trupanea pseudovicinia Hering (Diptera: Tephritidae) on Porophyllum gracile Bentham (Asteraceae) in southern California. Pro- ceedings of the Entomological Society of Wash- ington 100: 361—372. . 1999a. Life history and description of im- mature stages of Trupanea arizonensis Malloch (Diptera: Tephritidae) on Trixis californica Kellog var. californica (Asteraceae) in southern Califor- nia. Proceedings of the Entomological Society of Washington 101: 75—85. . 1999b. Life history and description of im- mature stages of Trupanea wheeleri Curran (Dip- tera: Tephritidae) on Asteraceae in southern Cal- ifornia. Proceedings of the Entomological Society of Washington 101: 414—427. 1999c. Life history and description of im- mature stages of Trupanea vicina (Wulp) (Diptera: Tephritidae) on wild and cultivated Asteraceae in southern California. Proceedings of the Entomo- logical Society of Washington 101: 742-755. Goeden, R. D., J. A. Teerink, and D. H. Headrick. 1998a. Life history and description of immature stages of Trupanea jonesi Curran (Diptera: Te- phritidae) on native Asteraceae in southern Cali- fornia. Proceedings of the Entomological Society of Washington 100: 126—140. . 1998b. Life history and description of im- mature stages of Trupanea actinobola (Loew) (Diptera: Tephritidae) on Acamptopappus sphaer- ocephalus (Harvey and Gray) Gray (Asteraceae) in southern California. Proceedings of the Ento- mological Society of Washington 100: 674-688. Headrick, D. H. and R. D. Goeden. 1994. Reproduc- tive behavior of California fruit flies and the clas- sification and evolution of Tephritidae (Diptera) mating systems. Studia Dipterologica 1(2): 194— Dy. . 1998. The biology of nonfrugivous tephritid fruit flies. Annual Review of Entomology 43: 217-241. Headrick, D. H., R. D. Goeden, and J. A. Teerink. 1995. 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 88: 58-71. . 1996. Life history and description of imma- ture stages of Dioxyna picciola (Bigot) (Diptera: Tephritidae) on Coreopsis spp. (Asteraceae) in southern California. Proceedings of the Entomo- logical Society of Washington 98: 332-349. Hickman, J. C., ed. 1993. The Jepson Manual. Uni- versity of California Press. Berkeley and Los An- geles. Knio, K. M., R. D. Goeden, and D. H. Headrick. 1996. Descriptions of immature stages of Trupanea ni- gricornis and T. bisetosa (Diptera: Tephritidae) from southern California. Annals of the Entomo- logical Society of America 89: 1-11. VOLUME 102, NUMBER | 81 Shreve, FE and I. L. Wiggens. 1964. Vegetation and ams). Proceedings of the Entomological Society Flora of the Sonoran Desert. Stanford University of Washington 100: 431—438. Press, Stanford, California. Teerink, J. A. and R. D. Goeden. 1998. Description of the immature stages of Trupanea conjuncta (Ad- . 1999. Description of the immature stages of Trupanea imperfecta (Coquillett). Proceedings of the Entomological Society of Washington 101: 225-232. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 82-98 REVIEW OF THE NEW WORLD TREEHOPPER TRIBE STEGASPIDINI (HEMIPTERA: MEMBRACIDAE: STEGASPIDINAE): II: FLEXOCENTRUS GODING, STYLOCENTRUS STAL, AND UMBELLIGERUS DEITZ JASON R. CRYAN AND LEwiIs L. DEITZ Department of Entomology, Box 7613, North Carolina State University, Raleigh, NC 27695-7613, U.S.A. (LLD e-mail: lewis.deitz@ncsu.edu); JRC current address: Depart- ment of Zoology, 574 Widtsoe Bldg., Brigham Young University, Provo, UT 84602, U.S.A. (JRC e-mail: jrc233 @email.byu.edu) Abstract.—Three genera in the treehopper tribe Stegaspidini Haupt—Flexocentrus God- ing, Stylocentrus Stal, and Umbelligerus Deitz—are redescribed and illustrated based on adult and nymphal morphology. One new synonymy is included in this work: Flexocen- trus brunneus Funkhouser is a new junior synonym of F. felinus (Haviland). Flexocentrus is now a monotypic genus; Stylocentrus and Umbelligerus have three valid species each. Updated taxonomic keys are given for Stylocentrus and Umbelligerus and a complete species checklist, including synonymies, is given for each genus. Key Words: onomy This work, the third in a series of three publications on the tribe Stegaspidini, in- cludes redescriptions of the genera Flexo- centrus Goding, Stylocentrus Stal, and Um- belligerus Deitz. The genera Bocydium La- treille, Lirania Stal, and Smerdalea Fowler were addressed in part I (Cryan and Deitz 1999a); Lycoderes Germar, Oeda Amyot and Serville, and Stegaspis Germar were treated in part II (Cryan and Deitz 1999b). Part I also included an introduction to this review series, explanations and illustrations of relevant morphological features, a redef- inition of the tribe Stegaspidini, and a tax- onomic key for the identification of includ- ed genera. MATERIALS AND METHODS Methods used in this work were de- scribed in part I (Cryan and Deitz 1999a). A superscript 1 denotes species distribution records from Metcalf and Wade (1965a) Membracidae, Stegaspidini, Flexocentrus, Stylocentrus, Umbelligerus, tax- that have not been confirmed in this work. Unverified distribution records from the lit- erature should be used with caution, as some may be based on misidentified spec- imens. The following codens are used here- in to refer to the collections in which rele- vant specimens are located or have been de- posited. Arnett et al. (1993a) listed the full postal addresses for most of the institutions; those not found in that publication are in- dicated by a dagger (+) following the cod- en. AMNH: American Museum of Natural History, New York, New York, USA. Department of Entomology, The Natural History Museum, Lon- don, United Kingdom. Entomology Section, Monte L. Bean Life Science Museum, Brigham Young University, Pro- vo, Utah, USA. BMNH: BYUE: VOLUME 102, NUMBER 1 ENGL Canadian National Collection of Insects, Eastern Cereal and Oil- seed Research Centre, Agricul- ture and Agri-Food Canada, Re- search Branch, Ottawa, Ontario, Canada. Museu de Entomologia Pe. Jesus Santiago Moure, Universidade Federal do Parana, Departamen- to de Zoologia, Curitiba, Parana, Brazil. Entomological Museum, Depart- ment of Biology, Utah State Uni- versity, Logan, Utah, USA. Instituto Nacional de Biodiver- sidad, Santo Domingo, Costa Rica. Instituto de Zoologia Agricola, Universidad Central de Venezue- la, Maracay, Aragua, Venezuela. Museum of Zoology, Lund Uni- versity, Helgonavagen, Lund, Sweden. North Carolina State University Insect Collection, Department of Entomology, North Carolina State University, Raleigh, North Carolina, USA. Quito Catholic Zoology Muse- um, Departamento de Biologia, Pontificia Universidad Catolica del Ecuador, Quito, Ecuador. Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA. S. H. McKamey Collection, cur- rently at the United States De- partment of Agriculture, Agri- cultural Research Service, Sys- tematic Entomology Laboratory, % National Museum of Natural History, MRC-168, Washington, D.C., USA. T. K. Wood Collection, currently at the Department of Entomolo- gy and Applied Ecology, Uni- versity of Delaware, Newark, Delaware, USA. The Bohart Museum of Ento- DZUP: EMUS: INBC: IZAV: MZLU: NCSU: QCAZ: SEMC: SHMC?: Rw Ci: UGDC: 83 mology, University of California at Davis, Davis, California, USA. Department of Entomology, Na- tional Museum of Natural His- tory, Smithsonian Institution, Washington, D.C., USA. USNM: The location and structure of suprahu- meral horns vary greatly within the tribe Stegaspidini, and even within some genera; nevertheless, the nature of these pronotal extensions usually provides excellent taxo- nomic features at the specific and generic levels. We consider any pronotal extensions located above the humeral angles to be su- prahumeral horns. Thus, the unbranched ‘horns’ of Flexocentrus felinus (Haviland) (Figs. 2—3), the stalked bulbs of Bocydium spp. (Cryan and Deitz 1999a: figs. 9, 11, 13), and the spinelike lateral projections of Stylocentrus spp. and Umbelligerus spp. (Figs. 11, 18, 24, 30—32) are homologous. Genus Flexocentrus Goding, 1926a Flexocentrus Goding 1926a: 106. Type spe- cies: Centruchoides felinus Haviland 1925a: 257, by original designation and monotypy. Diagnosis.—The genus Flexocentrus dif- fers from other stegaspidine genera in hav- ing foliaceous tibiae and a long, triquetrous posterior pronotal process close to the scu- tellum for its entire length. The suprahu- meral horns are stout and unbranched. Adult.—Dimensions (mm): Total length 5.1-7.6. Structure: Thorax: Pronotum (Figs. 2-3): Suprahumeral horns and pos- terior process simple, unbranched. Pronotal surface sculpturing (Fig. 38): Punctate; each puncture (pit) associated with 1 long, narrow seta. Scutellum (Figs. 2—3): Emar- ginate apically. Forewing (Fig. 5): Vena- tion simple; vein R,,, fused basally with R,. Genitalia: ¢: 2" valvulae (Fig. 6) curved dorsally. 6: Lateral plates (Fig. 7) fused to pygofer. Range.—Brazil to Venezuela. Remarks.—This genus superficially re- 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 3mm Figs. 1-9. Flexocentrus felinus. 1, Head of lectotype, anterior aspect (face). 2-3, Head, pronotum, and scutellum, anterolateral aspect (female lectotype of F. felinus and male holotype of F. brunneus, respectively). 4, Left metathoracic femur, tibia, and tarsus, ablateral aspect. 5, Left forewing of lectotype. 6, Female second valvulae, lateral aspect. 7, Male left lateral plate (fused to pygofer), lateral aspect. 8, Male aedeagus and left style, lateral aspect. 9, Late-instar nymph, lateral aspect. sembles a new genus to be described in the tribe Microcentrini (Cryan and Deitz, in press), but differs in the characters diag- nostic for the tribe Stegaspidini (forewing with one r-m and one m-cu crossvein, and vein R,,,; fused basally with R,; Cryan and Deitz 1999a). The generic name Flexocen- trus 1s a combination of the Latin terms “flexo” (from “‘flexus,’’ meaning ‘‘bend’’) and “‘centrus” (from “‘centrum,”> meaning ‘spur or spike’’), probably referring to the slight bend at the midpoint of the posterior pronotal process. Flexocentrus felinus (Haviland) (Figs. 1—9, 38) Centruchoides felinus Haviland 1925a: 276. Flexocentrus felinus: Goding 1926a: 106. VOLUME 102, NUMBER 1 Flexocentrus brunneus Funkhouser 1930a: 410, new synonymy. Type locality—Kartabo, Guyana. Diagnosis.—Flexocentrus felinus has fo- liaceous tibiae, unbranched suprahumeral horns, and a long, triquetrous posterior pronotal process close to the scutellum for its entire length. Adult.—Dimensions (mm): Total length 2 5.8-7.6, 3d 5.1-6.4; width between hu- meral angles 2 2.1—2.9; 6 1.9-2.4; pron- otal length 2 4.6-6.3, ¢ 4.3-5.1; wing length @ 5.4-6.7, 3 4.4—5.4; maximum width of head across eyes 2 2.0-—2.5, 6 1.8—2.3. Coloration: Body ranges from light brown to dark brown; posterior pron- otal process often with a pale stripe basally; legs uniformly brown except for metatho- racic tibiae, with distal % light brown; fore- wings either entirely brown or with basal % brown and distal % hyaline (with brown spots); abdomen light brown. Structure: Head: Face (Fig. 1) bearing fine pubes- cence; postclypeus trilobed, lateral lobes extending beneath foliate lobes; dorsal pro- jections indistinct. Thorax: Pronotum (Figs. 2-3): Covered with fine pubescence; metopidial sulci more or less distinct; hu- meral angles weakly produced; suprahu- meral horns simple, extending laterally (su- prahumeral horns of females are longer than those of males); posterior process with slight dorsal swelling at base; posterior pro- cess triquetrous, spinelike, extending to about % length of forewings (at rest); dorsal margin slightly sinuate, with low median carina extending entire length. Pronotal surface sculpturing (Fig. 38): Pits generally round, moderately deep; bases of pit-asso- ciated setae slightly raised. Scutellum (Figs. 2-3): Short, elevated basally, then flattened to emarginate apex. Legs (Fig. 4): Tibiae foliaceous; metathoracic femur with 2 api- cal, cucullate setae; metathoracic tibiae with enlarged setal rows I and II distinct (row II obscure), with cucullate setae in row II only; first metathoracic tarsomere with 1 apical cucullate seta. Forewing (Fig. 5): 85 Wing shape quadrate, basal % thickened, punctate (except area between vein Cu and claval suture); venation simple; r-m cross- vein often in direct line with r. Genitalia: ?: 2" valvulae (Fig. 6) curved dorsally, slightly broadened at midlength, tapering distally, with small dorsal serrations on broadened area. ¢: Lateral plates (Fig. 7) fused to pygofer; styles (Fig. 8) hooked api- cally; anterior margin of aedeagus (Fig. 8) sinuate in lateral aspect, anterior face of posterior arm with preapical area denticu- late. Late-instar nymph (Fig. 9).—Pronotum produced medially, bearing raised ridges in the approximate positions of the adult su- prahumeral horns; body brown, surface granulate; tibiae strongly foliaceous (almost triangular) and fringed with setae; lateral la- mellae, present on abdominal segments 4— 8, also fringed with setae. Distribution.—Brazil [DZUP]; Ecuador [USNM]; French Guiana [NCSU]; Guyana [SHMC]; Venezuela [IZAV]. Material examined.—Flexocentrus feli- nus (Haviland): Lectotype [2] [BMNH] with) labels: Type*;! “LECTO-/1 YPE”, **Kartabo,/Brit. Guiana./B.M. 1924-519.”’, **Kartabo, Brit. Guiana/July, 1922/e coll. M.D. Haviland/d.d. Collegium Newnha- mense”’, ‘*‘Centruchoides/felinus/Havi- land”, and “LECTOTYPE/?/Centruchoi- des/felinus/Havil./P.S. Broomfield, 1968.”’. Paralectotype [46] [BMNH] with labels: *“‘Para-/type’’, ““PARA-/LECTO-/TYPE’’, **Kartabo,/Brit. Guiana./B.M.1924-519.”’, ‘“*Kartabo, Brit. Guiana/July, 1922/e coll. M.D. Haviland/d.d. Collegium Newnha- mense’’, ‘‘Centruchoides/felinus/Havi- land’, and ““PARA-LECTOTYPE/Centru- choides/felinus/Havil./P.S. Broomfield, 1969... Flexocentrus brunneus Funkhous- er: Holotype [d] [USNM] with labels: ““Mackenzie/Demerara R./BRIT.GUIANA/ June 724 £27.74 Comell, Wi/Lote760/Sub OSH 48" Deitze Research/7le2936-.6 7; *“WDFunkhouser/Collection/1962”’, and “*“HOLOTYPE/Flexocentrus brunneus/W.D. Funkhouser’’. Other specimens: 4 @ from 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SHMC; 2 2 and 1 3 from USNM; | @ and 2 6 from BMNH; 10 @ and 5 ¢ from IZAV; 5 2, 1 d, and 1 2 nymphal exuviae (including Deitz Research #71-260c 2 and Deitz Research #71-260b 3) from NCSU; and 1 2 (Cryan Research #93-204b @) from DZUP. Remarks.—Haviland (1925a) described Centruchoides felinus from four female specimens and one male. In 1930, Funk- houser described Flexocentrus brunneus from a single male collected in British Gui- ana, but no comparison was made to F. fel- inus in his original description. Flexocentrus brunneus is here considered to be a junior synonym of F. felinus, based on examinations of 14 male and 26 female specimens (including the types of both nominate species). Moreover, F. felinus is sexually dimorphic with respect to pronotal shape. Females are more robust, with lon- ger suprahumeral horns (Fig. 2) than males (Fig. 3). The male holotype of F. brunneus is identical with the male paralectotype of F. felinus. Although Deitz (1975a) reported that males lack lateral plates, the presence of distinct lateral lobes on the posterior mar- gins of the pygofer (Fig. 7) suggest rather that the lateral plates are fused to the py- gofer, with the lateral lobes representing the apices of the original plates. Adults and nymphs can be found togeth- er on host plants in small colonies that are attended by ants (Haviland 1925a); Mc- Kamey (personal communication) observed nymphs with ant attendants, whereas near- by adults were apparently not attended. The late-instar nymph bears siightly raised ridg- es on the pronotum in the same position as, and apparently corresponding to, the adult suprahumeral horns. Host plant records are limited to the following: McKamey (per- sonal communication) collected F. felinus from Vismia spp. (Guttiferae), and Haviland (1925a) described one host as a “slender, straggling tree, common in open places, [which] had the twigs and undersides of the leaves covered with rusty brown powder.” McKamey noted that Haviland’s description is consistent with Vismia sp. The Latin specific name, ‘‘felinus,’’ translates as “‘of cats,’ perhaps comparing the pronotal horns to the ears of cats. Genus Stylocentrus Stal 1869a Stylocentrus Stal 1869a: 49. Type species: Bocydium ancora Perty 1833a, by mon- otypy. Diagnosis.—The genus Stylocentrus has the suprahumeral horns and the posterior pronotal process simple, spinelike, and elongate. The posterior pronotal process is elevated far above the scutellum. The ocelli are stalked. Adult.—Dimensions (mm): Total length 5.4—7.5. Structure: Head (Figs. 10, 17, 23): Face usually bearing waxy secretions in variable patterns; ocelli on raised tubercles; eyes slightly stalked. Thorax: Pronotum (Figs. 11, 18, 24): Usually bearing pale waxy secretions (Fig. 39) in variable pat- terns; suprahumeral horns and_ posterior process elongate, simple, with slightly raised ridges; posterior process raised above scutellum for its entire length; humeral an- gle produced to various degrees; metopi- dium with distinct callosities. Pronotal sur- face sculpturing (Fig. 39): Punctate, each pit associated with | long, narrow, seta; some pits appearing as oblong slits rather then round punctures; pronotal surface tu- berculate between pits. Scutellum (Figs. 11, 18, 24): Short, elevated anteriorly, then flattening to acuminate apex. Legs (Fig. 12): Metathoracic femur with 1-3 apical, cucullate setae dorsally; metathoracic tibiae with enlarged setal rows I, II, and III dis- tinct; row I with 1—3 cucullate setae distal- ly, row II with cucullate setae throughout, row III with setae cucullate in apical %; first metathoracic tarsomere with | apical cu- cullate seta. Forewing (Figs. 13, 19, 25): Venation simple; 1 r-m crossvein; 1 m-cu crossvein. Genitalia: 2: 2™ valvulae either of uniform width (Fig. 14) or slightly broadened at midlength (Figs. 20, 26). ¢: VOLUME 102, NUMBER 1 Lateral plates either free (Fig. 21) or fused to pygofer, lacking hooks on posterior mar- gins (Figs. 15, 27); styles (parameres) either hooked (Figs. 22, 28) or clubbed (Fig. 16) apically. Range.—South America (from about 15°S) to Costa Rica. Remarks.—Morphologically, Stylocen- trus ancora appears to have more derived features than either S. championi or S. rub- rinigris. For example, both S. championi and S. rubrinigris have the pronotal pro- cesses unstalked and weakly produced hu- meral angles, whereas S. ancora has the pronotal processes stalked and strongly pro- duced humeral angles. The male lateral plates of S. championi are free, whereas those of S. rubrinigris are fused to the py- gofer (as in the more derived Stegaspidini), indicating that of these two species, S. championi is probably the more plesio- morphic taxon. Although Deitz (1975a) reported that Stylocentrus species lack cucullate setae in metathoracic tibia setal row I, all three spe- cies of this genus bear 1—3 cucullate setae near the apex of this row. They are, how- ever, smaller than the cucullate setae in rows IJ and III. The generic name is a combination of the terms “‘stylo”” (from the Greek “‘stylos,”’ meaning “‘piller or column’’) and “‘cen- trus” (from the Latin “‘centrum,’’ meaning ‘spur or spike’’). KEY TO SPECIES OF ADULT STYLOCENTRUS 1. Pronotum with suprahumeral horns and poste- rior process arising from very short stalk (Figs. 18, 24); humeral angles not well developed — Pronotum with suprahumeral horns and poste- rior process arising from long stalk (Fig. 11); humeral angles acute, well developed .... Se heaters the ola sae Se eep oe att, oe S. ancora (Perty) . 6 lateral plate free, not fused to pygofer (Fig. 21); abdomen red with a black tip (in most specimens); forewing and body coloration blackish; tibiae entirely dark i) pe saagree tude oh. rouge «chi as nah’ Stay Sipe S. championi Fowler — 6 lateral plate basally fused to pygofer (Fig. 27); abdomen uniformly light brown; forewing 87 and body coloration brownish; tibiae light with dark band proximally Stylocentrus ancora (Perty) (Figs. 10—16) Bocydium ancora Perty 1833a: 179. Bocydium trispinosum Guérin-Méneville 1844a: 367. Stylocentrus ancora: Stal 1869a: 49. Type locality—Minas Gerais, Brazil. Diagnosis.—Stylocentrus ancora has the suprahumeral horns and the posterior pron- otal process arising from a slender stalk; the humeral angles are produced into acute spines. Adult.—Dimensions (mm): Total length 2 6.6-7.5, 3 6.4—7.2; width between hu- meral angles @ 2.1—2.4, d 2.0—2.3; pron- otal length 2 6.2-6.6, 6 6.1-6.4; wing length 2 5.6-5.8, ¢ 5.4—-5.7; maximum width of head across eyes @ 1.5-1.6, 6 1.4-1.5. Coloration: Head and thorax black, usually with pale, waxy secretions; scutellum very lightly pigmented (nearly yellow; elevated area may or may not bear waxy secretions); legs light brown proxi- mally, dark from base of tibia to apex of tarsi; forewing brown basally, hyaline from about 0.3 their length with brown pig- ments surrounding veins, making veins ap- pear thickened; abdomen light brown to light red. Structure: Head: Face (Fig. 10) with fine pubescence; ocelli (on tubercles) located below strongly produced dorsal projections; frons with expanded, anteriorly directed plates to either side of postclypeus. Thorax: Pronotum (Fig. 11): Suprahumer- al horns and posterior process arising from long stalk; suprahumeral horns curving pos- teriorly; posterior process extending to tips of forewings; metopidial region (posterior to dorsal border with head) sunken. Pron- otal surface sculpturing: Bases of pit-as- sociated setae slightly raised. Scutellum (Fig. 11): Short, elevated anteriorly, then flattening to acuminate apex; flattened re- gion rising slightly from body. Forewing (Fig. 13): Basal % coriaceous, excluding 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 10-16. Stylocentrus ancora. 10, Head, anterior aspect (face). 11, Head, pronotum, and scutellum, anterolateral aspect. 12, Left metathoracic femur, tibia, and tarsus, ablateral aspect. 13, Right forewing. 14, Female second valvulae, lateral aspect. 15, Male left lateral plate, lateral aspect. 16, Male aedeagus and left style, lateral aspect. VOLUME 102, NUMBER 1 area between vein Cu and the claval suture. Genitalia: 2: 2"! valvulae (Fig. 14) narrow throughout; dorsal ridge of apical % without distinct serrations. 6: Lateral plates (Fig. 15) fused to pygofer, represented by lobe with truncate apex; styles (Fig. 16) elon- gate, clubbed apically, with preapical, dor- sal irregularities; aedeagus (Fig. 16) U- shaped, anterior face of posterior arm not denticulate preapically; anterior arm of ae- deagus arcuate. Late-instar nymph.—Unknown. Distribution.—Brazil [USNM]; French Guiana [NCSU]; Panama!'; Suriname’; Guyana [NCSU]; Peru'; Bolivia'; Ecuador’; Colombia [USNM]; Venezuela [IZAV]. Material examined.—Holotype not ex- amined. Other specimens: 1 ° from CNCI; Vee woy arom, IZAV; ly 9541 6, from MZLU; 10 2, 11 6 from NCSU (including Cryan Research #94—076a 2 and Deitz Re- search #70—211i 3); 1 6 from SEMC; 1 ¢ from SHMC; 9 2, 6 do from USNM. Remarks.—The pronotum and female second valvulae of S. ancora (Figs. 11, 14), the largest of the three Stylocentrus species, differ markedly from those of its congeners (Figs. 18, 20, 24, 26). Although it seems to be the most common species of Stylocen- trus encountered in insect collections, there is no recorded information on its biology or life history. The: Latin specific name, ““ancora,”’ translates as “‘anchor,” probably referring to the resemblance of the suprahumeral horns and an anchor. Stylocentrus championi Fowler (Figs. 17-22, 39) Stylocentrus championi Fowler 1896e: 164. Type locality.—Volcan de Chiriqui, Bug- aba, Panama. Diagnosis.—Stylocentrus championi has the suprahumeral horns and the posterior pronotal process arising from a very short stalk. The posterior pronotal process ex- tends to near the apices of the forewings. Body coloration is generally black; abdom- 89 inal segments 1-7 red and 8-9 usually black. Adult.—Dimensions (mm): Total length 2 5.8-6.0, ¢ 5.4—-5.6; width between hu- meral angles @ 1.4-1.5, o 1.4—1.5; pron- otal length 2 5.1-5.2, ¢ 4.8-5.1; wing length 2 4.6-4.9, 5 4.6—-4.9; maximum width of head across eyes ? 1.4—-1.5, 6 1.4—1.5. Coloration: Head. and thorax black (or, rarely, dark brown); facial area and metopidium may or may not bear pale, waxy secretions; legs dark brown to black (coxae and tarsi may be lighter); forewing black basally, hyaline from about % of their length with dark pigments surrounding veins, making veins appear thickened; ab- domen red, usually with segments 8—9 black.. Structure: Head: Face (Fig. 17) with fine pubescence; ocelli (on tubercles) located on dorsal projections; frontal plates not expanded. Thorax: Pronotum (Fig. 18): Suprahumeral horns and posterior process arising from very short stalk; suprahumeral horns rising dorsally and curving posteri- orly; posterior process extending nearly full length of forewings. Pronotal surface sculpturing (Fig. 39): Bases of pit-associ- ated setae strongly raised. Scutellum (Fig. 18): Short, black, elevated anteriorly, then flattening to acuminate apex; produced area without waxy secretions. Forewing (Fig. 19): Basal % coriaceous, excluding areas between veins M and Cu and between vein Cu and the claval suture; apical limbus nar- row. Genitalia: 2: 2 valvulae (Fig. 20) slightly broadened at about % of their length, tapering to apex; dorsal ridge of broadened area bearing distinct serrations. 6: Lateral plates (Fig. 21) free, without hook; styles (Fig. 22) stout, hooked apical- ly; aedeagus (Fig. 22) U-shaped, anterior face of posterior arm denticulate preapical- ly. Late-instar nymph.—Unknown. Distribution.—Ecuador [QCAZ]; Vene- zuela [IZAV]; Panama [BMNH]; Hondu- ras!; Costa Rica [BYUC]. Material examined.—Lectotype [BMNH] with labels: ““Type’’, [2] “LECTO- 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 3mm Figs. 17-22. Stylocentrus championi. 17, Head of holotype, anterior aspect (face). 18, Head, pronotum, and scutellum of holotype, anterolateral aspect. 19, Right forewing. 20, Female second valvulae, lateral aspect. 21, Male left lateral plate, lateral aspect. 22, Male aedeagus and left style, lateral aspect. TYPE”, “V. de Chiriqui,/2—3,000 ft./Cham- pion.”, “Brit. Mus./1904-55.”, “‘Stylocen- trus/championi. Fowler/Type”, and ‘‘LEC- TOTYPE/®?/Stylocentrus championi/Fowl./ PS. Broomfield, 1969; Paralectotype [¢ with suprahumeral horns partly broken] [BMNH] with labels: ““Para-/type”’, ““PARA-/LECTO- ITYPE”, “V. de Chiriqui,/25—4,000 ft./ Champion.”, “Brit. Mus./1904-55.’’, “B.C.A. Homopt.IL./Stylocentrus/championi,/ Fowl.”’, and ““PARA-LECTOTYPE/Stylo- centrus/championi/Fowl./P.S. Broomfield, 1969.”’; Paralectotype [2 with suprahumeral horns and posterior pronotal process broken] [BMNH] with labels: “‘Para-/type’’, ““PARA- /LECTO-/TYPE”, ‘“‘Bugaba,/800—1,500 ft./ VOLUME 102, NUMBER 1 Champion”, “B.C.A. Homopt.II./Stylocen- trus/championi,/Fowl.”’, ““Brit. Mus./1904- 55.”, and “PARA-LECTOTYPE/Stylocen- trus/championi/Fowl./P.S. Broomfield, 1969.”’. Other specimens: 1 2 from BYUC; 1 2 from EMUS; 2 2, 2 3 from IZAV; 2 @ from QCAZ; 7 2, 4 3 from TKWC (includ- ing Cryan Research #94-159b 2 and Cryan Research #94-160a ¢); 1 6 from UCDC; 4 2,1 3 from USNM. Remarks.—Despite Funkhouser’s (1940a) observation to the contrary, the pronota of S. championi and S. rubrinigris appear quite similar (Figs. 18, 24). How- ever, the color patterns on the body and forewings are plainly different, as are the male lateral plate structures. Stylocentrus championi, the smallest of the three Stylo- centrus species, tends to be blackish in col- or; the wings are usually hyaline between veins R and M, although some specimens from Venezuela (IZAV) have this space uniformly pigmented black, and two spec- imens (one from Ecuador, one from Peru, both in the NCSU collection) have brown- ish forewing pigmentation. The abdomen is red, with segments 8—9 black (in most spec- imens). The male lateral plates are free in S. championi, but are basally fused to the pygofer in S. rubrinigris. Host plant records for S. championi are limited to Hamelia sp. (Rubiaceae) (Wood 1984a) and Miconia sp. (Melastomataceae) (TKWC). Fowler named S. championi in honor of the collector who provided the type series of this species, as well as other stegaspidine species. Stylocentrus rubrinigris Funkhouser (Figs. 23-28) Stylocentrus rubrinigris Funkhouser 1940a: 216: Type locality.—Callanga, Peru. Diagnosis.—Stylocentrus rubrinigris has the suprahumeral horns and the posterior pronotal process arising from a short stalk. The posterior pronotal process extends to 9] nearly % the length the wings. Body col- oration is generally brown. Adult.—Dimensions (mm): Total length 2 6.2-6.4, ¢ 6.2; width between humeral angles 2 1.7—1.8, 5 1.6; pronotal length 9 6.7—-6.8, d—(pronotum broken); wing length @ 5.2-5.4, ¢ 5.2; maximum width of head across eyes 2 1.6—1.7; 6 1.6. Col- oration: Head and thorax black with pale, waxy secretions in defined patterns; pron- otum with brown patch above humeral an- gles; legs light brown with dark band at ba- ses of tibiae; forewings brown basally, hy- aline in distal %4, with brown pigment form- ing a bridge between veins R and M; abdomen uniformly light brown. Structure: Head: Face (Fig. 23) with fine pubescence; ocelli (on tubercles) located on dorsal pro- jections; frontal plates not expanded. Tho- rax: Pronotum (Fig. 24): Suprahumeral horns and posterior process arising from short stalk; suprahumeral horns rising dor- sally and curving posteriorly; posterior pro- cess extending to about % length of fore- wings. Pronotal surface sculpturing: Bases of pit-associated setae moderately raised. Scutellum (Fig. 24): Short, elevated ante- riorly, then flattening to acuminate apex; el- evated area dark (may or may not bear waxy secretions), flattened area lighter. Forewing (Fig. 25): Basal % coriaceous, excluding areas between veins M and Cu and between Cu and claval suture; apical limbus narrow. Genitalia: 9: 24 valvulae (Fig. 26) abruptly broadened at % length, tapered to apex; dorsal ridge of broadened area bearing distinct serrations. 6: Lateral plates (Fig. 27) fused basally to pygofer, represented by lobe with rounded apex; styles (Fig. 28) stout, hooked apically; ae- deagus (Fig. 28) weakly U-shaped, anterior face of posterior arm denticulate apically (teeth minute). Late-instar nymph.—Unknown. Distribution.—Peru [USNM]; Ecuador [NCSU]; Venezuela [CNCI]; Panama [CNCI]. Material examined.—Holotype [°] [USNM] with labels: ‘‘Callanga/Peru’’, 92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 3mm i oR j |: rf f j ei o a ee a Figs. 23-28. Stylocentrus rubrinigris. 23, Head of holotype, anterior aspect (face). 24, Head, pronotum, and scutellum of holotype, anterolateral aspect (dashed lines indicate broken portion of pronotum). 25, Left forewing of holotype (reversed). 26, Female second valvulae, lateral aspect. 27, Male left lateral plate, lateral aspect. 28, Male aedeagus and left style, lateral aspect. “991.” ““WDFunkhouser/Collection/1962”’, and ““HOLOTYPE/Stylocentrus rubrinigris/ W.D. Funkhouser’’. Other specimens: 1 @ from AMNH; 2 6 from CNCI; 1 ¢, 1 ¢ from NCSU (including Cryan Research #94- 160b 3); 1 2 from USNM. Remarks.—In many respects, S. rubrini- gris resembles S. championi, but is paler in color, appearing brownish. The abdomen is not so brightly colored as in S. championi, and none of the distal segments are black. The tibiae are generally lightly colored, with a narrow, dark band proximally. In ad- dition, a “pigment bridge” is usually con- spicuous in the forewing of S. rubrinigris, extending from R to M, and completely di- viding the cell between those two veins. The host plants for this species are un- known. The Latin specific name is a combination of “rubri”’ (from “‘ruber,”’ meaning “‘red’’) and “‘nigris’’ (from ‘‘nigror,’’ meaning ‘“‘blackness’’), probably referring to the col- oration of the type specimen. VOLUME 102, NUMBER | Genus Umbelligerus Deitz 1975a (Figs. 29-37) Umbelligerus Deitz 1975a: 137. Type spe- cies: Umbelligerus peruviensis Deitz 1975a: 137, by original designation. Diagnosis.—Umbelligerus has stalked ocelli, and the pronotal processes are um- belliform, antler-like, without inflated bulbs. Adult.—Dimensions (mm): Total length (from anterior branches of suprahumeral horns to apex of forewings) 6.8—9.6. Struc- wre: Head (Fig. 29)" ‘Ocelli and Jeyes stalked. Thorax: Pronotum (Figs. 30-32): Humeral angles stout; metopidium and lat- eral areas either brown or black and may have white longitudinal bands; suprahumer- al horns and posterior process black, arising from long, central stalk; suprahumeral horns variably branched; posterior process long, sinuate, spinelike. Pronotal surface sculpturing (Fig. 40): Surface granulate and punctate; pits round or slitlike, widely spaced, each associated with one short seta. Scutellum (Figs. 30-32): Inflated anterior- ly, flat and acuminate apically. Legs (Fig. 33): Metathoracic tibiae without cucullate setae in rows I and III (except U. woldai, which has cucullate setae in rows II and Ill). Forewings (Fig. 34): Base weakly co- riaceous (except area between vein Cu and claval suture); 1 rm and | m-cu crossvein present. Genitalia: @: 2™ valvulae (Fig. 35) short and broad, curved dorsally, distal ¥ of dorsal ridge with small serrations. 6: Lateral plates (Fig. 36) slim, fused to py- gofer basally; aedeagus and styles (Fig. 37) relatively stout; aedeagus with anterior face of posterior arm denticulate preapically. Late-instar nymph.—Unknown for all species. Range.—Brazil [SHMC]; Peru [NCSU]; Guyana [BMNH]; Venezuela [IZAV], Pan- ama [DZUP]; Costa Rica [INBC]. Material examined.—Umbelligerus pe- ruviensis Deitz: Holotype [¢] [USNM] with labels: ““Hacienda Maria/Cusco, Peru/ March 12, 1952/EL. Woytkowski’’, *“*Along 93 R. Cosnipata/tropical jungle/900 m.a.s.1.”’, ‘‘Deitz Research/70-211f 3’’, ‘““HOLO- TY PE/Umbelligerus/peruviensis/Deitz/ 1975”, and “ON INDEFINITE LOAN/ FROM N. CAROLINA/ST. UNIV. RA- LEIGH”’’. Paratype [6] [NCSU] with la- bels: “‘Hacienda Maria/Cusca, Peru/March 19, 1952/EL. Woytkowski’’, “Along R. Cosnipata/tropical jungle/900 m.a.s.1.’’, ‘““Deitz Research/72-13a 3”, and ““PARA- TY PE/Umbelligerus/peruviensis/Deitz/ 1975”. Paratype [2, dissected] [USNM] with labels: ““Hacienda Maria/Cusco, Peru/ March 21, 1952/EL. Woytkowski’’, *‘Along R. Cosnipata/tropical jungle/900 m.a.s.l.”’, ‘“Deitz Research/70—21lg @2’’, and “‘PAR- ATY PE/Umbelligerus/peruviensis/Deitz/ 1975”. Other specimens: 1 ¢ from BMNH; 12 from: INBC: 2,2, 2 6 from IZAV; 3 Oo, 16 from: SHME; I ¢ from USNM. Umbelligerus furcillatus Sakakibara: Holo- type [2] [DZUP] with labels: ‘“‘Caruaru.PE/ II-1972/M. Alvarenga’”’ and ‘‘HOLOTY- PUS/Umbelligerus/furcillatus/A.M. Sakaki- bara 1981”. Umbelligerus woldai Sakaki- bara: Type not examined. Other material: 1 2 from BMNH; 1 ¢ from DZUP; 4 2 from INBC. Remarks.—Sakakibara (198la) pub- lished a review of the genus Umbelligerus, describing two new species. Umbelligerus species are remarkable insects, similar in appearance to Stylocentrus and, to a lesser degree, Bocydium. Although records indi- cate that they are widely distributed, Um- belligerus specimens seem to be uncommon in collections. No records of host plants or life histories are available for this genus— labels on some specimens indicate that they were collected at mercury vapor and black lights. If the lowest number of suprahumeral horn branches represents the plesiomorphic condition, then U. furcillatus has the most primitive state of the three Umbelligerus species. Nevertheless, this species has cu- cullate setae only in row II of the metatho- racic tibia, whereas U. woldai has cucullate setae in both rows II and III (a more ple- 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 29-37. Umbelligerus species. 29, U. peruviensis, head, anterior aspect (face). 30—32, Head, pronotum, and scutellum, anterolateral aspect of U. peruviensis, U. furcillatus holotype, and U. woldai, respectively. 33, U. peruviensis, left metathoracic femur, tibia, and tarsus, ablateral aspect. 34, U. peruviensis, right forewing. 35, U. peruviensis, female second valvulae, lateral aspect. 36, U. peruviensis, male left lateral plate and pygofer, lateral aspect. 37, U. peruviensis, male aedeagus and left style, lateral aspect. VOLUME 102, NUMBER 1 Figs. 38-40. Pronotal surface sculpturing of Stegaspidini. 38, Flexocentrus felinus. 39, Stylocentrus cham- pioni. 40, Umbelligerus peruviensis. siomorphic condition). Umbelligerus peru- viensis has cucullate setae only in row II and a pronotal structure that suggests it is an intermediate between the other two, re- gardless of which of those is the more ple- siomorphic species. The generic name is a combination of the Latin terms ‘“‘umbell” (from ‘“‘umbella,” meaning ‘“‘umbrella’’) and “‘igerus”’ (a suf- fix from “‘gero,”’ meaning “‘bearer or car- rier’), referring to the umbrella like supra- humeral horns. KEY TO SPECIES OF ADULT UMBELLIGERUS 1. Each suprahumeral horn with 4 distinct branch- CRT SINER, SY2))\ 5 dad Ben eee D U. woldai Sakakibara 96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mexico: Sme, Lyc bf o* ae © Trinidad: Boc, Lyc, Ste i gery a as _-® Venezuela: Boc, Lyc, Oed, a Ee Ste, Fle, Sty, Umb we e Se ae © Guyana: Boc, Lyc, Oed ; oe ge Ste, Fle, Sty, Umb eee wee _.© Suriname: Boc, Lyc, Oed, ‘ a Ste, Sty Guatemala: Sme, Lyc © Costa Rica: Boc, Sme, Lyc, Oed, Sty, Umb ---o French Guiana: Boc, Sme, Lyc, Oed, Ste, Fle, Sty Panama: Sme, Lyc, Ste, Sty, Umb ° Colombia: Boc, Lyc, Oed, Ste, Sty % Ecuador: Boc, Sme, Lyc, Oed, Ste, Fle, Sty Peru: Boc, Sme, Lyc, Oed, Ste, Sty, Umb oo” Bolivia: Boc, Lyc, Oed, Ste, Sty & > Brazil: Boc, Lir, Lyc, Oed, Ste, Fle, Sty, Umb “Ss Paraguay: Boc, Oed © Argentina: Lyc, Oed Fig. 41. Geographic distribution of Stegaspidini genera. Distribution records are summarized for all genera of Stegaspidini. Latitudinal limits are not indicated. Genera abbreviations are as follows: Boc = Bocydium; Fle = Flexocentrus; Lir = Lirania; Lyc = Lycoderes; Oed = Oeda; Sme = Smerdalea; Ste = Stegaspis; Sty = Stylocentrus; Umb = Umbelligerus. — Each suprahumeral horn with fewer than 4 dis- SPECIES CHECKLIST OF UMBELLIGERUS tinctybranchesi(Eigss.30—31) erie ee eee 2 : 2. Pronotum black with pale waxy secretions; furcillatus Sakakibara each suprahumeral horn with 3 distinct branch- Umbelligerus furcillatus Sakakibara esc(Bigs 3) 5736 ie ae: U. furcillatus Sakakibara 1981la: 67. — Pronotum brown with black processes, and of- ten with pale waxy secretions; each suprahu- meral horn with 3% branches (posterior-most branches with small spines; Fig. 30) ..... 137. SHUM: Parse G TAY jo, Shc seus de U. peruviensis Deitz Woldai Sakakibara peruviensis Deitz Umbelligerus peruviensis Deitz 1975a: VOLUME 102, NUMBER 1 Umbelligerus woldai Sakakibara 198ilay O77 SUMMARY We summarized recent taxonomic chang- es in the tribe Stegaspidini in the first part of this review series (Cryan and Deitz 1999a, Table 1). Currently, two included genera are monotypic: Flexocentrus (F. fel- Table 1. Index to host plants of Stegaspidini. Bellucia sp., Melastomataceae: Lycoderes serraticornis Cecropia sp., Moraceae: Oeda inflata Hamelia sp., Rubiaceae Stylocentrus championi Melastomataceae, unidentified: Bocydium astilatum Bocydium nigrofasciatum Lycoderes serraticornis Lycoderes petasus Miconia sp., Melastomataceae: Bocydium globulare Lycoderes phasianus Stylocentrus championi Pithocarphya poeppigiana (D.C.) Baker aff., Astera- ceae: Bocydium cubitale Unidentified green vines and shoots in shaded places: Stegaspis fronditia [as Stegaspis galeata] Unidentified low shrubs in shaded places: Lycoderes hippocampus Unidentified shrub, in clearings, with reddish powder on twigs and leaf undersides (possibly Vismia sp.): Stegaspis fronditia [as S. laevipennis | Unidentified slender, struggling tree: Stegaspis fronditia [as S. laevipennis | Unidentified small tree, in clearings, with reddish powder on twigs and leaf undersides (possibly Vis- mid sp.): Flexocentrus felinus Vismia sp., Guttiferae Bocydium sp. Flexocentrus felinus Lycoderes sp. (?) Stegaspis spp. 97 inus, for which both sexes are known) and Lirania (L. bituberculata, known only from a female specimen). Immature stages are unknown for all species of the genera Lir- ania, Oeda, Stylocentrus, and Umbellige- rus. The geographical distributions for the genera of Stegaspidini are summarized in Fig. 41. Information concerning host plants used by species of Stegaspidini is woefully deficient. Reliable host records are available for only a few species in the genera Bocy- dium (Cryan and Deitz 1999a), Lycoderes (Cryan and Deitz 1999b), Oeda (Cryan and Deitz 1999b), and Stylocentrus (above). We present an index to the known host plants of Stegaspidini in Table 1. Stegaspidini is the nominate tribe of Ste- gaspidinae, one of the more plesiomorphic membracid subfamilies (Dietrich and Deitz 1993a). The evolutionary relationships among the tribes and genera of Stegaspi- dinae will be explored in a forthcoming phylogenetic analysis of the subfamily (Cryan and Deitz in preparation). ACKNOWLEDGMENTS In addition to those workers and institu- tions acknowledged in parts I and II of this series, we are grateful to R. L. Blinn, W. M. Brooks, S. C. Mozley, H. H. Neunzig, and B. M. Wiegmann, who offered many helpful suggestions for improving this man- uscript; S. H. McKamey and C. H. Dietrich, who supplied invaluable information, ad- vice, and specimens; U. K. Cryan, who helped to validate distribution records and assemble SEM plates; and A. M. Sakaki- bara (DZUP), who lent specimens. Funded in part by the North Carolina Agricultural Research Service (North Carolina State University, Raleigh) and by NSF grant DEB-9815867, this work is based on a por- tion of the thesis submitted by J. R. C. in partial fulfillment of his M.S. degree in en- tomology at North Carolina State Univer- sity. 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED For consistency within membracid liter- ature, we conform to the letter designations for publications listed in bibliographies by Metcalf and Wade 1963a, Deitz and Kopp 1987a, Deitz 1989a, and McKamey 1998a. Arnett, R. H. Jr, G. A. Samuelson, and G. M. Nishida. 1993a. The Insect and Spider Collections of the World. 2"¢ edit. Flora & Fauna Handbook 11. Sandhill Crane Press, Gainesville, 310 pp. Cryan, J. R. and L. L. Deitz. In press. Revision of the New World treehopper tribe Microcentrini (Ho- moptera: Membracidae: Stegaspidinae). Thomas Say Monographs (Entomological Society of America). . 1999a. Review of the New World Treehopper Tribe Stegaspidini (Hemiptera: Membracidae: Ste- gaspidinae): I: Bocydium Latreille, Lirania Stal, and Smerdalea Fowler. Proceedings of the Ento- mological Society of Washington 101: 469—489. . 1999b. Review of the New World Treehopper Tribe Stegaspidini (Hemiptera: Membracidae: Ste- gaspidinae): II: Lycoderes Germar, Oeda Amyot and Serville, and Stegaspis Germar. Proceedings of the Entomological Society of Washington 101: 760-778. Deitz, L. L. 1975a. Classification of the higher cate- gories of the New World treehoppers (Homoptera: Membracidae). North Carolina Experiment Sta- tion Technical Bulletin 225: [i-iv], 1-177. 1989a. Bibliography of the Membracoidea (Homoptera: Aetalionidae, Biturritiidae, Membra- cidae, and Nicomiidae) 1981—1987. North Caro- lina Agricultural Research Service Technical Bul- letin 290: 1-31. Deitz, L. L. and D. D. Kopp. 1987a. Bibliography of the Membracoidea (Homoptera: Aetlionidae, Bi- turritiidae, Membracidae, and Nicomiidae) 1956— 1980. North Carolina Agricultural Research Ser- vice Technical Bulletin 284. [ii] + 39 pp. Dietrich, C. H. and L. L. Deitz. 1993a. Superfamily Membracoidea (Homoptera: Auchenorrhyncha). II. Cladistic analysis and conclusions. Systematic Entomology 18: 297-311. Fowler, W. W. 1896e. Order Rhynchota. Suborder He- miptera-Homoptera. (Continued). Biologia Cen- trali-Americana 2: 161—168. Funkhouser, W. D. 1930a. New genera and species of Neotropical Membracidae. Journal of the New York Entomological Society 38: 405-421. . 1940a. New Peruvian Membracidae (Homop- tera). Journal of the New York Entomological So- ciety 48: 275-292. Goding, E W. 1926a. New genera and species of Mem- bracidae. Transactions of the American Entomo- logical Society 52: 103-110. Guérin-Méneville, E E. 1844a. Insectes. Iconographie du Regne Animal de G. Cuvier, ou représentation d’apres nature de l’une des especes les plus re- marquables, et souvent non encore figurées, de chaque genre d’animaux. Avec un texte descriptif mis au courant de la science. Ouvrage pouvant servir d’atlas a tout les traités de Zoologie (Deux- iéme parte) 1844 (J.B. Baillére, Paris): 1-576. Haviland, M. D. 1925a. The Membracidae of Kartabo, Bartica District, British Guiana, with descriptions of new species and bionomical notes. Zoologica 6: 229-290. McKamey, S. H. 1998a. Taxonomic catalogue of the Membracoidea (exclusive of leafhoppers): second supplement to fascicle 1—Membracidae of the General Catalogue of Hemiptera. Memoirs of the American Entomological Institute 60: 1—377. Metcalf, Z. P. and V. Wade. 1963a. A Bibliography of the Membracoidea and Fossil Homoptera (Ho- moptera: Auchenorhyncha [sic]). North Carolina State [University], Raleigh, iv + 200 pp. . 1965a. General Catalogue of the Homoptera. A Supplement to Fascicle 1—Membracidae of the General Catalogue of Hemiptera. Membracoidea. In Two Sections. North Carolina State University, Raleigh, i—vi, 1—1552. Perty, J. A. M. 1833a. Familia: Cicadariae, pp. 176— 180, plate 35. Jn Delectus animalium articulato- rum, quae in itinere per Brasiliam Annis 1817— 1820 jussu et auspiciis Maximiliani Josephi I. Ba- variae regis augustissimi peracto collegerunt Dr. J.B. de Spix, et Dr. C.EPh. de Martius (Monachii) 3: 125-224; plates 25—40. Sakakibara, A. M. 1981la. Sobre o género Umbellige- rus Deitz, 1975, e descrigao de duas espécies no- vas (Homoptera, Membracidae). Revista Brasilei- ra de Entomologia (Rio de Janeiro) 25: 63-68. Stal, C. 1869a. Hemiptera Fabriciana. Fabricianska Hemipterarter, efter de i KGpenhamn och Kiel f6r- varde typexemplaren granskade och beskrifne. 2. Kongliga Svenska Vetenskaps-Akademiens Han- dlingar 8(1): 1—130. Wood, T. K. 1984a. Life history patterns of tropical membracids (Homoptera: Membracidae). Socio- biology 8: 299-344. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 99-104 DESCRIPTION OF THE LARVA OF HETAERINA INFECTA CALVERT (ODONATA: CALOPTERYGIDAE) RODOLFO NOVELO-GUTIERREZ Instituto de Ecologia, A.C. Departamento de Entomologia. Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico (e-mail: novelor@ecologia.edu.mx) Abstract.—The larva of Hetaerina infecta Calvert is described and illustrated based upon two exuviae of reared final instar larvae, and six last instar larvae. The larva of this species is compared to its close relative H. capitalis Selys. Data on distribution, habitat and habits, as well as a key to larvae for the Mexican species of Hetaerina are provided. Resumen.—Se describe e ilustra la larva de Hetaerina infecta Calvert con base en dos exuvias del ultimo estadio larval obtenidas por cria, y de seis larvas de Ultimo estadio. La larva de esta especie se compara con la de su pariente mas pr6ximo, H. capitalis Selys. Se proporcionan datos sobre su distribucion, habitat y habitos, asf como una clave para las especies mexicanas de Hetaerina. Key Words: Mexico The Neotropical genus Hetaerina cur- rently comprises 37 species (Garrison 1990). Of these, 15 have had their larvae described, although one of them by sup- position: H. americana (Fabricius) (Need- ham 1903), H. auripennis (Burmeister) (Santos 1970a), H. brightwelli (Kirby) (Santos 1972), H. caja (Drury) (Geijskes 1943; Zloty et al. 1993); H. capitalis Selys (De Marmels 1985; Zloty et al. 1993); H. cruentata (Rambur), H. fuscoguttata Selys (Zloty et al. 1993), H. hebe Selys (Santos 1970b), H. majuscula Selys, H. miniata Se- lys (Zloty et al. 1993); H. moribunda Ha- gen (Geijskes 1943 by supposition), H. oc- cisa Hagen in Selys (Geijskes 1946; Zloty et al. 1993); H. sempronia Hagen in Selys (Zloty et al. 1993); H. titia (Drury) (Byers 1930; Zloty et al. 1993); H. vulnerata Ha- gen in Selys (Provonsha and McCafferty 1973). In Mexico, 11 species have been re- corded (Gonzalez-Soriano and Novelo-Gu- Larva, description, key, Hetaerina, Calopterygidae, Odonata, Hidalgo State, tiérrez 1996) of which only three remain with larvae unknown: H. infecta Calvert, H. pilula Calvert and H. rudis Calvert. Here, I provide the description of the larva of H. infecta. Hetaerina infecta Calvert (Figs. 1-12) Material examined.—2 exuviae (6, reared), 6 last instar larvae (2 6, 4 @). MEXICO: Hidalgo State, Pemuxtitla, Rio Zacuala (800 m asl), 1O-MARCH-1994, R. Novelo leg., 1 6, 2 2; 8-APRIL-1994, R. Novelo leg., 2 6, 2 2; Calnali (1,350 m asl), 9-APRIL-1994, R. Novelo leg., 1 &. All deposited in the Entomological Collec- tion of Instituto de Ecologia, A.C., Xalapa (IEXA). Description.—Exuviae yellowish brown; mature larva yellowish with dark bands and dots (Fig. la); hind legs, when fully ex- tended, exceeding caudal appendages; and 100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON f--4 AAS Vay PAL dal reich cecocat Wetec SECT ST Figs. 1-4. Details of morphology of Hetaerina larvae. l1a,b>—4, H. infecta; \c, H. capitalis. 1a, Dorsal view of last instar larva (2) (left legs and antennae omitted, left paraproct and epiproct detached). 1b, Ic, Dorsolateral view of pronotum showing lateral protuberance. 2, Right antenna, dorsal view. 3a, Ventrointernal view of right mandible. 3b, Internal view left mandible. 4a, Prementum, dorsal view. 4b, Detail of left labial palp, dorsal view. VOLUME 102, NUMBER 1 long, triquetral, pointed paraprocts; with lateral spines on the last four abdominal segments. Head: Moderately flattened, maculation pattern as in Fig. la, maximum width 4.68— 4.71 mm; cephalic lobes with a postero-lat- eral low rounded tubercle; occiput widely concave. Antenna 7-segmented (Fig. 2): length of scape 4.06—4.11 mm, 85% of head width, with no setae on inner side ex- ceeding one-third width of scape; relative sizes of antennomeres: 1.0, 0.33, 0.22, 0.14, 0.08, 0.06, 0.04. Mouthparts creamy col- ored; labrum widely notched at apical mar- gin. Mandibles with a separated molar lobe (Fig. 3), right mandible with five incisor cusps, subventral cusp largest and ventral one with a small cusp at its base, molar lobe poorly developed, consisting of two small cusps (Fig. 3a); incisor area of left mandible with five cusps, subventral cusp largest, molar lobe with seven cusps, dorsal one largest (Fig. 3b). Maxilla: Galeolacinia with seven teeth, three long slightly incurved teeth on dorsal margin, three short slightly incurved teeth on ventral margin, apical tooth largest; maxillary palp shorter than galeolacinia, ending in a robust blunt spine. Labium: Prementum-postmentum articula- tion reaching posterior margin of mesoster- num; prementum long (Fig. 4a), basal half narrow then suddenly widened at 65% of length, maximum width of prementum 3.5 mm; depth of median cleft 2.0 mm or 57% of maximum premental width; two setae on internal side of each lobe of ligula, external margin of ligula’s lobes serrate. Labial palp (Fig. 4b) ending in three, long, sharply- pointed hooks, median longest, with a short seta near articulation of movable hook, and a row of 4—5 spiniform setae on apical 25% of dorsal margin; internal margin finely ser- rulated with small spines subequally distrib- uted along it. Thorax: Nota pale, yellowish brown, pleura dark brown; anterior margin of pron- otum concave at middle, posterior margin straight at middle; lateral margin with a low, rounded lateral slightly upturned pro- 101 tuberance (Fig. 1b); inferior margin of pro- pleura with two protuberances, that on proepimeron biggest; inferior margin of meso- and metapleura with small digitiform protuberance near pleural suture. Legs long and slender (Fig. 1), pale, with three dark brown rings on all femora and tibiae; pro- femur with many, terminally curved, point- ed spines on anterior border, neither long thin hairs nor prominent spines on posterior border. Anterior and posterior wing pads wholly pale except for dark stripe on apical third of costal margin, reaching posterior margin of abdominal segment five and bas- al half of abdominal segment six respec- tively. Abdomen: Without middorsal protuber- ances; lateral carinae with row of spines on segments 2—9 increasing in length caudally, those of 7—9 ending in a spine which is small on 7, stout and slightly incurved on 8-9 (Fig. 5). Posterior border of tergites 8— 9 lacking row of spines. Posterior margin of segment 10 with 22—23 spines arranged as follows (Fig. 6): 2—3 small middorsal spines, one large dorsolateral spine lateral to midline; 3 dorsolateral spines, 2 large lat- eral spines and 2 large lateroventral spines on each side, and 2 large ventrolateral spines lateral to midline. Male gonapophys- es pyramidal (Fig. 7), not reaching posterior margin of sternite 9. Female gonapophyses (Fig. 8) almost reaching posterior margin of sternite 10; in ventral view (Fig. 8a), apical half of lateral valvae digitiform, slightly convergent, and with small acutely-pointed tips; in lateral view as in Fig. 8b. Caudal appendages: Male cercus digi- tiform (Fig. 9), bluntly-pointed (Fig. 9a), convergent in dorsal view (Fig. 9b), and as long as tergite 10. Female cercus conical (Fig. 10), acutely-pointed (Fig. 10a), slight- ly convergent (Fig. 10b), 50% as long as tergite 10. Paraprocts triquetral (Fig. 11), in lateral view parallel-sided at basal 70%, then edges gradually convergent; all carinae armed with irregularly sized and spaced spines (Fig. 11b); length:width proportion 5:1, all surfaces densely pigmented except 102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5-13. Details of morphology of Hetaerina larvae. 5—12, H. infecta. 13, H. capitalis). 5a, Ventral view of sternites 7-10 (¢). 5b, Dorsolateral view of lateral carinae of abdominal segments 7—9. 6, Caudal view of 10th abdominal segment showing distribution of marginal spines. 7, Ventral view of male gonapophyses. 8, Female gonapophyses: a, ventral view; b, left lateral view. 9, Male cerci: a, left lateral view; b, dorsal view. 10, Female cerci: a, left lateral view; b, dorsal view. 11, Right paraproct: a, right lateral view; b, detail of part of ventral border. 12, Epiproct: a, left lateral view; b, detail of part of dorsal border. 13, Epiproct, left lateral view (redrawn from De Marmels 1985). VOLUME 102, NUMBER 1 for central, longitudinal, pale band, some- times with pale oval areas on margins. Epi- proct largely foliaceous, laterally com- pressed except for basal 40% which is tri- quetral in cross-section, parallel-sided in lateral view with apex broadly rounded; pigmentation as in paraprocts, although ex- uviae with color pattern as in Fig. 12; length:width proportion 3:1, 66-72% as long as paraprocts. Measurements (in mm).—Exuviae (in al- cohol): Total length (excluding caudal ap- pendages) 21.5—22; paraprocts 9.1—9.3; epi- proct 6.2; hind femur 8—8.1; male cerci 0.7. Last instar larva (in alcohol): Total length (excluding caudal appendages) 20—20.3; paraprocts 9.4—9.5; epiproct 6.2—6.3; hind femur 7.6—7.9; male cercus 0.65, female cercus 0.4. Distribution, habitat and habits.—In Mexico, H. infecta together with H. capi- talis and H. rudis form the group of “‘giant ruby-spots’’ which inhabit mountainous shaded streams at altitudes from 800 to 1800 m running through cloud forests. He- taerina infecta has been recorded from the states of Chiapas, Hidalgo, Nayarit, Oaxa- ca, San Luis Potosi, and Veracruz (Gonza- lez-Soriano and Novelo-Gutiérrez, 1996). In Hidalgo State, H. infecta coexisted sym- patrically with H. capitalis and H. cruen- tata; larvae of the first two species were found among decayed leaves and twigs in zones of moderately flowing water, and also on big rocks at the bottom of small, clear, still water pools, in the same manner as lar- vae of Archilestes spp. Larval populations of H. infecta are apparently smaller than those of the closely related H. capitalis. Only two adult specimens of H. infecta were reared; they emerged two days after capture as ultimate instar larvae on 10- April-1994, very early in the morning when still dark (ca. 0500 h). DISCUSSION The larva of Hetaerina infecta closely re- sembles that of H. capitalis in several de- tails of its morphology, although they are 103 easily separated by the following features (those of H. capitalis in parentheses): Max- imum width of head 4.7 mm (4 mm); depth of median cleft 57% of the maximum pre- mental width (45%); lateral protuberance of pronotum widely rounded (Fig. 1b) (more acute [Fig. lc]); lateral carinae of abdomi- nal segments 4—9 spiny, those of 7—9 end- ing in a spine (only segments 8—9); length: width ratio of paraprocts 5:1 (4.25:1); epi- proct 66-72% of length of paraprocts (60%). Neither De Marmels (1985) nor Zloty et al. (1993) described the maculation pattern of the caudal appendages of Hetaerina cap- italis (although De Marmels, provided an excellent illustration). I found that color pattern usually is the best feature to sepa- rate larvae of these species in the field. He- taerina capitalis have pronounced oval pale areas along the margins of epiproct (Fig. 13) and paraprocts, while H. infecta usually do not, although sometimes these pale areas are barely visible. In H. infecta the caudal appendages are concolorous (Figs. 11—12). KEY TO LARVAE OF MEXICAN HETAERINA This key follows Zloty et al. (1993), modified to include only Mexican species. i; ZOne pain ofsprementali setae.) ..55 4c) ene 4 — Two pairs of premental setae ............ 2 i) . Row of spines on posterior borders of abdom- inal tergites 8—9; without lateral pronotal pro- tuberances; paraprocts covered with short, tu- berculanm-spimesms 17.212 745 ot aeaiaG ee sempronia — Row of spines on posterior borders of abdom- inal tergites 8—9 lacking; with lateral pronotal protuberances; paraprocts without tubercular spines 3 oS) . Lateral pronotal protuberances broadly round- ed in dorsolateral view (Fig. 1b); lateral carinae of abdominal segments 2—9 with row of spines which are more conspicuous on 4—9, postero- lateral margins of 7—9 ending in a spine which is small on 7, flat and large on 8—9; posterior margin of segment 10 with 22—23 spines (Fig. lo) eke Rs US DOA Oy Ae c onl 6) Bat ca ol AOR Al infecta — Lateral pronotal protuberances more acute in dorsolateral view (Fig. lc); lateral carinae of segments 8-9 with a row of conspicuous spines, ending in a postero-lateral flat large 104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON spine; posterior margin of segment 10 with 16— IWS PINGS). teocseeet oe sas Gk o Ola capitalis 4. Mid-dorsal protuberances on abdominal seg- ments 1-9 cruentata — Mid-dorsal abdominal protuberances lacking 5 5. Anterior border of profemur with a few, short, thom=-likelspines) eee ee eeicnen > eee ee caja — Anterior border of profemur without conspic- WOUSSSPINES: 35 so cw Ea ace ee es 6 6. Postero-lateral abdominal spines on segments 8-10 GENO oe. Boy ee is sc eae ee ES 7 7. Labial palp with 3—4 spiniform setae at base of movable hook; each lobe of ligula almost twice as wide at base than maximum width of medianvclett=erse eee americana (in part) — Labial palp with 5—7 spiniform setae at base of movable hook; lobes of ligula narrower basally than the greatest width of the cleft ... 8. Lobes of ligula wider basally than maximum width of median cleft; basal half of prementum 0.33 as wide as length of prementum americana (in part) — Lobes of ligula almost as wide at base as max- imum width of the median cleft; basal half of prementum 0.25 as wide as length of premen- CUIENA Sfeceeeewisreremeec thp te ep ee al tye cy ene fe, Suse aah s uehinles 9 9. Antennal scape with some stiff setae on inner side; pronotal protuberances small and situated at midsegment vulnerata titia — Antennal scape with some long, fine hairs; pronotal protuberances broadly-based and ex- tending to posterior end of segment ..... occisa ACKNOWLEDGMENTS I thank Gordon Pritchard (Calgary, Al- berta), Rosser Garrison (Azusa, California), Enrique Gonzdalez-Soriano (México, D.F) and an anonymous reviewer for their in- valuable comments which improved the fi- nal manuscript. LITERATURE CITED Byers, C. EF 1930. A contribution to the knowledge of Florida Odonata. University of Florida Publica- tions 1: 1-327. De Marmels, J. 1985. On the true Hetaerina capitalis Selys, 1873 and its sibling species Hetaerina smaragdalis spec. nov. (Zygoptera: Calopterygi- dae). Odonatologica 14(3): 177-190. Garrison, R. W. 1990. A synopsis of the genus He- taerina with descriptions of four new species (Odonata: Calopterygidae). Transactions of the American Entomological Society 116: 175-259. Geijskes, D. C. 1943. Notes on Odonata of Surinam. IV. Nine new or little known zygopterous nymphs from the inland waters. Annals of the Entomolog- ical Society of America 36(2):165—184. . 1946. Observations on the Odonata of Toba- go, B.W.I. Transactions of the Royal Entomolog- ical Society of London 97(9): 213—235. Needham, J. G. 1903. Life histories of Odonata, sub- order Zygoptera. Bulletin of the New York State Museum 68(18): 218-279. Gonzalez-Soriano, E. y R. Novelo-Gutiérrez. 1996. Odonata, pp. 147-167. /n Llorente-Bousquets, J., A.N. Garcia-Aldrete, y E.Gonzalez-Soriano, eds., Biodiversidad, taxonomia y biogeografia de artr6- podos de México: Hacia una sintesis de su cono- cimiento. UNAM-CONABIO, México. Provonsha, A. W. and W. P. McCafferty. 1973. Previ- ously unknown nymphs of Western Odonata (Zygoptera: Calopterygidae, Coenagrionidae). Proceedings of the Entomological Society of Washington 75(4): 449—454. Santos, N. D. 1970a. Contribuigao ao conhecimento da fauna do estado da Guanabara 74. Descri¢ao da ninfa de Hetaerina auripennis (Burmeister, 1839) Selys, 1853 e notas s6bre 0 imago (Odon- ata: Agrionidae). Atas da Sociedade de Biologia do Rio de Janeiro 13(3—4): 115-117. . 1970b. Contribuigaéo ao conhecimento da fau- na do estado da Guanabara e Arredores. 77. Des- crigdo do Hetaerina hebe Selys, 1853 (Odonata: Agrionidae). Atas da Sociedade de Biologia do Rio de Janeiro 14(3—4): 89-90. . 1972. Contribuigao ao conhecimento da fauna do estado da Guanabara. 78. Descrigao da ninfa de Hetaerina brightwelli (Kirby, 1823) Selys, 1953 [sic — 1853](Odonata: Agrionidae). Atas da Sociedade de Biologia do Rio de Janeiro 15(2): 75-76. Zloty, J., G. Pritchard, and C. Esquivel. 1993. Larvae of the Costa Rican Hetaerina (Odonata: Calopter- gidae) with comments on distribution. System- atic Entomology 18: 253-265. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 105-107 DESCRIPTION OF A NEW SPECIES OF THE GENUS STIRICORSIA KONOW (HYMENOPTERA: ORUSSIDAE) FROM JAPAN IcHIJI TOGASHI 1-chome, Honmachi, Tsurugi-machi, Ishikawa Prefecture 920-2121, Japan Abstract.—Stiricorsia apicalis, n. sp., from Japan is described and illustrated. A key is given to separate the two species of Stiricorsia known from Japan. Key Words: Orussidae, Stiricorsia, Japan While studying specimens of the orus- sid genus Stiricorsia Konow, I found an undescribed species which is closely al- hed to S. tosensis (Tosawa and Sugihara 1934). The new species is separated from S. tosensis by characters of the head, the wing maculation, the length of the hind basitarsus, the apical margin of the su- bgenital plate, and the penis valve of the male genitalia. Though some characters may be variable in S. tosensis, the mag- nitude of differences in the new species appear sufficient to justify describing it as new based on the single specimen avail- able. This is the second species of Stiricorsia known from Japan. Stiricorsia may be sep- arated from other genera of Orussidae by using the keys in Benson (1938) and Tsu- neki (1963). KEY TO MALES OF STIRICORSIA FROM JAPAN 1. Apical portion of forewing infuscate (Fig. 4); stigma of forewing dark brown (Fig. 4); apical margin of subgenital plate with rather long pro- jection (Fig. 10); penis valve as in Fig. 11 PO eR GR POR apicalis, new species — Apical portion of forewing clear (Fig. 7); stig- ma of forewing rather white (Fig. 7); apical margin of subgenital plate with short projection (igs): penis*valvejasunFion 15 54... = ebay ear eee eae tosensis (Tosawa and Sugihara) Stiricorsia apicalis Togashi, new species (Figs. 1-4, 8-11) Male.—Length, 8 mm. Head and body including antenna and legs black. Wings hyaline; apical portion of forewing infus- cate (Fig. 4); stigma and costa of forewing dark brown (Fig. 4); other veins dark brown to black. Head: Face with 2 longitudinal carinae and with transverse carina in front of an- terior ocellus and with transverse carina on clypeus; postorbital carina distinct, reaching near top of eyes (Fig. 2); ocellar region flat- tened, surrounded by a series of 4 pairs of tubercles (Fig. 1); anterior tubercle forming lateral end of upper carina of front shield (Fig. 1); postocellus completely in touch with eye; occipital carina distinct. Head in profile as in Fig. 2. Antenna with basal 5 segments as in Fig. 3, relative lengths of basal 5 segments about 1.8:1.0:2.2:1.8:1.6. Thorax: Pronotum narrow, anteriorly rounded and posteriorly roundly emargin- ate; mesoscutum with slightly raised medi- an longitudinal carina; lateral side of me- soscutum with a distinct carina. Mesoscu- tellum slightly raised, anterior margin dis- tinctly crenulate. Wing venation as in Fig. 4. Legs with hind tibia, in lateral view, as in Fig. 8; hind basitarsus nearly as long as following 3 segments combined (ratio about 1.0:1.0) Gee Figs 9). 106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-7. antennal segments, lateral view. 4, 7, Forewing. Abdomen: Apical margin of subgenital plate, in ventral view, with rather long projection (Fig. 10); penis valve as in Fig. Lele Punctation: Head very closely, deeply, coarsely reticulate; anterior area of front ocellus impunctate, shining (see Fig. 1); thorax very closely, deeply and coarsely re- ticulate; central portion of mesoscutellum practically impunctate, shining; mesepime- ron covered with minute, rather deep and reticulate punctures, matt; abdominal ter- gites and sternites very closely, coarsely re- ticulate, posterior margin of each tergite nearly impunctate; central portion of su- bgenital plate practically impunctate, shin- ing. Female.—Unknown. Distribution.—Japan (Honshu). Bee 1—4, Stiricorsia apicalis. 5—7, S. tosensis. 1, Head, dorsal view. 2, 5 Head, profile. 3, 6, Basal 5 Holotype.—Male, 9. VIII.1974, Mt. Hakusan, Ishikawa Pref., I. Togashi leg. Deposited in the collection of the National Science Museum (Natural History), To- kyo. Remarks.—This new species is closely allied to S. tosensis, but it is easily separat- ed from the latter by the characters given in the key and the following additional characters: length of the 3rd antennal seg- ment (in S. tosensis, the third antennal seg- ment is nearly twice as long as the second); characters of the mesoscutellum (in S. to- sensis, the mesoscutellum is nearly flattened and the punctures on the mesoscutellum are sparse with the interspaces between them impunctate and shining); and the lengths of the hind basitarsus (in S. tosensis, the hind basitarsus is slightly longer than the follow- VOLUME 102, NUMBER 1 ie Figs. 8-15. 8-11, Stiricorsia apicalis. 12-15, S. tosensis. 8, 107 12 11 12, Hind tibia, lateral view. 9, 13, Hind tarsus, lateral view. 10, 14, Apical margin of subgenital plate, ventral view. 11, 15, Penis valve. ing three segments combined, the ration about 1.0:0.9) (see Fig. 13). ACKNOWLEDGMENTS I thank Dr. David R. Smith, U.S. De- partment of Agriculture, Washington, D.C., for reviewing this manuscript. I also thank Dr. A. Shinohara, National Science Muse- um (Natural History), Tokyo, for lending me the valuable specimens. LITERATURE CITED Benson, R. B. 1938. On the Australian Orussidae, with a key to the genera of the world (Hymenoptera, Symphyta). Annals and Magazine of Natural His- tory, Series 11, 2: 1-15. Tosawa, N. and Y. Sugihara. 1934. Ueber eine neue Oryssus Art aus Japan. Transactions of the Kansai Entomological Society, No. 5, pp. 1—2. [In Japa- nese. | Tsuneki, K. 1963. A contribution to the knowledge of Orussidae in Japan, with the description of a new species (Hym., Symphyta). Etizenia, No. 2, pp. 1-5. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 108-119 DESCRIPTION OF THE PUPAE OF FIVE SPECIES IN THE SUBGENUS ARMIGERES, GENUS ARMIGERES THEOBALD, WITH A KEY TO SPECIES OF THE KNOWN PUPAE OF THE SUBGENUS (DIPTERA: CULICIDAE) RICHARD E DARrsIE, JR. Florida Medical Entomology Laboratory, University of Florida, 200 9th Street, SE, Vero Beach, FL 32962, U.S.A. (e-mail: rfd@icon.vero.ufl.edu) Abstract.—Descriptions and illustrations of the unknown pupae of Armigeres (Armi- geres) aureolineatus (Leicester), Ar. (Arm.) durhami Edwards and Ar. (Arm.) kuchingensis Edwards and the partially described pupae of Ar. (Arm.) subalbatus (Coquillett) and Ar. (Arm.) baisasi Stone and Thurman are presented. A key to 15 species with known pupae of the subgenus Armigeres is included. Key Words: There are 33 species in the subgenus Ar- migeres Theobald, genus Armigeres Theo- bald. The pupae of only 15 are adequately described (Belkin 1962, Penn 1949, Ra- malingam 1972, 1987, Steffan 1968, Toma et al. 1994, 1995) including this work in which the unknown pupae of Armigeres (Armigeres) aureolineatus (Leicester), Ar- migeres (Armigeres) durhami Edwards and Armigeres (Armigeres) kuchingensis Ed- wards and partially described Armigeres (Armigeres) subalbatus (Coquillett) and Ar- migeres (Armigeres) baisasi Stone and Thurman (Delfinado 1966, Baisas 1974, LaCasse and Yamaguti 1950, Bohart and Ingram 1945) are presented. This work began as a study of the Ar- migeres species occurring in Nepal (Darsie 1998) and the former four species listed above are found there. Pupae of all species of Armigeres known from Nepal have now been described, save Ar. (Lei.) omissus (Ed- wards) which will be described later. For- tuitiously, I have the opportunity to de- scribe also the pupa of Ar. baisasi, occur- ring only in the Philippines, but belonging to the subgenus treated here. genus Armigeres, subgenus Armigeres, pupae, key The identity of Ar. durhami and Ar. ku- chingensis has been a problem in this study. The first specimens of ‘“‘durhami’’ studied were from Surat Thani Province, Thailand, located on the southern coastal plain. They were finally identified as Ar. kesseli Ra- malingam. The valid durhami is a montane species and the specimens used in the pupal description were collected at Fraser Hill and Cameron Highlands, Malaysia, at eleva- tions above 920 m (Ramalingam 1987). A more vexing enigma was the identifi- cation of Ar. kuchingensis. According to Thurman (1959), the larval description by Barraud (1934) refers to Ar. durhami, and in her account, the larva has seta 1-X sin- gle, stout and attached to the saddle. My study specimens, loaned through the kind- ness of Dr. S. Ramalingam, were individu- ally reared, mounted larval and pupal exu- viae. The larvae had seta 1-X tiny, 4- to 6- branched, attached at the border or ventral to the saddle. A check of the associated adults, 3d’s and 2°’s, confirmed that they indeed were kuchingensis. Thurman’s de- scription of the larva was taken from Borel (1930) which she had not seen to confirm its identity. VOLUME 102, NUMBER 1 Except for Ar. baisasi, these species are widely distributed throughout south and southeast Asia. They breed in a variety of natural and artificial containers, especially cut bamboo nodes. METHODS AND MATERIALS The methods are the same as described by Darsie (1998). Those pupae collected in Nepal are in the author’s collection at the Florida Medical Entomology Laboratory, Vero Beach, Florida. The others from Bru- nei, Malaysia, Singapore and Thailand were borrowed from the national mosquito col- lection at the Walter Reed Biosystematics Unit, Smithsonian Institution, Washington, D.C. and from the private collection of Dr. S. Ramalingam. The ‘‘b’’ in the descriptions means branches and Le and Pe refers to the exu- viae of larvae and pupae. DESCRIPTIONS Armigeres (Armigeres) aureolineatus (Leicester) (Fig. 1) Position and size of setae as figured; range and modal number of branches in Ta- ble 1. Cephalothorax: Seta 1-CT with thin branches; 3-CT stout, usually double; seta 7-CT 0.78—0.83 or, as in one specimen, 1.25—-1.86 length of 6-CT; trumpet length 0.8—0.93 mm, index 3.0—4.0, pinna 0.12— 0.23 of total length. Abdomen: Seta 1-II with single short stem, branched in outer 0.8-0.9, 3-6b, 3-II 1.86—2.3 longer than 2- II, 0.89—1.39 length of tergum III; 3-IIl 0.94—-1.0 length of tergum IV; 5-IV 1.0— 1.45 longer than tergum V; 5-V 0.9-1.18 length of tergum VI; 6-VI with 4—6 b, stouter than 5-V; 9-VII aciculate, with 9— 19 b, all branches without distal branching; 9-VIII aciculate, with 9-15 b. Paddle: length 1.02—1.34 mm; index 1.45-1.6; fringe in apical 0.58—0.75 of outer margin, in apical 0.65—0.77 of inner margin, fringe length 0.22 mm; seta 1-P single, 0.40—0.45 length of paddle. 109 The description is based on the following specimens from Thailand: Nakornayok Province, Hongking, IX-25-63, 22 LePe (slides 8567, 8568), 1d LePe (slide 8563), ex artificial container; Kanchanburi Prov- ince, Saeng, VI-4-65, 12 LePe, ex coconut on ground. In Nepal, the only confirmed re- cord is a single larva from SUNSARI, Gho- pal, [IX-X-84 (Darsie and Pradhan 1990). Armigeres (Armigeres) durhami EDWARDS (Fig. 2) Position and size of setae as figured, range and modal number of branches in Ta- ble 2. Cephalothorax: Setae 1, 3-CT stout, 1 single, 3 double; 7-CT 0.46—0.61 length of 6-CT, with 2-5 b; trumpet length 0.65— 0.82 mm, pinna 0.23—0.39 total length; in- dex 2.12—2.61. Abdomen: Seta 1-II pedun- culate, brush-like, with 17-27 b in outer 0.65—-0.77; 3-II 2.02—3.04 length of 2-II; 0.95—1.08 length of tergum II; 3-III 0.90— 1.15 length of tergum IV; 5-IV 1.19-1.41 length of tergum V; 5-V 1.16—1.31 length of tergum VI; 6-VI single to triple, stouter than 5-V, except in one specimen in which 6-VI was poorly developed; 9-VII acicu- late, stout, with 3-5 b; 9-VIII aciculate, stout, with 7—10 b. Paddle: ovoid, emargin- ate apically, length 1.11—1.31 mm, index 1.15—1.35, fringe on outer margin in apical 0.48—0.57, on inner margin in apical 0.31— 0.59, fringe length 0.17—0.22 mm; 1-P sin- gle, 0.24—0.40 length of paddle. The description is based on the following specimens from Malaysia: Pahang, Camer- on Highlands, 1970, 12, 1d LePe (Nos. 1511-10, 11); Fraser’s Hill, 1970, 12, 2¢ LePe (Nos. 787—36, 40, 88). Nepal speci- mens without immatures are recorded from Makwanpur District, Hetaura, I-90, 12; Sunsari District, Bhotepur, [X-X-83, 22; Manglabre, IX-X-83, 22; Morang District, Dubai, [X-X-85, 12 (Darsie and Pradhan 1990). Armigeres (Armigeres) kuchingensis EDWARDS (Fig. 3) Position and size of setae as figured; range and modal number of branches in Ta- 110 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pore Foder-7edly, / / ys i aN Fig. 1. Pupa of Armigeres aureolineatus. A, Cephalothorax. B, metanotum and abdomen. Abbreviations: Bu = external buttress; CT = cephalothorax; GL = genital lobe; Mr = midrib; Pa = paddle; T = trumpet. VOLUME 102, NUMBER 1 Table 1. 11] Pupal chaetotaxy of Armigeres aureolineatus. SSE aS... 2.2 2:°9»°00eo° = Abdominal Segments Seta Cephalothorax I II Il IV Vv VI VII VIII a 0 = = 1 1 iee 2) s0=19" CM) 3-65) 1-44) me 225.6) | 1 Ey Be i=?) 1 1 1-4 (1) As 2-612) 520?) 4-14), 223-0) 50) 64=8)G) Ot Dt) 23-456)" 41-3413) 6 1 3°8.66)' 9 328.4)’ 3-6-6) 72 TIED) DEA (A) EPO)" 1-43) Bilt 2=51(2) = 2 2-6 (4) 9 344 ! 1 1 10, 62514) = I4(1) 24 (4) 11 1 = 1-3 ©) I 1209 3-7) — — -- 14 _ _ Zs 1 ] l 1 1 l 3-5 (4) 3-6(4) 3-6 (3) 1-4 (2) — 1 1 1 ] ~ = 4-6 (6) 47 (7) 1-6 (4) 3-5 (3) = 12, @) 4110902), 2956) 2-6 (5) 3-5 (4) l 1 1-4 (3) 2-3 (3) = 3-4 (4) 3-5(5) 46 (4) 2-6 (5) — 1-3 (2) 3-8 (6) 2-7 (?) 3-7 (6) — 13 (3) 230) 2-46) 5-8 (7) — ] 1 1 9-12 (10) 9-15 (10) 24 (3) 2-3 (3) 1-4 (3) 2-4 (2) = ] 1 1 1-3 (2) == l l 1 l l ' Range followed in parentheses by the mode. ble 3. Cephalothorax: Setae 1, 3-CT long, stout, | single, 3 single or double; 7-CT 0.55—-1.6 length of 6-CT, usually double; trumpet length 0.42—0.54 mm, pinna 0.04— 0.22 total length, index 1.68—2.25. Abdo- men: Seta 1-II small with thin branches, 2- to 7-branched; 3-II 3.0—4.23 longer than 2- II, 3-II 1.03—1.15 length of tergum III; 3- III 1.04-1.19 length of tergum IV; 5—IV 0.96—-1.10 length of tergum V; 5-V 0.92-— 1.10 length of tergum VI; 6-VI single, stouter than 5-V; 9-VII stout, aciculate, usu- ally double (2—4); 9-VIII stout, aciculate, with 6—9 b, some branches of 9-VII, VIII divided apically. Paddle: ovoid, slightly emarginate apically, length 0.85—1.10 mm, index 1.24—1.46; fringe on outer margin on apical 0.43—0.60, inner margin on apical 0.40—0.66; fringe length 0.12—0.16 mm; seta 1-P strong, 0.23—0.40 length of paddle. The description is based on the following specimens: Singapore Island, 1970?, 1°, 13 LePe, (Nos. 1200-10, 18); Brunei, Be- lait District 1970; 19, Pe,126 LePe. CNos: 1295-19, 1312-15, 1341-102). Nepal spec- imens have been collected Rupandehi Dis- trict, Bhairahawa, IV-7-88, 1d (Darsie and Pradhan 1990); Makwanpur District, Hetau- ra, VI-VII-55, 22; Nayagaon, I-V-55, 12 (Peters and Dewar 1955). Armigeres (Armigeres) subalbatus (Coquillett) (Fig. 4) Position and size of setae as figured; range and modal branching in Table 4. Cephalothorax: Setae 1, 3-CT stout, dou- ble; 7-CT 0.58—0.81 length of 6-CT; trum- pet length 0.36—0.50 mm, pinna 0.11—0.30 of total length, index 1.0—1.72. Abdomen: 1-II branched from base, 8—13 b; 3-II 1.63— 1.90 length of 2-II; 3-H 1.05—1.20 length of tergum III, 3-III 1.04—1.26 of tergum IV, 5- IV 1.32-1.39 of tergum V, 5-V 0.98—1.28 of tergum VI; 6-VI single, rarely double, thicker than 5-V; 9-VII with 3—7 b, some branches shorter; 9-VIII with 9—14 b, outer branches shorter. Paddle: ovoid, emarginate apically, length 0.93—1.17 mm, fringe of outer margin on apical 0.51—0.81, of inner margin, on apical 0.23—0.53; index 1.16— 1.35, fringe length 0.15—0.18 mm; 1-P sin- gle or seldom double, 0.33—0.43 length of paddle. The description is based on the following specimens from Nepal: Kaski District, Pok- hara, Male Patan, VII-27-92, 32 Pe, 1d Pe, ex treehole; Pokhara, VII-28-92, 1d Pe, ex bamboo stump; Pokhara, Lamachaur, VII- 29-92, 12 Pe, ex bamboo stump. 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 102, NUMBER 1 113 Table 2. Pupal chaetotaxy of Armigeres durhami. Abdominal Segments Seta Cephalothorax I I Ul IV Vv VI VI Vul 0) a — 1 1 ] l 1 l 1 1 l 9-11 (10) 17-26 (?) 2-5 (4) 2-6 (3) 2-6 (3) 1-5 (3) 14 (1) = D 2-5 (3)! 1 ] l ] 1 1 1 a 3 2 1 l 1 2-7 (5) 2-5 (4) 2-5 (3) 1-6 (2) = 4 2-5 (3) 5—8 (6) 5-8 (6) 2-5 (4) 24(4) 3-7(6) 2-5(2) 24 (3) 1+ (2) 5) 3-7 (5) 1-3 (1) 2-3 (2) 1-5 (4) I | 1-4 (1) 1-3 (3) — 6 | 2-4 (2) 24 (3) 2-5 (2) 2-4 (3) 1-4 (3) 1-3 (1) 1-4 (2) = i 2-5 (2) 3-5 (5) 4-7 (4) 2-5 (4) 3-5 (5) 47 (4) 2-4 (3) 1-2 (2) — 8 2-5 (3) = — 2-5 (3) 24 (3) 3-5 (3) 3-5 (3) 34 (3) = 9 2-6 (4) 1 l 1 | 1 l 3-4 (4) 7-12 (9) 10 5-8 (5) — 2-3 (2) 2-5 (3) 24 (3) 2-3 (3) 2-3 (2) 1-3 (1) oa 11 l — — 1 | l 1-2 (2) 1-2 (2) 1 3-5 (4) — oa — — -= — — — 14 _- oo — ] l l ] 1 l ' Range followed in parentheses by mode. Armigeres (Armigeres) baisasi STONE AND THURMAN (Fig. 5) Position and size of setae as figured; range and modal number of branches in Table 5. Cephalothorax: Seta 1-CT and 3- CT long, stout, 1 single, 2 usually double; 7-CT 0.54—0.82 length of 6-CT; trumpet length 0.63—0.75 mm, pinna 0.22—0.44 to- tal length; index 1.96—2.27. Abdomen: Seta 1-II pedunculate, with 3—4 main stems, each with 2-8 branches in apical 0.14-— 0.43, totalling 10-18 b; 3-II 1.59—1.83 longer than 2-II, 0.95-1.07 length of ter- gum III; 3-II 0.96—1.11 length of tergum IV; 5-IV 1.14—1.34 length of tergum V; 5- V 0.89-1.38 length of tergum VI; an ad- ventitious seta marked (A) found in one specimen just anterior to seta 5-V, see Fig. 5; 6-VI aciculate, single or double; 9-VII aciculate, with 2-5 b; 9-VIII aciculate, with 6-8 b. Paddle: ovoid, emarginate api- cally, length 0.90—1.24 mm, index 1.06— 1.37, fringe on apical 0.32—0.68 of outer and on 0.36—0.52 of inner margins, fringe length 0.13 mm; 1-P strong, 0.26—0.31 length of paddle. The description is based on the following specimens from the Plilippines: Palawan, Puerto Princesa, [X-20-45, 1d LePe, ex co- conut shell (No. P-307, 19th MGL); Davao, Lasang, V-7-46, 12 LePe, ex coconut husk (No. P-945-3) (J. Enke and H. Hoogstraal); without locality data, 12 LePe (No. 2010- >): KEY TO THE KNOWN PUPAE OF THE SUBGENUS ARMIGERES, Genus ARMIGERES Key formulated partly using key of Stef- fan (1968), descriptions by Ramalingam (1972, 1987), Toma et al. (1994, 1995) and descriptions herein. Ie Paddle without seta; seta 7-CT shorter thant SetaiG=CI, 6 cc y.4.6 He cde ar Abie ee ewe 2 — Paddle with strong seta; seta 7-CT sube- qual to, or longer than, 6-CT Setae 3-II, HI and 5-IV, V shorter than 0.25 length of following tergum ...... theobaldi Barraud - Setae 3-H, III and 5-IV, V subequal to length of following tergum SL ioe: alkatirii Toma, Miyagi and Syafruddin 3(1). Paddle fringe on external margin extend- Ime to neéaribasew © 210m. @ ees eee 4 ~ Paddle fringe on external margin on apical OWSrOrdess as i 3.5 cats ee Be eee 5 4(3). Seta 1-P usually triple; seta 6-VI mostly with more than 2 branches malayi (Theobald) = Seta 1-P usually double; seta 6-VI usually 114 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 102, NUMBER 1 LS Table 3. Pupal chaetotaxy of Armigeres kuchingensis. Abdominal Segments Seta Cephalothorax I II Hl IV Vv VI VIL VI 0) —_ — 1 1 1 l l 1 1 l 8=85(6) =, 2=7'G) —5 (4) 2-5(3) 24(3) 2-6(4) 2-5 (3) = 2 1-3 (3)! 1 l 1 l 1 1 l = 3 1-2 (1) 1 1 1 4-8 (4) 24(2) 1-3(3) 2-3 (2) — 4 2-3 (3) 4-95) A-6\(6) - 2-353) 2-4@) 3-6@ 1536) 2-3.) 223 @) 5 24 (3) 137), 1-272) 1-311) 1 l 1-3 (2) ~=1-3 (1) = 6 ] 3-7 (3) 2-8(3) 2-8(3) 24(3)_ 1-6(3) ] 1=5 (2) = 7 e=3"(2) 1-9 (2) 3-7(4) =2-8(?%) 24() 3-6(65) 2-3(@) 1-2(@) — 8 24 (3) — 34 (3) 3403) 244) 3464) | 3-6(3) —= 9 243) 123) 1 i 1 l 1 24 (2) 6-9 (8) 10 2-4 (3) — 2-Ai(2) 2 2 24 (4) 1-3() 14 (1) — 11 ] — 1+ (3) 1-3 (1) 2d) e@) 12@) 1Q) = 12 1-4 (2) == = = = a = = = 14 — — — 1 1 1 l 1 ' Range followed in parentheses by mode. single or double, rarely triple).........-. 10(9). Seta 1-If small, branches thin, 0.36 or less By ike Capen artes aes breinli (Taylor) length of seta 3-II ... kuchingensis Edwards 5(3). Seta 1-CT with branches thin, usually = Seta 1-II large, branches strong, 0.5 or double; seta 9-VII with 9 or more branch- more length of seta 3-II ........... 11 CS tren rare 2c aureolineatus (leicester): 7710). ‘Seta O-VIl with some. branches split be- yj pe Cl Sn, Stout: siuzle Or Geuble, yond middle; seta 2-I about 0.7 length of Sete an or sewer Pranehes 222 6 Sl Beet ee See eels ware lenbesteni Brug 4 ae 5 6(5) Seta 1-CT with 2 or more branches; seta ; : : 3-CT with 3 or more branches; seta 1-II a Shee fag sp ote sample tee less than 0.5 length of 3-II ........... about.0:5 or less length of 3-1) % s4o.% 12 Aiobee Crttach crichton hore op Cuca crrekt papuensis Peters 12(11). Seta I-IT pedunculate, with 2-3 main = Seta 1-CT mostly single; seta 3-CT single stems, branched in apical 0.7-0.9 .. baisasi or double; seta 1-IH 0.5 or more length of Stone and Thurman S=lIK(EXcept ‘KUCGHINGENSIS) .. 3.25%. = 3s i, Se Seta 1-II with single main stem or sub- 7(6). Seta 1-I] pedunculate, brush-like, with sin- pedunculate, usually branched from near gle main stem, branched at least in apical BASE 5.95 soos es Sse some owes Stan cue ts Geka 13 OS Rade Or aes = Whee yes ered os 8 13(12). Trumpet index 1.7 or less, length 0.5 mm % Seta I-If branched from Deak base, if OF less, 4,.hoale Pee subalbatus (Coquillett) SOE OES DS MUO nO ae = Trumpet index 2.3 or more, length 0.51 Se geese gies sep ae ws ey Oe a as 2 MUM OL AMOTEM Gs ow ako 5 sic ions iy ache rae 14 8(7). Seta 10-VI usually double; trumpet length : OSSimmysees as kinabaluensis Ramalingam ECT pe 7-8 Myatt branches: sce We ~ Seta 10-VI usually single; trumpet length less than 0.5 length of ©-CT;, pina 0-5 0.65 or greater ......... iphan, Bawards length of trumpet ..2..«.< milnensis Lee 9(7). Seta 7-CT generally 3-branched; seta 6-VI ‘a Seta 9-VII with 5 or more branches; seta no stouter than 5-V ..... lacuum Edwards 7-CT more than 0.5 length of 6-CT; pinna ~ Seta 7-CT usually single or double; seta 0.4 length of trumpet .. kesseli Ramalingam 6-Vilt'stouten thane5-W! Aes = eles ae 10 C= Fig. 3. Pupa of Armigeres kuchingensis. A, Cephalothorax. B, Metanotum and abdomen. For abbreviations, Sees ione 116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Pupa of Armigeres subalbatus. A, Cephalothorax. B, Metanotum and abdomen. For abbreviations See bilcanille VOLUME 102, NUMBER 1 117 Table 4. Pupal chaetotaxy of Armigeres subalbatus. Abdominal Segments Seta Cephalothorax I II Il IV Vv VI Vil Vill 10) — — 1 | l | 1 l 1 2 8-12 (9) 8-13 (9) 2-6(4) 47(5) 47(6) 24(3) 23) — 2. 2—4 (2)! 1 l l l l 1 l — 3 2 1 1 | 3-6 (4) 2-5(4) 24(2) 2-5 (2) 2-3 (3) 4 2-4 (4) 4-9 (5) 48(6) 24(3) 2-5(3) 3-6(4) 2-6(4) I+ (2) = 5 3-5 (4) 1 (3) 1-3 (2) 1—3:(3) 1-2 (1) 121) 24() 1=3"(@) = 6 ] 3-7 (6) 3-8(4) 3-6(4) 3-6(?) 2-6 (5) 12 (1) 2-4@) = il 2-3 (2) 2-6 (3) 476 354 364 476 %35@4 1—2 (2) = 8 1=3.,(3) = = 3-4 (3) 2-5(3) 340) 3-503) 2-6(3) 9 2-5 (3) 1—3 (2) l | 1 1 1 3-7 (?) 9-14 (10) 10 2-4 (4) = 14 (3) 2-5(3) 2-3(3) 2-33) 1-3 (2) 1—3 (2) 11 | — — l l l 1-3 (2) = 2) 34 (3) = == —— = = —— = — 14 — — — l ] 1 l ' Range followed in parentheses by mode. PUPAL CHARACTERS FOR THE GENUS AND SUBGENERA Genus Armigeres presently has 48 spe- cies, 33 in subgenus Armigeres and 15 in subgenus Leicesteria. Pupae have been ad- equately described for 15 species of sub- genus Armigeres and 7 of subgenus Leices- teria (including Ar. omissus, description pending). These 22 descriptions offer an opportunity to formulate tentative pupal characters to distinguish the genus and sub- genera. Belkin (1962), Definado (1966), Thurman (1959), and Mattingly (1971, by virtue of his generic key to pupae), have mentioned generic characters. The former three mention that they resemble Aedes pu- pae and that they have a paddle fringe. Only Belkin also noted the following: seta 8-C caudad of the trumpet base, setae 5-VI, VII very small and 4-VIII short, multiple. Genus.—The trumpet is normal, 0.36— 0.71 mm in length, pinna rather short, trun- cate apically; seta 6-CT long, stout, usually light tan; 8-CT not longer and stouter than 9-CT, located well posterior to base of trumpet; 5-III not longer than following segment; 5-IV-V simple, not aciculate; 5- VI, VII very small; seta 6-VI well devel- oped, usually stouter than 5-V; seta 9-VII long, stout, usually branched; 4-VIII single or branched; segments IX and X without setae; paddle usually emarginate apically, with long fringe on outer and inner mar- gins, with or without seta 1-P, when present, less than 0.5 length of paddle. Subgenera.—Pupae of subgenus Armiger- es have seta 6-CT usually as long as or lon- ger than 7-CT; 3-CT and usually 1-CT long and very stout; seta 1-P mostly present and at least 0.3 length of paddle. In subgenus Leicesteria seta 7-CT at least 1.6 length of 6-CT; 1, 3-CT usually slender; 1-P absent or if present, no longer than fringe spicules. ACKNOWLEDGMENTS The author is indebted to G. W. Courtney and S. P. Pradhan for their collaboration in the project; to the National Geographic So- ciety for support in all aspects of the study; to T. Gaffigan and the Walter Reed Biosys- tematic Unit and Dr. S. Ramalingam for loan of specimens, to B. Bower-Dennis for preparing the illustrations, and to J. R. Rey and J. K. Nayar for reviewing the manu- script. This is Florida Agricultural Experi- ment Station Journal Series No. R-06298. LITERATURE CITED Baisas, F E. 1974. The mosquito fauna of Subic Naval Reservation, Republic of the Philippines. U.S. 118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Boule Bowler-enviz, x Fig. 5. Pupa of Armigeres baisasi. A, Cephalothorax. B, Metanotum and abdomen. For abbreviations see Eigaele VOLUME 102, NUMBER 1 119 Table 5. Pupal chaetotaxy of Armigeres baisasi. Abdominal Segments Seta Cephalothorax I II II IV Vv VI VII VIII 0) — — 1 1 1 1 1 1-3 (1)! 9-13 (12) 10-18 (2) 24 (3) 2-3(3) 34(4 2-363) 1-2 (1) 2 2-4 (3) 1 1 ] 1 1 1 1 —_ 3 1-2 (2) 1 1 1 1-5 (4) 3-5 (3) 2-4 (4) 2-4 (4) — 4 2-6 (?) 3-7 (?) 3-6 (4) 1-6 (6) 1-5 (1) 3-7(6) 3-6(4) 34(@3) 2-4 (?) 5 3—6:1(3) 2-4 (2) f=) (S51 1-3") 1 1 1-4 (?) 2 — 6 1 3-6 (3) 3-4 (4) 14 (3) 34 (4) 34 (3) 12(1) 24 (2) — ih 1—3 (2) 3-5 (5) 47 (4) 2-5(3) 24 (4) 3-6(4) 3-5 (3) 2 — 8 3-4 (7) = I2#(1) 45 (4) 3-5(3) 2-3(3) 2-5(3) 3-6 (4) = 9 2-4 (3) 1—2 (2) 1-3 (3) 1 1 1 I 2-5 (7?) 6-8 (?) 10 3-6 (5) = 1-3 (3) + 2-3(3) 2-3(2) 2-3 (3) 1-5 (?) 1-3 (2) — 11 l _ — 1-3 (1) l 1 1-2 (?) 1-3 (2) = 12 2-4 (?) — — — — — — _ — 14 aad = _ 1 l 1 l l 1 ' Range followed in parentheses by mode. Navy, First Medical Service Wing, Technical Re- port 72-2, 170 pp. Barraud, P. J. 1934. Family Culicidae. Tribes Megar- hinini and Culicini. The Fauna of British India, Including Ceylon and Burma. Diptera, Vol. V, London, Taylor and Francis, 473 pp. Belkin, J. N. 1962. The mosquitoes of the South Pa- cific (Diptera, Culicidae). Vols. I, 608 pp.; II, 412 figures. Berkeley, University of California Press. Belkin, J. N., R. X. Schick, P. Galindo, and T. H. G. Aitken. 1965. Mosquito studies (Diptera, Culici- dae) I. A project for a systematic study of the mosquitoes of Middle America. Contributions of the American Entomological Institute (Ann Ar- bor) 1(2): 1-78. Bohart, R. M. and R. M. Ingram. 1946. Mosquitoes of Okinawa and Islands in the Central Pacific. United States Navmed 1055, 110 pp. Borel, E. 1930. Les moustiques de la Cochinchina et du Sud-Annam. Collections Society Pathologique Exotique, Monograph 3: 1-123. Darsie, R. FE Jr. 1998. Description of the pupae of six species in subgenus Leicesteria, genus Armigeres, from Nepal (Diptera, Culicidae). Proceedings of the Entomological Society of Washington 100: 234-246. Darsie, R. F, Jr. and S. P. Pradhan. 1990. The mos- quitoes of Nepal: Their identification, distribution and biology. Mosquito Systematics 22: 69-130. Delfinado, M. D. 1966. The culicine mosquitoes of the Philippines, tribe Culicini (Diptera, Culicidae). Memoirs of the American Entomological Institute 7: 1-152. LaCasse, W. J. and S. Yamaguti. 1948. Mosquito fauna of Japan and Korea. Corps of Engineers, United States Army, 273 pp. Mattingly, P. FE 1971. Contributions to the mosquito fauna of Southeast Asia. XII. Illustrated keys to the genera of mosquitoes (Diptera, Culicidae). Contributions of the American Entomological In- stitute 7(4): 1-83. Penn, G. H. 1949. The pupae of the mosquitoes of New Guinea. Pacific Science 3: 3-85. Peters, W. and S. Dewar. 1955. A preliminary record of the megarhine and culicine mosquitoes of Ne- pal with notes on their taxonomy (Diptera: Culic- idae). Indian Journal of Malariology 10: 37-51. Ramalingam, S. 1972. A new species of Armigeres from Sabah, Borneo (Diptera: Culicidae). Pro- ceedings of the Entomological Society of Wash- ington 74: 459-467. . 1987. On the restriction of Armigeres durhami Edwards and the description of Armigeres kesseli n. sp. (Diptera: Culicidae). Tropical Biomedicine 4: 55-65. Steffan, W. A. 1968. Armigeres of the Papuan Subre- gion (Diptera: Culicidae). Journal of Medical En- tomology 5: 135-159. Thurman, E. B. 1959. A contribution to a revision of the Culicidae of northern Thailand. University of Maryland Agricultural Experiment Station Bulle- tin A-100, 182 pp. Toma, I., I. Miyagi, and N. Benjaphong. 1994. Rede- scription of Armigeres (Armigeres) theobaldi (Diptera: Culicidae). Mosquito Systematics 26: 11-18. Toma, T., I. Miyagi, and Syafruddin. 1995. Description of Armigeres (Armigeres) alkatirii, a new species from Sulawasei, Indonesia (Diptera: Culicidae). Mosquito Systematics 27: 110—117. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 120-128 COMPARATIVE STUDY OF SPERMATHECAE IN ELEVEN RHAMMATOCERUS SAUSSURE 1861 GRASSHOPPER SPECIES (ORTHOPTERA: ACRIDIDAE: GOMPHOCERINAE: SCYLLININI) CRISTIANE VIEIRA DE ASSIS-PUJOL AND MICHEL LECOQ (CVAP) Museu Nacional, UFRJ, Departamento de Entomologia, Quinta da Boa Vista, Sao Crist6vao, 29.940-040, Rio de Janeiro, Brazil (e-mail: cpujol @ acd.ufrj.br); (ML) Cen- tre de coopération internationale en recherche agronomique pour le développement, Prifas, BP 5035, 34032, Montpellier Cedex 1, France (e-mail: lecog @cirad.fr) Abstract.—The spermathecae of 11 Rhammatocerus Saussure 1861 grasshopper species are described [R. brasiliensis (Bruner 1904), R. brunneri (Giglio-Tos 1895), R. cyanipes (Fabricius 1775), R. guerrai Assis-Pujol, 1997, R. palustris Carbonell 1988, R. pictus (Bruner 1900), R. pratensis (Bruner 1904), R. pseudocyanipes Assis-Pujol 1997, R. schis- tocercoides (Rehn 1906), R. suffusus (Rehn 1906) and R. viatorius viatorius (Saussure 1861)]. This taxonomic character, which has never been assessed in the tribe Scyllinini, is critical for determining grasshopper species. It also may be helpful in a taxonomic reclassification of species and provide a basis for reviewing the genus Rhammatocerus and the entire tribe Scyllinini. Key Words: In most taxonomic analyses of acridians, the phallic complex is the main morpholog- ical character used to distinguish species (Dirsh 1956, 1961). However, this character sometimes proves impracticable, e.g., for diagnosing some Gomphocerinae genera (Carbonell 1995). In such cases, the female genitalia, especially the spermatheca, may provide a basis for determination. Slifer (1939) described this distinguishing feature for many acridian species, and to a lesser extent by Dirsh (1957) and other authors, but these studies are almost solely descrip- tive. Spermatheca design has never been used as a basis for a review of current tax- onomic groups. Nevertheless, as early as 1939, Slifer (1939) stated that the sperma- theca would be a relevant criterion for de- termining families and subfamilies. Uvarov (1966) point out that the distal part of the spermathecal duct is subject to considerable Acrididae, Scyllinini, Rhammatocerus, spermatheca variations, characteristic for some taxonom- ic groups. Amedegnato (in litt.) noted that the spermatheca and diverticula are useful diagnostic characters and could even pro- vide a basis for a phylogenic reclassifica- tion. However, Dirsh (1957) regard the spermatheca as a subsidiary taxonomic character of the higher categories to be used with caution. In fact, very few studies have been carried out to assess the potential rel- evance of spermatheca design for differen- tiating species within an Acrididae genus. Grasshoppers of the genus Rhammato- cerus Saussure 1861 are distributed from southern USA to Argentina (Otte 1981). This group is characterized by high intra- specific variation and a relatively hetero- geneous external morphology. The econom- ic impact of these grasshoppers has in- creased markedly over the last 15 years in Brazil and other South American countries, VOLUME 102, NUMBER 1 especially in Colombia (Lecoq and Assis- Pujol 1998), where they devastate many different crops and pastures (Cardenas and Devia 1995, Guagliumi 1958, 1959, 1962, 1973, Carbonell 1988, Lecoq and Balanga 1998, Lecoq and Pierozzi Jr 1994, 1995a, b, Lecog et al. 1997, Leén 1996, Martinez and Gomez 1996, Miranda et al. 1996). Very few taxonomic studies have been carried out on Rhammatocerus. Since this genus was first described the most note- worthy studies on the genus were by Jago (1971) and Carbonell (1995), but no taxo- nomic revisions were involved. There are currently 13 species included in the genus (Carbonell 1995, Assis-Pujol 1997a, b, c, 1998, 1999). Most of them are known only from the original descriptions, which are generally ill-defined and based on charac- ters that cannot be readily used or com- pared. Moreover, some studies have been carried out without reference to previous ar- ticles on the genus, which has led to con- siderable synonymy, erroneous species identifications and misinterpretations of geographical distributions. These latter spe- cies are currently distinguished on the basis of pigmentation features, which is a serious constraint for accurate identifications, 1.e., pigmentation is often unclear on type spec- imens, due partly to poor preservation con- ditions. A critical problem with using color is also that it varies within a species as shown clearly in R. schistocercoides for which color change from brown to green with sexual maturation (Lecoq and Pierozzi Jie, 1996): As part of a general project to revise Rhammatocerus (Assis-Pujol 1997a), we at- tempted to pinpoint relevant diagnostic cri- teria for species identification. It was found that interspecific morphological differences in male genitalia are so minor, or even non- existent, that they are unsuitable for species determination (Assis-Pujol 1997a). How- ever, internal female genitalia, especially the spermatheca, were found to be a critical taxonomic character. Here we present a comparative analysis of the spermatheca. MATERIALS AND METHODS Spermathecae of 11 of the 13 species currently included in Rhammatocerus were studied. Specimens of R. varipes (Bruner 1905) and R. alticola (Hebard 1923) were not available. The 264 female specimens analyzed were obtained from the following institutions: Facultad de Ciencias Naturales y Museo, La Plata, Argentina (FCNM); Museu Nacional do Rio de Janeiro, Brazil (MNRJ); Universidade Federal de Pernam- buco, Brazil (UFPE); Universidade Federal Rural de Rio de Janeiro (UFRRJ); Colecao Entomoldgica Costa Lima, Universidade Federal Rural do Rio de Janeiro, Brazil (CECL); and Facultad de Ciencias, Univ- ersidad de Montevideo, Uruguay (FCUM). A detailed list of specimens studied is given for each species. We adopt the terminology of Slifer (1939). Spermathecae were dissected and carefully prepared according to the follow- ing technique. The specimens were placed in a wet chamber for 24—36 h. Once soft- ened, the abdomen was cut off at the 6th segment and a longitudinal incision was made along the intersegmental abdominal membrane. The spermatheca was removed and incubated in a 10% potassium hydrox- yde solution (KOH) for 24 h. It was then washed to remove all KOH in a solution containing 10 ml acetic acid/100 ml 70% alcohol. All tissues surrounding the sper- matheca were removed, and it was then washed and incubated for about 24 h in a Mercurochrome/70% alcohol solution (1:10 v/v) in order to stain the structures. The spermatheca was washed again in water and placed between a glass slide and a coverslip in a drop of glycerine. All spermathecae il- lustrations were made with a camera lucida under a dissecting microscope (50). After dissection, the abdomen was dried and put with the specimen. The genitalia were placed in a microtube in glycerine along side the specimen after the analysis. RESULTS AND DISCUSSION The general morphology of Rhammato- cerus spermathecae was found to be in line 122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with that described by Slifer (1939) for Ac- rididae spermathecae. All of them have a well-developed preapical diverticulum (PD) and a distinct apical diverticulum (AD) (Figs. 1-11). Spermatheca of the different species studied are described below. Rhammatocerus brasiliensis (Bruner 1904) Material—BRASIL, PARA: Santarém, V.1978, M. V. Cerdeira, 3, MNRJ; bidos, IV.1978, B. Silva, 1, MNRJ; Rio Grande do Norte: Jancico, [.1952, M. Alvarenga, 1, MNRJ; PARAIBA: Brandao, Santa Luzia, O8.XII.1955, A. G. A. Silva, 2, MNRJ; Joazeirinho, Soledade, VII.1956, A. G. A. Silva, 1, MNRJ; PERNAMBUCO: Arari- pina, VI.1993, M. J. S. Lopes & C. S. Car- bonell, 1, FCUM; Bonito, I.1978, B. Silva, 2, MNRJ; Serra das Russas, BR 232, Km j5a(cerca ‘dexnGravat)y) VIEI9932 "MiMi: S: Lopes & C. S. Carbonell, 3, UFPE; 8 Km E de Sanharo, 30.IV.1991, C. S. Carbonell & M. J. S. Lopes, 14, UFPE; ESPIRITO SANTO: Cariacica, XII.1977, B. Silva, 1, MNRJ; MINAS GERAIS: Diamantina, XJI.1977, Seabra, Roppa & Monné, 3, MNRSJ; Aguas Vermelhas, X.1977, O. Rop- pa, 5, MNRJ; Aguas Vermelhas, Curral de Dentro, XI.1977, O. Roppa, 1, MNRJ; Cur- velo, VII.1977, Seabra, Roppa & Monné, 1, MNRJ; Pirapora, XI.1977, Seabra, Roppa & Monné, 2, MNRJ; GOAIS: Minagi, 29.V-04.VI.1987, Monné & Roppa. 1, MNRJ; Mineiros, II.1979, Roppa & Silva, 1, MNRJ; SAO PAOLO: Piracicaba, 19.X.1961, A. Mesa, 2, FCUM.; Piracicaba, Volta Grande, 4.V.1962, A. Mesa, 2, FCUM;; Indaiatuba, 22.XI.1961, A. Mesa, 2, FCUM; Mato Grosso: Diamantino, BR 364, Km 348, II.1978, B. Silva, 1, MNRJ; MATO GROSSO DO SUL: Campo Grande, 08.XI.1952, M. Alvarenga, 1, MNRJ; RIO GRANDE DO SUL: Traman- dai, 14.11.1964, C. S. Carbonell, A. Mesa & M. A. Monné, 2, FCUM; PARAGUAIT, CENTRAL? Wuque; 03: 1965; (Cy S:"Car- bonell, A. Mesa & M. A. Monné, 1, FCUM; PARAGUARI: Paraguarf, 10.111.1965, C. S. Carbonell, A. Mesa & M. A. Monné, 8, FCUM; CAAGUAZU: 40 Km N de Caaguazu, Ruta Caaguazt, Yhu, 13.11.1965, C. S. Carbonell, A. Mesa & M. A. Monné, 3, FCUM; ARGENTINA, SAL- TA: San Lorenzo, 20.1.1965, A. Mesa & R. Sandulski, 1, FCUM; P. del Carmen, 22.11.1936, E Langman col., 1, FCNM; BUENOS AIRES: San Nicolas, 15- 20.1.1949, Sonia Rivas leg., 2, FCNM; SANTIAGO DEL ESTERO: Beltran, 8.X1.1941, Maldonado col., 1, FCNM. Description of spermatheca.—Short preapical diverticulum, about 1.6 mm in length; apical diverticulum short, about % as long as preapical diverticulum, with its tip oriented in opposite direction relative to that of former structure (Fig. 1). Rhammatocerus brunneri (Giglio-Tos 1895) Material.— BRASIL, GOIAS: Minagcu, 7.X.1987, Monné & Roppa, 1, MNRJ. SAO PAULO: Serra da Bocaina, 1660 m, S. J. Barreiro, IV.1978, Seabra, Monné & Rop- pa, 1, MNRJ. Mato Grosso: Chapada, mar- ¢o, Bruner cln, Ex. Carn. Mus., A.N.S.P., 1, FCUM. BR 364, KM 616, II.1978, B. Silva, 1, MNRJ; MINAS GERAIS: Varginha, 11.1955, E M. Oliveira, 1, UFRRJ; RIO GRANDE DO SUL: 5, 33 km N de Passo Fundo, 25.I].1964, C. S. Carbonell, A. Mesa & M. A. Monné, 1, FCUM; Pelotas, 14.11.1962, C. Biezanko, 1, FCUM; Santa Maria, 5.1.1955, E. Corseuil, 02676, 1, FCUM;; Ronda Alta, 24.11.1964, C. S. Car- bonell, A. Mesa & M. A. Monné, 3, FCUM; Tramandai, 14.11.1964, C. S. Car- bonell, A. Mesa & M. A. Monné, 1, FCUM. URUGUAI, LAVALLEJA: Alredores Pueblo Sauce, 22.11.1953, C. S. Carbonell, 1, FCUM; RIVERA: Cuchilla Cunapiru, 21.1.1956, C. S. Carbonell, 4, FCUM; Si- erra de La Aurora, 14.III.1961, C. S. Car- bonell, A. Mesa & P. San Martin, 3, FCUM. PARAGUAI, CAAGUAZU: Caaguazu, 12.111.1965, C. S. Carbonell, A. Mesa & M. A. Monné, 1, FCUM; 40 Km N de Caa- guazu, Ruta Caaguazu, Yhu, 13.]]1.1965, C. S. Carbonell, A. Mesa & M. A. Monné, 4, VOLUME 102, NUMBER 1 Dy F Figs. 1-11. Spermathecae. 1, Rhammatocerus brasiliensis. 2, R. brunneri. 3, R. cyanipes. 4, R. guerrai. 5, 11, R. palustris. 6, R. pictus. 7, R. pratensis. 8, R. pseudocyanipes. 9, R. schistocercoides. 10, R. suffusus. viatorius viatorius. Abbreviations: DA = apical diverticum; DP = preapical diverticulum. 124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON FCUM; Sapucay, 26.XII.1904, Foster, 1, FCUM; 26.1.1905, Foster, 1, FCUM. AR- GENTINA, MISIONES: Caraguatay, 1.1960, Ronderos & Trotta col., 1, FCNM; CHACO: Resistencia, 20.III.1939, Denier cols aL, FCNM: Description of spermatheca.—Long preapical diverticulum, about 2.0 mm in length; very short rounded apical divertic- ulum (less than %4 length of preapical di- verticulum) with its tip oriented towards posterior part of spermatheca (Fig. 2) (if we consider anterior region with preapical di- verticulum on right side of observer and apical diverticulum on left side, as indicated on Figs. 1-11). Rhammatocerus cyanipes (Fabricius 1775) Material—_COLOMBIA, SANTA MAR- TA: Valencia (Alt. 500 ft.), O08. VIII.1920, FE M. Gaige col., 2, FCNM. Description of spermatheca.—Very short preapical diverticulum, about 1.0 mm in length; long apical diverticulum (if com- pared with preapical diverticulum), with its tip oriented slightly in opposite direction relative to that of former structure (Fig. 3). Rhammatocerus guerrai Assis-Pujol 1997 Material.—Paratype: 1, BRASIL, MATO GROSSO: Campo Novo do Parecis, 08.V.1996, C. Pujol, W. Guerra & O. Ri- beiro, MNRJ. Description of spermatheca.—Short preapical diverticulum, about 1.6 mm in length; very long, thin and sinuous apical diverticulum about half as long as preapical diverticulum, with its tip oriented in oppo- site direction relative to that of former structure (Fig. 4). Rhammatocerus palustris Carbonell 1988 Material. BRASIL, MATO GROSSO: Chapada dos Guimaraes, 25.1.1972, Ron- deros & Carbonell, 2, FCNM:; GOIAS: Minaci, M. A. Monné e O. Roppa, 1, MNRJ; MATO GROSSO: Chapada dos Guimaraes, Fevereiro, 24-25, 1972, M. Descamps, 2, MNRJ; same locality, July, 1983, O. Roppa & M. A. Monné, 1, MNRJ; GOIAS: Jatai, MNRJ. PARAGUAI, CAA- GUAZU: 40 km N. de Caaguazu (at present Ihu), Marco, 1965, C. S. Carbonell, A. Mesa & M. A. Monné, 2, MNRJ. Description of spermatheca.—Long preapical diverticulum, about 2 mm in length; long apical diverticulum slightly shorter than half of preapical diverticulum, with its tip oriented slightly in opposite di- rection relative to that of former structure (Rigs): Rhammatocerus pictus (Bruner 1900) Material. BRASIL, RIO GRANDE DO SUL: Lagoa Vermelha, 18.II.1964, C. S. Carbonell, A. Mesa & M. A. Monné, 3, FCUM; Nonoai, 20.11.1964, C. S. Carbo- nell, A. Mesa & M. A. Monné, 1, FCUM; Ronda Alta, 24.11.1964, C. S. Carbonell, A. Mesa & M. A. Monné, 2, FCUM; Palmeira das Missies, II. 1978, O. Roppa, 1, MNRJ. URUGUAIT, ARTIGAS: Arroyo Tres Cru- ces Grandes (Potrero Sucio), 12, 14.11.1955, Fac. de Hum. Y Cienc., 5, FCUM; Arroyo Tres Cruces, Timbauba, 14.11.1955, FE de H. Y Ciencias; 2; FECUM:; Bellay Union: 281.0952, Li Ca Zolesst, 3, FCUMs Yacares 21.1.1952; EB de H. Y-Ciencias. 125FCUNME Canelones: Las Piedras, 20.III.1964, 1, FCUM; Las Piedras, 5, 6.II.1966, A. Car- menes, 2, FCUM; Las Piedras, 15.11.1966, A. Carmenes, 2, FCUM; Lavalleja: Cercan- ias Pueblo Sauce, 22.11.1953, C. S. Carbo- nell, 2, FCUM; Paysandu: Puerto Pepeaji, IV.1954, C. S. Carbonell, 1, FCUM; Rivera: Cerro Chato Dorado, 23.III.1963, A. Mesa & €. S. ‘Carbonell, 1, FE@UM: Guchilla Cunapiru, 210.1956, C, Ss Carbonell; FCUM;; Sierra de La Aurora, 14.11.1961, C. S. Carbonell, A. Mesa & P. San Martin, 1, FCUM. PARAGUAIT, San Bernardino: 14.1.1939, Denier col., 1, FCNM. ARGEN- TINA, SANTIAGO DEL ESTERRO: Bel- tran 8.XI.1941, Maldonado col., 1, FCNM; BUENOS AIRES: sd, J. Bosq col., 2, FCNM; 5.1.1907, P. Jorgensen, 1, FCNM; La Plata; sd, sc, 1, FCNM; Campo del Ci- clo, 1.1954, sc, 1, FCENM; CATAMARCA: VOLUME 102, NUMBER 1 Andalgala, 8.III.1962, Torres & Ferreyra col., 1, FCNM; CHACO: Resistencia, 20.11.1939, Denier col., 4, FCNM:; COR- DOBA: Alta Gracia, I. 1937, M. Birabén, 2, FCNM; Cabana, 12.11.1937, M.Birabén col., 1, FCNM; Oncativo, 17.1.1959, Bira- bén & Scott leg., 2, FCNM; Rio Los Sauc- es, 28.11.1942, Maldonado col., 1, FCNM; Rio Tercero, 14.]J.1942, Birabén col., 3, FCNM; Tanti, III.1968, Bulla & Grosso leg., 1, FCNM; Corrientes: 15—30.11.1959, M. Birabén col., 1, FCNM; Manantiales, II.1946, Birabén col., 1, FCNM; 1- 10.111.1959, Birabén col., 1, FCNM; LA RIOJA: Famatina, 22.11.1959, Torres & Gardellawcol.. fl FENM= -JOUJUY: ‘Yala; I2AMISt9S9%) Birabén.é Scott leg., 1, FCNM;; Reyes, 11.I].1939, Birabén & Scott leg., 1, FCNM; La Pampa: 10 Km W de Rancul, 1.I11.1973, A. Mesa & E. Cabella, ia wMNEI MISIONES= Cerro, Cora, 18.11.1965, C. S. Carbonell, A. Mesa & M. A. Monné, 1, FCUM; Seis Misiones, 10.11.1990, Balatti col., 1, FCNM; Parada Seis, 11.11.1990, Balatti col., 1, FCNM; SALTA: Abra Santa Laura, 11.III.1939, Birabén & Scott leg., 2 j, FCNM; Ciudad de Salta, 19.1.1965, A. Mesa & R. Sandul- ski, 1, FCUM; Juramento, 14.III.1939, Bir- abén & Scott leg., 1, FCNM; San Lorenzo, 20.11.1965, A. Mesa & R. Sandulski, 1, FCUM;; San Lorenzo, 3.11.1965, A. Mesa & R. Sandulski, 5, FCUM; Tartagal, 29- 31.1.1965, A. Mesa & R. Sandulski, 1, EEUM:.- SAN) UIS:. .Piedray-Blanca, 19.11.1960, V. Sarmiento & Trotta, 1, FCNM,; Rincon del Este, Comechingones, 19.111.1960, V. Sarmiento & Trotta col., 2, FCNM; Concaran, 16—17.1II.1960, V. Sar- miento & Trotta col., 1, FCNM; Puerta Co- lorada, 14.1I].1960, V. Sarmiento & Trotta, 1, FCNM; SANTA FE: Barrancas, 22.11.1965, C. S. Carbonell, A. Mesa & M. A. Monné, 3, FG@UM:. Carcarana, 21.11.1965, C. S. Carbonell, A. Mesa & M. A. Monné, 7, FCUM; LA RIOJA: Vinchi- na, 22.11.1959, Torres & Gardella col., 4, FCNM. Description of | spermatheca.—Long 125 preapical diverticulum, about 2 mm in length; long apical diverticulum, bent at mid length, slightly shorter than half of preapical diverticulum, with its tip oriented in opposite direction relative to that of preapical diverticulum (Fig. 6). Rhammatocerus pratensis (Bruner 1904) Material— BRASIL, RIO GRANDE DO NORTE: Macaiba, VIHI.1951, M. Alvaren- ga, 1, MNRJ; PERNAMBUCO: 12 Km N de Buique, 30.X.1995, C. Ayres, 1, LGBE: 2422, UFPE; Serra das Russas, Km 116, 28.2%, 1995, C. “Ayres, 1, LGBE2019; UPFPE; 9 km U de Salsueiro, 3).X.1995, €. Ayres, 1, WRPE. Description of spermatheca.—Short preapical diverticulum, about 1.6 mm in length; straight apical diverticulum about % as long as preapical diverticulum, with its tip oriented almost in same direction rela- tive to that of former structure (Fig. 7). Rhammatocerus pseudocyanipes Assis- Pujol, 1997 Material.—Paratype 1, BRASIL, SER- GIPE: Areia Branca, 35 Km W de Aracaju, 19.11.1981, Roppa, Carbonell & Roberts, MNRJ; AMAZONAS: Humaitd, VIII. 1980, G. S. Andrade, 1, MNRJ; MINAS GERAIS: Aguas Vermelhas, X.1977, Sea- bra, Roppa & Monné, 1, MNRJ. Description of spermatheca.—Short preapical diverticulum, about 1.5 mm in length; short apical diverticulum about % as long as preapical diverticulum, with its tip clearly oriented in opposite direction rela- tive to that of former structure (Fig. 8). Rhammatocerus schistocercoides (Rehn 1906) Material— BRASIL, MATO GROSSO: Gustavo Dutra, 25.X.1953, C. R. Gongalves col., 1, UFRRJ; Serra do Roncador, Posto Pimentel Barbosa, 25.IX.1947, EK Meirelles col, 4, UFRRJ; Tangara da Serra, Dest. Al- cool Branca, 22.VI.1947, R. C. Mauro, 3, MNRJ; GOIAS: Aragareas, 2.VII.1953, H. Sick col., 1, UFRRJ; Margens do Rio Ar- 126 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON aguaia, 10.VII.1953, C. S. Carbonell, 2, FCUM; MINAS GERAIS: Sacramento, 30.1V.1996, G. W. Cosenza col., 3, MNRJ; Pedregulho, 30.1V.96, G. W. Cosenza col., 4, MNRJ. COLOMBIA, ARAUCA: Cravo Norte, 14.X.1996, M. Lecogq, J. A. Jimenez Gomez & O. J. Ramirez cols, 22, MNRJ. Description of spermatheca.—Long preapical diverticulum, about 2.0 mm in length; long bulky apical diverticulum about % as long as preapical diverticulum, with its tip clearly oriented in opposite di- rection relative to that of former structure (Fig. 9). Rhammatocerus suffusus (Rehn 1906) Material.— BRAZIL, AMAZONAS: Hu- maita, VIII.1980, G. S. Andrade, 1, MNRJ; RIO DE JANEIRO: Octobre, A.N.S.P, Ex. Carn. Mus., Bruner Cln, 1, FCUM; MINAS GERAIS: Corinto, XI.1977, Seabra, Roppa & Monné, 2, MNRJ; Curvelo, XI.1977, O. Roppa, 4, MNRJ; Gouveia, XI.1977, Sea- bra, Roppa & Monné, X.1982, L. Reys, 1, MNRJ; Minacu, 23.V—04.VI.1987, Monné & Roppa, 1, MNRJ; MATO GROSSO: Chapada near Cuyaba, May, June, August, AGNES: P Ex.'i€amsMusy) Bruner €in.; 2; FCUM. BOLIVIA, SANTA CRUZ: Santa Cruz de la Sierra, 26.1X.1953, M. Alvar- enga, 1, MNRJ. Description of spermatheca.—Short preapical diverticulum, about 1.5 mm in length; long thin apical diverticulum slight- ly longer than half of preapical diverticu- lum, with its tip oriented slightly in oppo- site direction relative to that of former structure (Fig. 10). Rhammatocerus viatorius viatorius (Saussure 1861) Material —MEXICO, SINALOA: Rosa- rio (J. A. Kusche), A.N.S.P, 1, FCUM; TE- COJA: Jatate, Rio Chiapas, 1800 ft, Iif.1934, D. W. Amram Jr., 1, MNRJ. GUA- TEMALA, LA UNION: Zacapa, 85 m., 10.X1.1972, E. Welling, 1, MNRJ. Description of spermatheca.—Short piri- form preapical diverticulum, about 1.6 mm in length; extremely large apical diverticu- lum, about *% of preapical diverticulum, scythe-shaped and its tip oriented in oppo- site direction relative to that of former structure (Fig. 11). These 11 species can be pooled into four different groups on the basis of the above descriptions. Group | includes R. brasilien- sis, R. cyanipes, R. guerrai, R. pseudocy- anipes, R. suffusus and R. viatorius viato- rius, Characterized by a short preapical di- verticulum (1.6 mm or less) and with the tip of the apical diverticulum oriented in the opposite direction relative to that of the preapical diverticulum (Figs. 1, 3, 4, 8, 10, 11). Group 2 includes only R. pratensis characterized by a short preapical divertic- ulum and an apical diverticulum oriented in the same direction (Fig. 7). Group 3 include R. palustris, R. pictus, and R. schistocer- coides with a long preapical diverticulum and an apical diverticulum with its tip clear- ly oriented in the opposite direction relative to that of the preapical diverticulum (Figs. 5, 6, 9). Group 4 includes a single species, R. brunneri, with a long preapical divertic- ulum and an apical diverticulum with its tip oriented towards the posterior part of the spermatheca (Fig. 2). CONCLUSION This is the first attempt to distinguish Scyllinini grasshoppers on the basis of sper- matheca design, and the results indicate that this character considerably facilitates iden- tifications. This criterion should be helpful in a taxonomic revision, while overcoming the problem of making identifications based on external male genital morphology. The genus Rhammatocerus and the entire tribe Scyllinini could be better revised by con- ducting systematic diagnoses on the basis of classical characters including also sper- mathecal morphology. ACKNOWLEDGMENTS This study was funded by FAPESP (Proc. No. 97/00264-3). For the specimen loans, interesting discussions, and review of VOLUME 102, NUMBER 1 the manuscript, we are especially grateful to Professors Miguel A. Monné (MNRJ), Carlos S. Carbonell (FCUM), Francisco Racca Filho (UFRRJ), Maria Marta Cigli- ano, Juan Schnack (FCNM), Maria José S. Lopes (UFPE), José Roberto Pujol-Luz, (UFRRJ), and Dr. C. Amedegnato (Muséum National d’Histoire Naturelle, Paris). We also thank Luis Antonio A. Costa (MNRJ) for the illustrations. LITERATURE CITED Assis-Pujol, C. V. 1997a. Revisao das espécies brasi- leiras do genero Rhammatocerus Saussure, 1861 (Orthoptera, Acrididae, Gomphocerinae, Scyllini- ni). Mestrado Thesis. Museo Nacional, Rio de Ja- neiro, Brazil. 124 pp. 1997b. Notas sinonimicas e redescrig¢ao de duas espécies de Rhammatocerus Saussure, 1861 (Orthoptera, Acrididae, Gomphocerinae, Scyllini- ni). Boletim do Museo Nacional de Rio de Janei- ro, N.S., Zoologia 376: 1-12. 1997c. Duas novas espécies brasileiras de Saussure, 1861 (Acrididade, Gomphocerinae, Scyllinini). Boletim do Museo Nacional de Rio de Janeiro, N.S., Zoologia 380: 1-10. . 1998. Aspectos morfologicos, taxonomicos e Rhammatocerus distribuigao geogrdfica de cinco espécies de Rhammatocerus Saussure, 1861 (Acrididae, Gom- phocerinae, Scyllinini). 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Colegao canavieira, Instituto do Acucar e do Alcool, Brasil, N°10. 622 pp. Jago, N. J. 1971. A review of the Gomphocerinae of the world, with a key to the genera (Orthoptera, Acrididae). Proceedings of the Academy of Nat- ural Sciences of Philadelphia 123: 205-343. Lecoq, M. and C. V. Assis-Pujol. 1998. Identity of Rhammatocerus schistocercoides (Rehn, 1906) forms south and north of the amazonian rain forest and new hypotheses on the outbreaks determinism and dynamics [Acrididae, Gomphocerinae, Scyl- linini]. Transactions of the American Entomolog- ical Society 124: 13-23. Lecogq, M. and G. Balanga. 1998. Field trials of fipronil for control of Rhammatocerus schistocercoides (Rehn, 1906) hopper bands in Brazil. Crop Pro- tection 17: LO5—110. Lecoq, M. and I. Pierozzi Jr. 1994. Rhammatocerus schistocercoides (Rehn, 1906), criquet ravageur de l'état du Mato Grosso (Bresil). Essai de syn- thése bibliographique. Centre de coopération in- ternationale en recherche agronomique pour le dé- veloppement, Montpellier, France. 89 pp. 1995a. Rhammatocerus schistocercoides \o- cust outbreaks in Mato Grosso (Brazil): A long- standing phenomenon. The International Journal of Sustainable Development and World Ecology 2: 45-53. 1995b. Le criquet du Mato Grosso: Lagriculture est-elle responsable? Tropicultura 1332-33. . 1996. Chromatic polymorphism and geopha- gy: Two outstanding characteristics of Rhamma- tocerus schistocercoides (Rehn 1906) grasshop- pers in Brazil [Orthoptera, Acrididae, Gompho- cerinae]. Journal of Orthoptera Research 5: 13— 17. 128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Lecoq M., Miranda E. E. de, and I. Pierozzi Jr. 1997. A new approach to the control of Rhammatocerus schistocercoides (Rehn, 1906) in Brazil, pp. S5O5— 506. Jn Krall S., R. Peveling and Ba Daoule Dial- lo, eds. New strategies in locust control. Birkhau- ser Verlag, Basel, Switzerland. Leon, M. G. 1996. 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Harvard Uni- versity Press, Cambridge, Mass. and London, England. 275 pp. Slifer, E. 1939. The internal genitalia of female Acri- dinae, Oedipodinae and Pauliniinae (Orthoptera, Acrididae). The Journal of Morphology 65: 437— 469. Uvarov, B. P. 1966. Grasshoppers and locusts. Vol. 1. Cambridge University Press, London. 481 pp. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 129-134 BIOLOGY AND BEHAVIOR OF THE STRAWBERRY GUAVA SAWELY, HAPLOSTEGUS EPIMELAS KONOW 1901 (HYMENOPTERA: PERGIDAE), IN SOUTHERN BRAZIL J. H. PEDROSA-MACEDO Laborat6rio de Protegao FlorestahA—DSM/SCA-UFPR, Av. Pref. Lothdrio Meissner, 3400, 80.210-170 Curitiba, Parana, Brazil (e-mail: johpema@netpar.com.br) Abstract.—Results of four years of observation and study of the strawberry guava sawfly, Haplostegus epimelas Konow 1901, in two ecosystms, coastal plain and Parana first plateau of Brazil, are presented. The strawberry guava sawfly is trivoltine, feeding almost exclusively on sprouting araga, or strawberry guava, Psidium cattleianum Sabine (Myrtaceae). This plant is native to the Brazilian Atlantic forest but is currently found throughout the tropics where it is frequently a weed. Adult female sawflies are rarely found in the field and males have never been found. The eggs are laid when the sprouts are young. Larvae are gregarious and go through six or seven instars. Pupation occurs in the soil. Natural enemies attack all stages. The total parasitism frequency is 3.6%, mostly by Vibrissina sp. (Diptera: Tachinidae). Two main types of damage to strawberry guava are Oviposition wounds resulting in the death of twigs and young leaves consumed or damaged by larval feeding. The sawfly completes its life cycle on strawberry guava but has recently been observed feeding on commercial guava, P. guajava L., which may preclude its use as a biological control agent. Resumo.—Sao apresentados os resultados de quatro anos de observagoes e estudos sobre a vespa-serra, Haplostegus epimelas Konow 1901 (Hymenoptera: Pergidae) em dois ecossistemas: Litoral (Restinga) e Primeiro Planalto Paranaense. O dimorfismo sexual entre machos e fémeas é distinto por trés caracteristicas morfologicas. A vespa-serra procria-se e alimenta-se na brotagao sazonal do aragazeiro, Psidium cattleianum Sabine (Myrtaceae) e é trivoltina. Esta planta originalmente da Floresta Atlantica brasileira en- contra-se distribuida nas regides tropicais e subtropicais onde se tornou daninha. Fémeas adultas da vespa-serra sdo raramente encontradas no campo e os machos nunca foram vistos. Os ovos colocados em brotagoes tenras. As larvas sao gregarias e passam por 6— 7 intares. O empupamento ocorre no solo. Ha inimigos naturais em todas as fases do ciclo vital. O parasitismo por Vibrissina sp. (Diptera: Tachinidae) atinge 3, 6%, é o mais im- portante. Dois tipos de danos sao encontrados: lesGes nos ramos pelo ato de postura e os danos causados nas folhas jovens por larvas neonatas. A vespa-serra completa seu ciclo vital no aragazeiro e foi recentemente encontrada em goiabeira, Psidium guajava L. (Myr- taceae) fato que pode limitar 0 seu uso como um agente de controle bioldégico. Key Words: araga, biological control, biology, behavior, strawberry guava, sawfly For almost two centuries, araga or straw- (Myrtaceae), has attracted the attention of berry guava, Psidium cattleianum Sabine _ horticulturists who have used it as an or- 130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON namental plant and fruit tree. It has been introduced into several parts of the world, where it has spread aggressively competing with the local vegetation (Smith 1985). In its native Brazilian Atlantic forest, it grows harmoniously with the other vegetation of the woods. Two types of strawberry guava are known, one with red fruit and the other with yellow fruit. In Brazil, the red-fruited form is confined to higher elevations above 850 m, whereas the yellow-fruited form is found throughout the range. Elsewhere in the world, the two forms are found at all elevations. In its native woods, this tree can be of medium height (up to 5 m), but it has a low density per hectare (<1% cover). In contrast, the trees along roads, urban areas, and agricultural or pasture areas are abun- dant and bushy. Numerous natural enemies and competition from other plants effec- tively control strawberry guava in Brazil. This study describes the biology of Hap- lostegus epimelas Konow, the associated phenology of strawberry guava, and the im- pact of the sawfly’s damage on the plant. The objective is to evaluate the potential of the sawfly as a biological control agent in Hawaii. Some other insect species, mainly gall midges, are important natural enemies of strawberry guava (Wikler 1995, Vitorino 1995, Angelo 1997). A species of Coleop- tera (Chrysomelidae) is also destructive, but it is not confined to strawberry guava (Cax- ambu 1998). According to Smith (1993), some Pergidae are considered forest pests in North and South America, New Guinea, and Australia. Benson (1940) and Smith (1990) reported that H. epimelas also at- tacks guava, Psidium guajava L., in the State of Pernambuco. MATERIAL AND METHODS Study area.—Field observations of the plant and sawfly were carried out in two regions: (1) First plateau of Parana (48°58’ to 49°38’ W; 25°27’ to 25°35’S), particularly the districts of Curitiba, Balsa Nova, Cam- po Largo, Almirante Tamandaré, Colombo, Piraquara, and Sao José dos Pinhais; and (2) the coastal plains of Parana and Santa Catarina (Restinga) (48°29’ to 48°46’W and 25°19’ to 26°03'S) especially the districts of Antonina, Morretes, Paranagua, Pontal do Parana, Guaratuba, and the coastal plain of Santa Catarina (only the district of Itapoa). According to Carpanezzi et al. (1986), the first plateau of Parana is characterized by altitudes from 650 to 1,100 m, and by a rain forest of Araucaria angustifolia (Araucari- aceae) and submontane fields. Within the Holdridge system, the climate is temperate, hot and humid, or very humid. The average annual temperature ranges from 15° to 19°C, with an absolute minimum of —10°C. One to forty frosts occur per year. The av- erage rainfall ranges from 1250 to 2500 mm and there is no water deficit. The coastal plains of Parana and Santa Catarina are characterized by native rain forests of low altitude, ranging from 0 to 500 m. The cli- mate is subtropical, humid or very humid, according to the Holdridge system. The av- erage annual temperature ranges from 18° to 22°C, with an absolute minimum of —0.9°C. Frosts are rare. The annual rainfall ranges from 1,600 to 2,000 mm, and there is no water deficit. Methodology.—All captured insects were taken to the nursery at the Forest Pro- tection Laboratory, Federal University of Parana, where they were kept alive for ob- servation. Prepupae were kept in containers of two different sizes (3 X 7 cm, vol. 45 ml; 5 < 11 cm, vol. 180 ml) to study adult and parasite emergence and to record life history data, e.g., adult longevity, oviposi- tion, depth of pupation, and pupal duration. Diameter of cocoons, period of oviposition, and number of eggs were measured on shoot tips or branches. The growth of strawberry guava was observed using 20 trees previously chosen at random and tagged, ten in the District of Colombo, Pa- rand (Estancia Betania), and ten from the District of Piraquara, Parana (Mananciais da Serra Mar). The presence of adult saw- flies, eggs, and larvae were counted and ob- served from September 1994 through July VOLUME 102, NUMBER 1 1998. Standard statistical tests were used. Damage caused by oviposition and larvae were evaluated during a year of observa- tions and collecting of field data. RESULTS AND DISCUSSION Host plants and host specificity.—In the states of Parana and Santa Catarina, Hap- lostegus epimelas was observed on yellow- (350) and red-fruited (25) strawberry guava plants during weekly visits to the field and in the University gardens. The red-fruited trees are not found in the coastal plains. In the two states cited above, there are several other Myrtaceae, such as Psidium spathu- latum Mattos, Campomanesia xanthocarpa Berg., Eugenia uniflora L., Eugenia invol- ucrata DC, Myrciaria trunciflora Berg., Gomidesia schaueriana Berg., and Acca sellowiana (Berg.) Burret. Sawfly eggs and larvae were not found on any of these trees. Sawfly life history—The female aver- ages 10.2 + 0.3 mm long (n = 13) and the male 8.3 + 0.3 mm long (n = 15). Both the male and female are black dorsally with the ventral part of the head and the poste- rior part of the abdomen dark brown, the thorax orange, and the front legs orange to the tibiae with the tarsi almost black. The female is larger than the male; otherwise, the usual sexual characters separate them. In the field, the female is rarely observed on the plant, and males were never seen. To Oviposit, the female turns upside down on young, soft branches to lay eggs. The di- ameter of these branches range from 2.1 to 3.9 mm (2.9 + 0.1 mm) (n = 61). Ninety egg masses on 20 strawberry guava trees were observed. Only eight females were observed during oviposition. The eggs are laid in a line, under the epidermis of young shoots that are slightly reclined. Between each egg, there is a septum, a fiber filament of the vegetable tissue, which separates them (Fig. 1). The number of eggs laid varies from 52 to 118, averaging 90.5 + 16.4 per female (n = 13). As many as 178 eggs were found in the ovary of one virgin female, suggest- 131 Necrotic bark Fig. 1. Oviposition site of Haplostegus epimelas. Eggs are distributed symmetrically and reclined. ing that they move to other branches to con- tinue oviposition. The eggs are rod-shaped, with ovoid tips. Eggs are white from ovi- position to eclosion. In the field, incubation lasts from 7 to 12 days with an average of 10 days (Fig. 2). The gregarious, newly hatched larvae move to young, recently expanded leaves to feed. Initially, they feed by scraping the un- der surface of the leaf. Fourth instar larvae consume the entire leaf. In the last instars they remain gregarious, but they sometimes split up into smaller groups. When larvae are disturbed they bend the posterior part of their abdomen over their dorsum showing an attitude of intimidation. If they are touched, they regurgitate a drop of fluid which hangs from their mouths. The smell of the regurgitated substance is strong and similar to the smell of the host tissue on which they are feeding. Presumably, the flu- id deters natural enemies. The growth of the larvae and the number of instars are not al- ways homogeneous. Observations in the laboratory show that the gregarious habit of the larvae is vital for the survival of the larval colony. When a group of first instar larvae is smaller than ten, they die before completing their larval development. Lar- vae were observed to go through 6 or 7 instars. Growth of some larvae are delayed in relation to others from the same egg batch. As a consequence, they go through at least one additional instar. When they reach the end of the larval period, the lar- vae are 22.1 + 0.7 mm long (n = 15). They stop feeding and start draining their diges- 132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON [| 4 days eo 10 days 22 days ADULTS EGGS LARVAE PUPAE Fig. 2. Life cycle of Haplostegus epimelas. tive tube, changing their tegument and their head capsule. The last exuvium is larger than the previous one and is adhesive and sticky to the touch. It loses its adhering property when it dries out. The prepupa is 11.8 + 0.4 mm long (n = 12). The dorsum is light brown and the venter is light beige; the head and the endo of the abdomen are black. It is easy to see that there is nothing in the prepupa intes- tine. Its body is thinner and shorter com- pared to that of the full-grown larva. The prepupa drops off the branch and digs into the organic soil within one hour, taking ad- vantage of natural depressions or holes. The average depth of penetration in sandy soil is 20.3 + 3.6 mm (n = 12) as measured from pupae inside plastic containers. After penetrating the soil, the prepupa produces silk to form a cocoon in which it remains free for a variable time. After a few hours, some prepupae lose the capacity to pene- trate the soil and become stiff, getting in- fected by fungi. The male cocoon is 7.7 + 0.2 mm long (n = 12) and has a diameter of 4.3 + 0.2 mm (n = 12). The female co- coon is 8.8 + 0.1 mm long (n = 20) and has a diameter of 5.0 + 0.2 (n = 20). The cocoon is spun with white, thin, soft silk fibers, but, in the soil or sand, it later be- comes dark and stiff. Particles of soil or sand adhere to it, making it look bigger than it actually is. For males, the average pupa- tion period is of 21.5 + 1.0 days (n = 51). For females, it is 21.6 + 1.5 days (n = 24). From 330 prepupae, 75 (51 males and 24 females) H. epimelas adults emerged, 6 adults of another sawfly (Truqus magnus Smith 1990, Hymenoptera: Pergidae, iden- tified by D. R. Smith) emerged, and para- sites emerged from 12 others. For unknown reasons, 237 pupae (71.8%) died during the pupation period. The larvae of Truqus mag- nus were found in Mananciais da Serra in Piraquara, Parana, on the yellow-fruited strawberry guava and in the neighborhood of Santa Felicidade, Curitiba, Parana, on the red-fruited strawberry guava. Smith (1990) described the species from samples collect- ed in Campina Grande do Sul, Parana, northern Curitiba. Adult females oviposit and live for 2 to 3 days in the laboratory according to Ped- rosa-Macedo (1998). No maternal behavior was observed. The larval phase is generally between 28-30 days; however, some feed up to 35 days (Fig. 2). The prepupal phase, inside a cocoon, has variable longevity. They can remain in the soil for several weeks. We have observed three peaks in H. epimelas populations for two years, sug- gesting that it is trivoltine (Fig. 3). Natural enemies.—The main parasitoid of H. epimelas is an undetermined species of Vibrissina (Diptera: Tachinidae). There are also two other species of the family Sar- VOLUME 102, NUMBER 1 1“ Cycle a" Cycle | October March December — January Fig: 3: cophagidae (Diptera). These parasites emerge from the cocoon after an average of 28.3 + 0.9 days (n = 12) after pupation of the host. The percentage of parasitism re- corded was 3.6% (n = 330). One mite egg predator was observed. Its presence was recorded in November 1997 at Mananciais da Serra, Piraquara, Parana. One female adult sawfly was found caught in a spider web. The fungi Fusarium sp. and Verticillium sp. were isolated from dead prepupae, but it is not known if they were responsible for deaths. Damage.—Three types of damage are caused by the sawfly on young shoots of strawberry guava. The first and most im- portant for biological control is the wound caused by the female during oviposition. The second is by the young larvae (first to third instars) which scrape the underside of young and soft leaves on the same sprout as the egg mass. The third is by the fourth to seventh instar larvae which eat mature leaves entirely elsewhere on the plant. The female destroys the soft tissue of the shoots when penetrating their epidermis with her saw-like ovipositor. The action of the young larvae further injures the young shoots re- sulting in the death of 31% of the shoots (28 of 89 observations) leaving 69% of the shoots alive. When the first and second types of damage occurred together, they caused irreparable loss to the strawberry guava, which survived but had permanent damage, i.e., the branches were bent. The activity of the sawfly is synchro- nized with flushing of the strawberry guava. The sprouts on the branches are abundant in the spring and summer. In autumn, new May June July September Annual seasonal activity of Haplostegus epimelas, 1997 to 1998. sprouts can be seen, but in winter they are scarce. Therefore, there are sprouts on the strawberry guava which enable the survival of sawflies in all four seasons of the year. Nevertheless, in the coast and plateau re- gions, there was a period of 120 + 7.9 days (n = 3, June 1995 to July 1998) in which no adult sawflies, eggs, or larvae were found. Use as a biological control agent.—The pupal phase of Haplostegus epimelas is easy to transport or export. A 45 ml capac- ity container (3 X 7 cm) is adequate for handling and exporting pupae. The natural enemies of the sawfly contribute to their population balance, but they do not limit the impact and damage caused to the plant. Unfortunately, the sawfly has recently been observed attacking two plants of Psidium guajava. This observation, though perhaps an unusual even, may curtail an attempt to use this species as a biological control agent. ACKNOWLEDGMENTS I thank Dr. Clifford W. Smith, Depart- ment of Botany, University of Hawaii at Manoa; Dr. David R. Smith, Systematic En- tomology Laboratory, USDA, National Museum of Natural History, Washington D.C.; Dr. Alvaro Figueiredo, EMBRAPA/ Colombo, Parana; the biologist Ronaldo Toma, the designer Ricardo Pedrosa Ma- cedo, the administration of SANEPAR in Piraquara, the Estancia Betania in Colom- bo, and all the members of the research group on biological control of plants of the Laboratory of Forest Protection of the De- partamento Silvicultura e Manejo-SCA- 134 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON UFPR, for their suggestions and contribu- tions. Finally, I thank Dr. Peter W. Price, Northern Arizona University, Flagstaff, for reviewing the manuscript. LITERATURE CITED Angelo, A. C. 1997. A Galha dos botdes do araga- zeiro—Psidium cattleianum Sabine, 1821 (Myr- taceae) e Insetos Associados. Curitiba, Disserta- ¢ao de Mestrado, Curso de Pds-Graduagao em Ciencias Biolégicas, Area de concentracgao em En- tomologia, Setor de Ciencias Biolégicas, Univer- sidade Federal do Parana, 95 pp. Benson, R. B. 1940. Three sawflies attacking guava in Brazil (Hymenoptera, Symphyta). Bulletin of En- tomological Research 30:463—465. Carpanezzi, A., A. Ferreira, C. A. Rotta, et al. 1986. Zoneamento ecoldgico para plantios florestais no Estado do Parana. Curitiba. 89 pp. (EMBRAPA- CNPE Documentos, 17.) Caxambu, M. G. 1998. Morfologia e aspectos bioe- cologicos de Lamprosoma azureum Germar, 1824 (Coleoptera: Chrysomelidae) associado a Psidium cattleianum Sabine (Myrtaceae). Curitiba, Disser- tacao de Mestrado do Curso de Pés-Graduagao em Ciencias Bioldgicas, Area de concentrgao em En- tomologia, Setor de Ciencias Biol6gicas, Univer- sidade Federal do Parana, 78 pp. Pedrosa-Macedo, J. H. 1998. Contribuigao aos estudos bioecologicos da vespa-serra, Haplostegus epi- melas Konoy, 1901 (Hymenoptera: Pergidae) e ao controle biold6gico do aragazeiro, Psidium cat- tleianum (Myrtaceae). In VI SICONBIOL—6° Simposio de Controle Biologico, ANAIS: Sessoes de Posteres. Rio de Janeiro 24—28 de maio de 1998. Smith, C. W. 1985. Impact of alien plants on Hawaii’s native biota, pp. 180—250. Jn Stone, C. P. and J. M. Scott, eds. Hawaii’s terrestrial Ecosystems: Preservation and Management. Cooperative Na- tional Park Resources Studies Unit, University of Hawaii, Honolulu. Smith, D. R. 1990. A synopsis of the sawflies (Hy- menoptera, Symphyta) of America South of the United States: Pergidae. Revista Brasileira de En- tomologia. 34: 7—200. . 1993. Chapter I. Systematics, life history, and distribution of sawflies, pp. 3-39. In Wagner, M. and K. EF Raffa, eds. Sawfly life history adapta- tions to Woody Plants. Academic Press. San Di- ego. Vitorino, M. D. 1995. Aspectos bioldgicos e de espe- cificidade de Tectococcus ovatus Hempel, 1990 (Homoptera: Eriococcidae) para 0 controle biolé- gio do aragazeiro, Psidium cattleianum Sabine (Myrtaceae). Curitiba, Dissertagao de Mestrado, Curso de Pos-Graduagao em Engenharia Florestal, Setor de Ciencias Agrarias, Universidade Federal do Parana, 58 pp. Wikler, C., M. D. Vitorino, and J. H. Pedrosa-Macedo. 1994. Projeto de Controle Bioldgico do Aragazei- ro—Psidium cattleianum Sabine, 1821. Revista Setor de Ciencias Agrarias (UFPR), pp. 247-254. Wikler, C. 1995. Aspectos bioecolégicos do euritomi- deo causador de galhas nos ramos do aragazeiro— Psidium cattleianum Sabine. Curitiba, Dissertagao de Mestrado, Curso de Pos-Graduagao em Engen- haria Florestal, Setor de Ciencias Agrarias, Univ- ersidade Federal do Parana, 62 pp. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 135-141 JORDANOPSYLLA BECKI (SIPHONAPTERA: CTENOPHTHALMIDAE), A NEW SPECIES OF FLEA FROM THE NEVADA TEST SITE MICHAEL W. HASTRITER Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM, PO. Box 20200, Provo, UT 84602-0200, U.S.A. (e-mail: hastritermw @ sprintmail.com) Abstract.—Jordanopsylla becki, a new species of flea collected 1.25 km north of Mer- cury, Nevada Test Site, Nye County, Nevada, from Neotoma lepida is described and illustrated. This flea, assigned to the subfamily Anomiopsyllinae, tribe Jordanopsyllini, is a nest flea found as an adult only during the winter months. Techniques for recovering this rare flea are described. Key Words: Traub and Tipton (1951) erected a new genus and tribe in the subfamily Anomiop- syllinae for two females of the rare flea, Jordanopsylla allredi. Only 12 specimens (9 3, 3 2) of this species have been col- lected and the male was only recently de- scribed by Hastriter, Egoscue and Traub (1998). Morphological details described by the latter authors also support the system- atic position of the genus in the subfamily Anomiopsyllinae, tribe Jordanopsyllini as previously described by Traub and Tipton (1951). Beck and Allred (1966) reported five (3 ¢6, 2 2) additional specimens of J. allredi from the northwest slopes of the Spotted Range and Red Mountain, Nevada Test Site (near Mercury), Nye County, Ne- vada and subsequently provided these spec- imens to the late Robert Traub. Traub con- cluded that they represented a new species. Prior to his death, he kindly submitted these five specimens to me for description. Be- cause of the dearth of material belonging to this genus, I sought more specimens of this new species by collecting on the northwest slope of Red Mountain during January 12— 14, 1999. The Spotted Range was not ac- cessible during this collection trip. flea, Nevada, Jordanopsyllini, Neotoma After nearly 50 years of intensive collec- tion efforts, Jordanopsylla is represented by only two taxa. Most of the specimens (with the exception of recent collections) of both taxa were either collected by, or under the direction of D. M. Allred and D. E. Beck, Brigham Young University. Since the nom- inate species was named after Allred, it seems fitting to name this new species after Beck, honoring the team efforts of both of these naturalists who contributed so much toward our understanding of the Siphon- aptera of the Great Basin and the northern fringes of the Mojave Desert. MATERIALS AND METHODS Aluminum collapsible Sherman" traps baited with oatmeal were set at three sites on January?) D1=13, 921999 BiCSiteo 1) 36°41.5'N, 115°58.8'W, (Site 2) 36°41.7'N, 115°58.3°W,. and. "(Site 3) 36°42-6'N, 115°59.5’W]. Sites 1 and 2 had steep slopes with rocky limestone outcroppings, cliffs, and talus. Dominant vegetation included sparsely distributed black brush (Coleogyne ramosissima Torr) and yucca (Yucca schi- digera Roezl ex Ortega). The slope of site 3 was more gentle than that of sites 1 and 136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2 with scattered basalt rock. Black brush, creosote (Larrea tridentata Coville), and yucca were the predominant vegetation at site 3. Mammals were shaken from traps direct- ly into white cloth bags (23 X 28 cm) each morning and processed immediately after collection for fleas. The number of mammal specimens of each species that could be re- tained was restricted by permit, necessitat- ing release of some. Only fleas that escaped in the bags were obtained from animals that were released. Other mammals were sacri- ficed by cervical fracture, while in the bags. The entire contents of each bag (mammal, fleas, and debris) were emptied into a white 19 liter bucket. The mammal was held be- low the level of the top of the bucket while blowing a gentle stream of CO, over the pelage to excite the fleas to vacate the an- imal. Because all fleas do not become agi- tated by CO,, each animal was also combed briskly with a toothbrush (with a single row of bristles). Hosts placed in individual plas- tic bags were frozen at the end of each day. Upon return to the laboratory, they were in- dividually washed by rapidly stirring the frozen animal back and forth in a 3.8 liter jar filled half full with tepid tap water. The water was poured into a white enamel pan and fleas flushed from the animal’s fur were collected. Nests of Neotoma lepida Thomas were collected from site 1 (N = 7) and site 3 (N = 4). Each was placed in a plastic bag, sealed, and returned to the laboratory. Fleas and flea larvae were collected by placing the 11 nests in Berlese funnels for 48 hours (two nests were combined in each of four funnels and three in a fifth funnel). RESULTS Seventy-three mammals [Peromyscus crinitus (Merriam) (N = 42), N. lepida (N = 19), Peromyscus eremicus Baird (N = 7), Peromyscus maniculatus (Wagner) (N = 2), Onychomys torridus (Coues) (N = 2), and Dipodomys ordii Woodhouse (N = 1)] were collected during 280 trap nights. Peromys- cus crinitus and N. lepida comprised 84% of the mammals trapped. One hundred six- ty-four fleas [Orchopeas sexdentatus agilis (Rothschild) (N = 82), Malaraeus sinomus (Jordan) (N = 74), Anomiopsyllus amphi- bolus Wagner (N = 4), J. becki n. sp. (N = 2), Stenistomera alpina (Baker) (N = 1), and Carteretta clavata Good, (N = 1)] were obtained from mammals (none from O. torridus or D. ordii) and 126 fleas [O. s. agilis (N = 91), A. amphibolus (N = 32), and J. becki (N = 3)] were obtained from nests. Two Jordanopsylla females were re- covered from the pelage of two adult N. lepida females during washing, while none were collected from the animals during thorough field processing. A male and fe- male Jordanopsylla were each collected from a composite of two nests from the same Berlese funnel and an additional fe- male from three N. lepida nests combined in another funnel. All five specimens of Jordanopsylla were collected between 1,225 m and 1,305 m. Nests of N. lepida are usually difficult to obtain, since they are frequently established in inaccessible rock areas or deep within ancient wood rat middens. The author dis- covered that the wood rats living among Y. schidigera almost always built their nests at the base of a dead yucca stump. Nests could seldom be located unless the healthy yucca stands (two or three stalks) contained a dead stump. The central core of the woody stock decay, leaving a hard outer surface covered with dead spear-like leaves. These stumps provide protection for N. lepida and an environment for flea development during extreme desert conditions. By carefully pulling the dead stumps over, the decayed root system exposes the wood rat nest cen- trally located beneath. With care, the stumps may be turned over, the nest col- lected, and the stump replaced, minimally disturbing the building site for future wood rat nests. Jordanopsylla becki Hastriter, new species (Figs. 2—6) Type data.—Holotype (1 6, field no. 734), ex Neotoma lepida, 1.25 km north of VOLUME 102, NUMBER 1 137 Figs. 1-6. Jordanopsylla spp. 1, J.allredi, ninth sternum and lateral lobe of aedeagus. 2—6, J. becki. 2, ninth sternum and lateral lobe of aedeagus. 3, aedeagus. 4, metatarsus (holotype). 5, spermatheca. 6, female sixth and seventh sternites (note: sinus in posterior margin of st. VI). Scale = 100 w. 138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mercury, Nevada Nuclear Test Site, Nye County, Nevada, October 28, 1961, collec- tom, SHR” allotype (to, field, nos 723), same data except collector ““MCD” and paratypes (2 d, field no. 670 each), same data except collector ““SHR’’; paratype (1 ?, field no. 647), same data except Novem- ber 10, 1961; and paratypes (1 6,1 9°, field no. MH-446/447, 1°, field no. 451, 1 °, field no. 464, and 1 9, field no. 472), 1.25 km north of Mercury on northwest facing slope of Red Mountain, Nevada Test Site, Nye County, Nevada (36°41'N, 115°58'W), elev. 1,225—1,305 m, January 12—13, 1999, M. W. Hastriter. Holotype, allotype, and three paratypes (2 d, 1 2) are deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C.; one paratype (2) in the Robert E. Lewis collection; and four paratypes (1 ¢,3 @) in the author’s collection. Diagnosis.—Males of Jordanopsylla be- cki differ from those of the nominate spe- cies in the shape and setation of st. IX and the extended length of the lateral lobe of the aedeagus. The ventral caudal margin of the apical lobe of st. [X is distinctly angular, bearing two or three well-developed setae, whereas in J. allredi it is evenly rounded without conspicuous setae. The apex of the lateral lobe (lateral lobe fused to st. [X in both species) extends well beyond the pos- terior margin of the apical lobe of st. IX in J. becki and extends only to the margin in J. allredi (Figs. 1—2). Caudal margin of st. VII of female is similar to J. allredi with a broad, shallow concavity subtended by a weakly developed ventral lobe (Fig. 6), however, the setation differs. The st. VII of the new species has two parallel oblique rows of setae per side (two anterior and three posterior). Jordanopsylla allredi has only a single oblique posterior row of five slender setae per side and four smaller setae per side anterior to the oblique row. The smaller setae are arranged in pairs along the ventral margin at the level of the ventral most seta of the oblique row. Jordanopsylla becki females also have a markedly scler- otized zone surrounding a distinct sinus on the caudal margin of the st. VI (Fig. 6). This sclerotization is not apparent in the fe- male holotype of J. allredi. Although the latter holotype was drastically over-cleared, it is doubtful that over-clearing could oblit- erate the heavy sclerotization noted in J. be- cki. Collection of additional females of J. allredi may substantiate or refute this ob- servation. It is noteworthy, that Jordanop- sylla is the only genus in North America with a distinct sinus in st. VI, a feature shared only by species of Tetrapsyllus (Te- trapsyllus) Jordan that occur in western South America from Ecuador south into Chile and Argentina. Description.—Male. Head: Frons smooth- ly rounded without frontal tubercle. Incras- sation at oral angle, 2 marginal placoid pits in preantennal region, and a single placoid pit in lateral occipital area. Depth of occip- ital groove nearly equal to width of first segment of maxillary palpus. Eye moder- ately well developed, ventrally notched. Tentorial arm anterior to eye. Preantennal setae with ocular row of 3, upper and lower longer than middle seta. Setae in occipital area restricted to posterior row of 4—5 setae per side, most ventral seta largest and set slightly anterior to row. Dorsal margin of antennal fossa with line of 23—24 minute coniform setae. Scape bearing | small dor- sal seta, 3 along apical margin, pedicel not over-lapping onto clavus, with 3 setae % length of clavus. Proximal he!lf of maxilla sclerotized, distal half transparent and dif- ficult to distinguish, rounded apex extend- ing half the length of first segment of labial palpus. Maxillary palpus of 4 segments, each similar in length. Labial palpus of 5 segments, terminal segment longest, ex- tending beyond trochanter in its entirety. Thorax: Without combs or spinelets. Pro-, meso-, and metanota subequal in length along dorsal margin, each with posterior row of 5—6 setae per side plus intercalaries; anterior row of 1—2 small setae; 5—6 pseu- dosetae per side beneath mesonotal collar. Proepisternum without setae, dorsal depres- VOLUME 102, NUMBER 1 sion sheaths terminal segments of antennal clavus. Mesosternum with 1 seta, anterov- entral margin acuminate and sclerotized, mesepimeron with 3 setae. Distinct lateral metanotal area with 2 setae. Well developed pleural arch. Metasternum with one seta, dorsoanterior margin convex, anteroventral margin lobate. Metepimeron bearing 3 se- tae, dorsal margin not fused to metanotum. Legs: Procoxa with 16—17 setae (including marginals), 2 setae on mesal side of anterior margin. Meso- and metacoxae with numer- ous slender setae along anteromesal margin from dorsoanterior base to apex. Pro-, meso-, and metafemora each with single lateral seta, setae along dorsal margins, and line of setae mesally from base to apex. Paired bristles guarding femoro-tibial joint of procoxa equal in length, outer bristles of pair shorter in meso- and metafemora. Pro-, meso-, and metatibia each with a lateral vertical row of 2, 4, 4 setae, respectively. Tarsal segment V of all legs with 4 lateral plantar bristles, 2 preapical plantar bristles (subspiniform), and 2 preapical plantar hairs (Fig. 4). Unmodified abdominal seg- ments: Tergites I-III with marginal spine- lets (includes both sides) 3—5, 2—3, and O— 1, respectively. Intercalary setae in all pos- terior rows of t. I-VII, which have 4, 6, 5— 6, 5—6, 5, 5, 5 setae per side, respectively. A single seta in posterior row of each ter- gite lies ventrad to each spiracle. Anterior rows of t. I-VII with 2-3, 2-3, 1-2, 1-3, 0, QO, O setae per side, respectively. One ante- sensilial bristle per side. Per side, st. II has 1 small setae, st. III-VII each has 2, and st. VIII has 4 long setae. Modified abdominal segments: Tergum VIII, reduced posterior- ly, not extending over body of clasper, bear- ing 2 small setae anterior to a vermiform spiracle and | stout seta posterior to the spi- racle. Body of clasper, movable process, and manubrium indistinguishable from those of J. allredi: characterized by a small movable process, 2 acetabular bristles, a distinct fovea, anterior margin of t. X and manubrium semicircular in outline with apex of manubrium truncately rounded. 139 Apical lobe of the distal arm of st. [IX an- gular at its ventral margin, truncate at the apex, and armed with a row of submarginal setae that enlarge towards the ventral mar- ginal angle (Fig. 2). The apodemal rod of st. [IX proximally fragile. The distal arm of the st. IX laterally fused to the greatly en- larged lateral lobe of the aedeagus, char- acteristic of the genus. Aedeagus: Very much like J. allredi, except for the pro- nounced lateral lobe (Figs. 2—3). Aedeagal apodeme broadest medially, tapering to a narrow neck near fulcral lobes. Penis rods exceeding aedeagal apodeme, but not coiled. Median dorsal lobe nipple-like, lat- erally expanded into paired distolateral lobes, lateral lobes, and a large single me- dian lobe. The apical lobes of the distal arm of st. IX envelops the large median lobe and the apical lobes are enveloped and fused to the lateral lobes. Sclerotized inner tube is long and slender, bearing a dorsal tooth ba- sally. Crochets lacking. Female. Characteristics similar to those of male except as follows: Frons smoothly rounded to posterior occipital margin with minute punctations and 4 marginal placoid pits. Dorsal margin of antennal fossa lack- ing line of minute coniform setae, pedicel with 3 setae extending to apex of clavus, clavus shorter than in male. Segment 2 of maxillary palpus distinctly longer than oth- ers. Mesonotal collar with 7—9 pseudosetae per side. Fewer spinelets on abdominal t. I— II than in male (1—4 and 0-2, respectively). Anterior row of setae on t. III-VII range from 3—5, 2—4, 2—4, 2—3, and 2-3 per side, respectively. Tergum VIII with 3-5 setae dorsad to vermiform spiracle, a single an- terior row of 7—9 setae below spiracle, and a group of 20—25 setae on caudal margin, many somewhat spiniform. Abdominal st. II with lateral patch of 6—9 setae and a sin- gle apical ventral seta per side, st. III-V each with 2-3 setae per side. Sixth sternum with distinct sinus in caudal margin sur- rounded by heavy sclerotization, 2 basal se- tae per side and a single seta above sinus (Fig. 6). Caudal margin of st. VII entire 140 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with slight broad concavity, each side with two parallel oblique rows (2 in anterior row and 3 in posterior row). Eighth sternum bluntly rounded with 5—6 minute terminal setae. Anal stylet with long apical bristle and 1—2 minute setae. Bursa copulatrix par- tially sclerotized. Single spermatheca with globular bulga, broad cribriform area, hilla upturned, apex extending slightly beyond bulga, indistinguishable from nominate spe- cies) (Pig. 5): Size: (total length mm, range followed by mean, mounted specimens) (N = 3 6, 2 2). Male = 1.8—2.2, 2.0; Female = 2.5— 2.7, 2.6. (alcohol specimens) (N = 1 6, 3 2). Male = 1.5, Female = 1.6—1.8, 1.7. DISCUSSION The Nevada Test Site has a faunal tran- sition zone between the Great Basin to the northwest and the Mojave Desert to the southeast. Jordanopsylla becki occurs to the south of this transition zone. It is lo- cated 225 km from the closest known western population of J. allredi and south 30 minutes latitude. Subspecific designa- tion is inappropriate as the two species are morphologically distinct and the two known populations of J. allredi occurring 50 km apart (within 04 minutes latitude) in Washington County, Utah do not demon- Strate a variable character cline from east to west. Habitats for both Utah populations are similar (Hastriter et al., 1998) and oc- cur within a narrow range of elevation (940—1,190 m). The new species occurs at slightly higher elevations (1,225—1,305 m) than the nominate species (author’s collec- tions). Beck and Allred (1966) provide only general reference as to where they collected J. becki without mention of ele- vation. Both species are restricted to the plant communities associated with com- mon flora of the Mojave Desert (creosote, yucca, prickly pear cactus, and pale cholla) opposed to those of the Great Basin. Neo- toma lepida, as suggested by Beck and Allred (1966), is undoubtedly the preferred host for J. becki, however, additional col- lections are needed to elucidate the host range of this new species. The distribution of both species of Jordanopsylla appears to be limited by micro-environmental re- quirements of their immature stages, but seasonal abundance and collecting tech- niques may also play a role in its rarity in collections. All known specimens in the genus have been collected during the months of November (N = 1), December (N = 7) and January (N = 10) indicating that Jordanopsylla is certainly a winter flea. Among the eight specimens of Jor- danopsylla collected by the author (J. all- redi, 36 and J. becki, 16, 4 ¢), four of the eight were obtained by reexamining bags or washing specimens more than 24 hours after initial attempts failed to collect them (three collected in nests). Similar te- nacity also has been demonstrated for S. alpina. Many collectors have sought to collect specimens of Jordanopsylla with- out success. The author would suggest the technique of washing animals thoroughly to maximize the opportunity to collect Jor- danopsylla. Although many specimens of P. crinitus have been collected in the same areas where Jordanopsylla has been col- lected from P. eremicus and N. lepida, fleas of this genus have never been col- lected from this host. The topography of much of the expanse separating the two species is lower in elevation than the type localities of either species. One might speculate that such natural isolation might have contributed to speciation of Jordan- opsylla. ACKNOWLEDGMENTS Appreciation is expressed to Nancy Ad- ams, Curator of Siphonaptera, National Museum of Natural History, Smithsonian Institution for loan of specimens and to Robert Furlow and Derek Hall, Bechtel Nevada, Nevada Test Site, for their assis- tance in facilitating on-site mammal col- lections. VOLUME 102, NUMBER 1 LITERATURE CITED Beck, D. E. and D. M. Allred. 1966. Siphonaptera (fleas) of the Nevada Test Site. Brigham Young University Science Bulletin, (Biological Series) 7(2): 1-27. Hastriter, M. W., H. E. Egoscue, and R. Traub. 1998. A description of the male of Jordanopsylla allredi 141 (Siphonaptera: Ctenophthalmidae). Proceedings of the Entomological Society of Washington 100(1): 141-46. Traub, R. and V. J. Tipton. 1951. Jordanopsylla all- redi, a new genus and species of fleas from Utah (Siphonaptera). Journal of the Washington Acad- emy of Science 41(8): 264-70. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 142-150 A NEW SUBGENUS, FOOTERELLIA, AND NEW DISTRIBUTION RECORDS OF NEASPILOTA OSTEN SACKEN (DIPTERA: TEPHRITIDAE: TERELLIINI) ALLEN L. NORRBOM AND BENJAMIN A. FOOTE (ALN) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. De- partment of Agriculture, % National Museum of Natural History, MRC 168, Washington, DC 20560-0168, U.S.A. (e-mail: anorrbom @sel.barc.usda.gov); (BAF) Department of Bi- ological Sciences, Kent State University, RO. Box 5190, Kent, OH 44242-0001, U.S.A. Abstract.—The subgenus Footerellia of Neaspilota is described based on N. (F.) retic- ulata, n. sp., from USA: Ohio and Michigan. It is unusual for the tribe Terelliini in having an extensively reticulate wing pattern. It breeds in flowers of Coreopsis tripteris L. (As- teraceae). Distributional data (new records or corrections) are given for three other species of Neaspilota: N. floridana is reported from the Bahamas, the first record of Neaspilota from the Neotropical Region; N. achilleae is recorded from Georgia; and N. vernoniae is reported from Kentucky, but not Newfoundland as previously reported. Updated couplets for the genera of Terelliini are also provided for the most recent key to Nearctic genera of Tephritidae. Key Words: The tribe Terelliini is a predominantly Old World (mostly Palearctic) group of 107 species divided among six genera, four of which have been reported to occur in North America: Chaetorellia Hendel (three intro- duced species); Chaetostomella Hendel (one native species); Neaspilota Osten Sacken (19 native species); and Terellia Robineau-Desvoidy (two native and three introduced species) (Norrbom et al. 1999a, b). The native Nearctic Terelliini have been relatively well studied taxonomically (Foote et al. 1993), especially the largest genus, Neaspilota, which was thoroughly revised by Freidberg and Mathis (1986). We were therefore rather surprised to dis- cover a very distinctive new terelliine spe- cies from the Midwest, which breeds in flowers of Coreopsis tripteris L. (Astera- ceae). It is unusual in having an extensive, reticulate wing pattern; most species of Ter- elliini have banded, spotted, or mostly or Neaspilota, Terelliini, Asteraceae, distribution, Bahamas. Georgia, Kentucky entirely hyaline wings. This species, which we here describe in the new subgenus Neas- pilota (Footerellia) as N. reticulata, n. sp., appears to be the sister group of the other species of Neaspilota. We also take this op- portunity to report distributional informa- tion for three species of Neaspilota and to update the couplets concerning the Terelli- ini in the most recent key to Nearctic genera of Tephritidae (Foote et al. 1993) to incor- porate changes in classification and from the introduction of species and genera. UPDATES TO KEY TO NEARCTIC GENERA OF TEPHRITIDAE AND KEY TO SPECIES OF NEASPILOTA Since the publication of the latest key to genera of fruit flies of America north of Mexico (Foote et al. 1993), three species of the Palearctic genus Chaetorellia have been introduced (C. acrolophi White & Mar- quardt and C. australis Hering for weed VOLUME 102, NUMBER 1 biocontrol, and C. succinea (Costa) acci- dentally), as were two species of Terellia (T. virens (Loew) for weed biocontrol, and T. fuscicornis (Loew) accidentally) (Norr- bom et al. 1999a). The three species for- merly classified in the genus Orellia Robi- neau-Desvoidy were transferred to Terellia by Korneyev (1985), but this change was not included by Foote et al. The following couplets, which can be substituted for cou- plets 18—20 of the generic key of Foote et al. (1993), incorporate these changes. Fig- ure numbers refer to Foote et al. (1993), not the present publication. 18. Posterior orbital bristle distinctly inclinate (fig. 44, a); first flagellomere rounded at apex dorsally; scutum with lyre-shaped dark PEAVY ePAtteLM PS) BS eke tank obey Seo eae — Posterior orbital bristle usually reclinate or absent, if inclinate, first flagellomere with dorsoapical point (Rhagoletis acuticornis) and/or scutum without lyre-shaped pattern (Campiglossa pallidipennis) Scutum with 2 pairs of dorsocentral bristles (one pre- and one postsutural); wing with 4 transverse bands ....... Chaetorellia Hendel (3 introduced Palearctic spp.) — Scutum with | pair of dorsocentral bristles (postsutural); wing pattern variable .... 19 19. Anteroventral margin of gena with a few well-developed bristles larger than adjacent setae (fig. 45, a); wing with 4 transverse bands (fig. 175) 18A. Shh 4 Chaetostomella Hendel (1 sp., western U.S.) — Anteroventral margin of gena with only small, more or less equal sized setae (fig. 44, b); wing pattern variable 20. Node of vein Rs (at fork of R,,; and R,,;) dorsally with | to several setulae (fig. 47, a); scutum with small, circular, dark brown spot at base of acrostichal bristle (in species in N. America); abdomen paired’ dark brownvspots™ =. Peet ees as. MgO OPO athe eee Terellia Robineau-Desvoidy (2 native and 3 introduced Palearctic spp., widespread) — Node of vein Rs usually bare, rarely with 1—2 setulae; scutum without small, circular, dark brown spot at base of acrostichal bris- tle; abdomen yellow, mostly brown, with brown bands, or occasionally (2 spp.) with paired dark brown spots usually with Neaspilota Osten Sacken (20 spp., Alaska, southern Canada, wide- spread U.S., also Mexico and Bahamas) 143 Chaetorellia was recently revised by White and Marquardt (1989), and their key is the best means of identifying the three species introduced to North America. The two recently introduced species of Terellia differ from the other three species in North America (treated under Orellia in Foote et al. 1993) in having entirely hyaline wings. These two species are easily distinguished using the key of Freidberg and Kugler (1989: 166): T. fuscicornis is a large species (wing length 3.6—6.2 mm), with, as its name indicates, the first flagellomere dark brown; T. virens (wing length 2.2—3.8 mm) has entirely yellow antennae. These species also differ significantly in genitalic charac- ters (see Freidberg and Kugler 1989, White 1989). The following couplet can be added to the beginning of the key to species of Neas- pilota in Foote et al. (1993) in order to in- clude N. reticulata. The figure numbers in the second part of the couplet refer to Foote et al. (1993), not the present publication. A. Wing pattern reticulate, predominantly dark with hyaline spots (Fig. 2). Male fore tarsus symmetrical. Female with aculeus relatively broad, less than 4 times as long as wide (Fig. 4A). Midwest; infesting flowers of Coreopsis (Footerellia, n. subgen.) ...... reticulata, n. sp. — Wing predominantly or entirely hyaline, at most with 3 bands or series of spots and some faint smaller spots (fig. 49, 266, 267). Male fore tarsus usually asymmetrical (fig. 275— 282). Aculeus more than 4 times as long as wide. Widespread; hosts various Genus Neaspilota Osten Sacken Footerellia Norrbom and Foote, new subgenus Type species.—Neaspilota reticulata, n. sp. Diagnosis.—The single known species of this subgenus is easily distinguished from other Terelliini, including the other two subgenera of Neaspilota, by its extensive, reticulate wing pattern (Fig. 2), which has more dark than hyaline areas. The largest of the hyaline areas do not extend over more than 2 wing cells each. Many other 144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Characters used in phylogenetic analysis of Footerellia. . Posterior orbital seta—0O) reclinate; 1) inclinate. WN . Scutum with dark, lyre-shaped pattern—O) no; 1) yes. Male fore tarsomere 5—O) symmetrical; 1) asymmetrical, with ‘“‘comb’’, or at least several, short, stout, subequal setulae anteroapically; 2) as state 1, but with anteroapicalmost setula of “‘comb’’ larger than others. . Glans with ‘‘apicodorsal rod’’—0) yes; 1) no. . Spermathecae relatively dark—O) no; |) yes. . Spermatheca shape—O) rounded apically; 1) truncate. Proctiger with lateroventral tuft of setae—O) no; 1) yes. . Distiphallus with apex of basal (non-glans) part with lappet-like projection—O) no; 1) yes. Terelliini have extensive wing patterns, but with either isolated dark spots or distinct bands, broadly delimited by large hyaline areas. Footerellia further differs from other Terelliini, except other Neaspilota, in lack- ing a subapical lobe (“‘apicodorsal rod”’) on the glans of the phallus, and from the other subgenera of Neaspilota in having a sym- metrical male fore tarsus and a broader acu- leus. The type species, N. (F.) reticulata is more likely to be confused with species of Tephritini, such as Euaresta or Neotephri- tis, or Xyphosiini, such as Gymnocarena, which have similar wing patterns. The Te- phritini differ as follows: postocular setae mixed black and white and varying in size; scutal color pattern absent, or if present, not lyre-shaped; and posterior orbital seta usu- ally reclinate or absent (in New World Te- phritini, convergent only in a few species of Campiglossa, which further differ from N. reticulata in having only 2 frontal setae Table 2. Character state distributions in taxa used in phylogenetic analysis. See Table 1 for explanation of character numbers and states. 12345678 Trypetinae OQOO00000 other Tephritinae 00000100 other Terelliini 11001000 N. (Footerellia) reticulata 11000110 N. (Neaspilota) vernoniae 11200110 N. (Neaspilota) floridana 11110110 other Neaspilota, s.str. 11210110 N. (Neorellia) callistigma 11101111 N. (Neorellia) isochela 11001111 other Neorellia 11100111 and the proboscis geniculate). Gymnocar- ena species differ from N. reticulata in hav- ing the posterior orbital seta reclinate and the scutum unicolorous, without the lyre- shaped pattern. Etymology.—The name of this subgenus, which is feminine, is a combination of the last name of Richard H. Foote, in recogni- tion of his vast contributions to the taxon- omy of the Nearctic Tephritidae, and Ter- ellia, the type genus of the tribe to which this subgenus belongs. Remarks.—We conducted a preliminary cladistic analysis of the phylogenetic rela- tionships of Footerellia based on the char- acters listed in Table 1 and the taxa listed in Table 2. For simplicity, only some spe- cies of the other two subgenera of Neaspi- lota were included. The matrix of Table 2 was analyzed using Hennig86 (ie* option), which resulted in four equally parsimonious trees (length 13 steps, consistency index = 0.69, retention index = 0.73). One of these trees is shown in Fig. 1. Characters 3 and 6 may have evolved differently than shown; character 3.2 could have arisen in Neaspi- lota vernoniae and in other Neaspilota spe- cies, and character 6 could have evolved in the Tephritinae and reversed in the other Terelliini. The other three trees differ from Fig. 1 only in the relationships of the spe- cies of Neaspilota (Neaspilota); in one tree the positions of N. vernoniae and N. flori- dana are reversed, and in the other two trees either N. vernoniae or N. floridana forms an unresolved trichotomy with the rest of Neaspilota (Neaspilota) and Neorel- VOLUME 102, NUMBER 1 £ AS) am 2 77) +. Wy) a fs s =e tale -— 140) —= = = = i = = ® 77) Spm Bias oeecr¥ nd = = = 8 = ® We = Lire << rom ro) ® atl ® = £ = : £ = re) O 2 ro) 5 6 fe o2 145 ec 8 © leet Eg 3 Ss > > = Ss = se) S D es = > a ro} ae es ininal ae S &§ & S&S § Saas > aes D o % Dice. @ 2 2 os os = 8S 8 — ee ie 2 Zoya. oO 2 2 Ce 3.0 4 5 3.2 8 3.1 6,17 Fig. 1. Possible phylogenetic relationships of Neaspilota (Footerellia) reticulata. One of four equally par- simonious trees that resulted from cladistic analysis (see text). Character numbers refer to Table 1. lia. In all of these trees, Footerellia was hypothesized to be the sister group of the rest of Neaspilota. We consider these results preliminary, however, because we have not studied the other Terelliini in detail. We did not include some characters used by Freid- berg and Mathis (1986) in the matrix be- cause their polarities could not be resolved. For example, Footerellia possesses several characters that Freidberg and Mathis (1986) 146 considered diagnostic for Neaspilota, s. str.: head more rounded than quadrate, height to length ratio greater than 1.25 and frontofa- cial angle greater than 120°, lower facial margin not strongly projected; frons sparse- ly setulose; arista mostly bare; and aculeus tip with sensilla closely approximated. The number of large anepisternal setae is vari- able. All of these characters are fairly var- iable in Tephritinae, for example, within the genus Terellia, head shape varies consid- erably and the arista varies from micropu- bescent to bare. Freidberg and Mathis (1986) considered the setulose frons a syn- apomorphy for Neaspilota, s. str., but N. (F.) reticulata has considerably fewer se- tulae than the species of that subgenus and the homology of this character is therefore uncertain. Further analysis of the entire Ter- elliini, which was beyond the scope of this study, should be conducted to confirm the phylogenetic position of N. (F.) reticulata. Neaspilota reticulata clearly appears to belong in the tribe Terelliini based on its having the two character states considered synapomorphies for this group by Freidberg and Mathis (1986): posterior orbital seta in- clinate (character 1); and scutum with dark, lyre-shaped pattern (character 2). It also has vein R,,; bare, which Korneyev (in press) considered another synapomorphy of the Terelliini. Like the other two subgenera of Neaspilota, Footerellia differs from other Terelliini in lacking a subapical lobe (“‘ap- icodorsal rod’’) on the glans of the phallus (character 6), which Korneyev (in press) considered an apomorphy that also occurred independently in the other tribes of Tephri- tinae. Footerellia, Neaspilota, s. str., and Neorellia also have relatively dark sper- mathecae (character 7), which is probably a synapomorphy. Footerellia lacks the character state that Freidberg and Mathis (1986) considered a synapomorphy for Neaspilota, s. str. + Neorellia in that the male of N. (F.) retic- ulata does not have asymmetrical fore tarsi as in most other species of Neaspilota (character 3). But it should be noted that PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON there is reversal in this character in one spe- cies of Neorellia (N. isochela) according to Freidberg and Mathis’ hypothesis of rela- tionships, which was confirmed by our analysis. Footerellia also lacks the charac- ter state that Freidberg and Mathis (1986) hypothesized as a synapomorphy for Neo- rellia; the spermathecae are rounded api- cally, not truncate (character 8). The male of N. (F.) reticulata does not possess two apomorphic character states that occur within the subgenus Neaspilota. It does not have a lateroventral tuft of setae on the proctiger (character 4), which occurs in the three species of N. (Neaspilota) other than N. vernoniae, nor does it have the anter- oapicalmost seta of the fore tarsal ‘“‘comb”’ enlarged as in the three species of N. (Neas- pilota) other than N. floridana (Footerellia entirely lacks the comb). Freidberg and Mathis (1986) considered the former char- acter a synapomorphy for the subgenus Neaspilota, and in combination these two characters support this group as monophy- letic in two of the four trees that resulted in our analysis, however, two additional trees with the relationships of N. (Neaspilota) un- resolved were equally parsimonious. Fur- ther study is thus needed to confirm that this subgenus is monophyletic. One additional character used in this analysis is worthy of mention. Footerellia lacks the lappet-like lateral extension of the apex of the basal (non-glans) part of the distiphallus (character 5) as do most other species of Neaspilota. This lobe is present, but variable in size, in several species of Neorellia and is present in most other spe- cies of Terelliini. Korneyev (in press) inter- preted this structure as homologous with the membranous basal lobe of the glans pre- sent in many Trypetinae, but due to its dif- ferent structure, this homology seems un- certain. Korneyev considered this character state to have been present in the ground plans of both the Tephritinae and Terelliini, in which case its loss could be interpreted as a synapomorphy for Footerellia + Neas- pilota (s. str.) + most species of Neorellia, VOLUME 102, NUMBER 1 but our analysis suggests independent evo- lution in some Neorellia and other Terelli- inl. The reticulate wing pattern in N. (F.) re- ticulata is probably autapomorphic, al- though similar patterns occur within many other tribes of Tephritinae and even in some Trypetinae, Phytalmiinae and Blepharoneu- rinae. Other Neaspilota species and other Terelliini have hyaline, spotted, or banded wings. Given the extreme variation in wing pattern that occurs within the Tephritidae it is difficult to determine whether or not the pattern in Footerellia is homologous with similar types of patterns in other taxa. Neaspilota (Footerellia) reticulata Norrbom and Foote, new species Holotype.—? (USNMS51657) USA: Ohio: Portage Co.: Towners Woods Park, 1.0 mile E. Kent, reared from Coreopsis tripteris L., emerged 27 May 1984, B. A. Foote, Biol. note no. 8021. (USNM = Na- tional Musuem of Natural History, Smith- sonian Institution, Washington, DC.) Paratypes.—USA: Ohio: Portage Co.: Towners Woods Park, 1.0 mile E. Kent, reared from Coreopsis tripteris L., emerged lis “eSep 1994.5 v1 8 » with puparium (USNM51656); same, except emerged 30 Sep 1996, 2 6 (USNMS51724-25). Michi- gan: Ingham Co., Dansville St. Wild. Area, 17 Jul 1987, J. Jenkins, Malaise trap baited with CO,, 1 2 (USNMS1655). Description.—Most setae and setulae golden. Head: In lateral view 1.28—1.45 times as high as long; ventral facial margin not strongly produced, angle between frons and face 130—140°. Frons with 2—4 setulae medially; 3 frontal setae, anterior pair sometimes smaller than posterior 2; 2 or- bital setae, posterior seta golden, nearly as large as anterior seta, and strongly inclinate. Face slightly silvery microtrichose. Gena in lateral view 1.0—1.4 times as high as height of first flagellomere. Postocellar seta and postocular setae slightly more whitish than other setae, very slightly lanceolate; pos- tocular setae uniform in color and size. 147 Arista 1.4 times as long as rest of antenna, mostly bare, minutely pubescent at base. Proboscis capitate. Thorax: Scutum with dark “‘lyre-shaped” pattern, obscured by dense microtrichia, but posteromedial yel- low area clearly extended anteriorly to dor- socentral seta; without any additional dark brown spots at bases of setae. Setulae gold- en and acuminate, a few on anterior margin and on postpronotal lobe more whitish and very slightly lanceolate. Microtrichia whit- ish anteriorly, golden posteriorly. Subscu- tellum, mediotergite, most of laterotergite, parts of anepimeron and meron, and most of katepisternum (except margins) also with dark ground color. One pair of dorsocentral setae, aligned with postsutural supra-alar seta. Anepisternum with 1—2 large setae in posterior row. Other thoracic chaetotaxy as described for Neaspilota by Freidberg and Mathis (1986). Male fore tarsus symmettri- cal, without modified setae. Wing (Fig. 2): With extensive reticulate pattern, mostly dark brown with hyaline spots, infuscate ar- eas fainter gray-brown in basal third of wing. Dark brown areas include broad area from pterostigma extended into cell cu,, covering crossvein R-M, and subapically a large area primarily in cells r,,, and r,,; as long as broad and without hyaline spots in its center. Cell r, with two hyaline basal marginal areas extended into cell r,,;, and with 1—2 small subapical hyaline spots. Cell r,,; with round hyaline spot near apex of vein R,,, and a subapical hyaline spot near posterior margin, sometimes fused with spot in cell r,,;. Cell r,,; with 2 subbasal spots (a large one anterior to crossvein DM- Cu and a smaller one slightly more distally) and 2—4 subapical spots. Pterostigma 2.0— 2.5 times as long as wide. Node of Rs and vein R,,; without setulae. Abdomen: Ter- gites entirely yellow, without dark spots. Setulae uniformly golden. Male genitalia: Epandrium and surstyli relatively broad versus high in posterior view (Fig. 3B), nearly circular; lateral surstylus short, strongly mesally curved. Proctiger (Fig. 3A) with setae evenly distributed. Glans 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON RRR AL eae, 2 (Figs. 3C—D) relatively stout, without basal membranous lobe, and without sclerites in vesica; acrophallus short, stout, with sharp subapical turn (Fig. 3C). Female genitalia: Aculeus (Fig. 4A) relatively broad, about 3.3 times as long as wide; tip relatively short and stout, evenly tapered, entire; 2 larger sensilla separated by approximately length of sensillum. 2 spermathecae (Fig. 4B) pear-shaped, dark brown. Etymology.—The name of this species, the Latin adjective reticulata, refers to its wing pattern. Biology.—The second author has reared this species from flowers of Coreopsis trip- teris L. (Asteraceae, tribe Heliantheae). This is the first confirmed record of a ter- elliine species breeding in this genus. Freid- berg and Mathis (1986) listed C. calliopsi- dea (DC.) Gray as a suspected host of Neaspilota wilsoni, but Goeden (1989) con- sidered it a doubtful host. No other terelli- ine species have reported hosts in the tribe Heliantheae. The four species of Neaspilota (Neaspilota), except for two records for N. floridana considered doubtful by Freidberg and Mathis (1986), so far as known breed only in species of Vernonia (tribe Vernon- ieae). Species of Neaspilota (Neorellia) have hosts in several tribes, but breed pre- dominantly in species of the tribe Astereae (Goeden 1989). The North American spe- SE ASaee Wing of Neaspilota (Footerellia) reticulata (Michigan paratype). cies of Chaetorellia, Chaetostomella, and Terellia breed in species of Cardueae. DISTRIBUTIONAL RECORDS FOR NEASPILOTA The previously published distributional data discussed below are from Freidberg and Mathis (1986) or Foote et al. (1993). Neaspilota achilleae Johnson The following records are the first from Georgia and eastern Newfoundland for N. achilleae, which is known from coastal ar- eas from Newfoundland to Alabama. The male from Newfoundland was misidentified as N. vernoniae. CANADA: NEWFOUND- LAND: Terra Nova Nat’]. Park, 6 Jul 1961, C. P. Alexander, 1 6 (USNMS51002). USA: GEORGIA: 14 mi. N Folkston, 10 Apr 1989, J. E. Swan, 1 2 (University of Guelph). Neaspilota floridana \brahim The following records extend the known range of this species, previously reported from Kansas, Missouri, Arkansas, Texas, Louisiana, Pennsylvania, Virginia, North Carolina, Georgia, and Florida. BAHA- MAS: Grand Bahama Island, Freeport, 20— 27 Jun 1987, W. E. Steiner, M. J. & R. Mol- ineaux, Malaise trap in Caribbean pine and palmetto scrub, 1 2 (USNMS51110). USA: Illinois: Bond Co.: 0.5 mi W Dudleyville, VOLUME 102, NUMBER 1 149 D Figs 3% Male genitalia of Neaspilota (Footerellia) reticulata (Ohio paratype). A, Lateral view of epandrium, surstyli, proctiger and base of phallus. B, Posterior view of epandrium and surstyli. C—D, Glans in ventral and lateral views. Bar = 0.10 mm. IS eAugel979.. E. cAs Lisowski,,.1 6 (USNMS51658); Clinton Co.: 1.5 mi NE Al- bers, 12 Aug 1980, E. A. Lisowski, 1 ¢ 1 2 (USNMS51659-60); Monroe Co.: 3 mi ESE New Design, 13 Aug 1980, E. A. Li- sowski, | d6 (USNMS51661). The Bahamas record, notable as the first record of Neas- pilota from the Neotropical Region, ought to be confirmed at the species level by the collection and identification of male speci- mens. The Illinois specimens were retained from larger series in the Illinois Natural History Survey. Neaspilota vernoniae (Loew) This species was previously known from an arc from Kansas, Nebraska and [owa to Michigan to Massachusetts and New Jersey, with one outlying record from Newfound- land which is erroneous. It should be noted 150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Female genitalia of Neaspilota (Footerel- lia) reticulata (Michigan paratype). A, Aculeus. B, Spermathecae. Bar = 0.10 mm. that the distribution map in Foote et al. (1993, map 36) has the color of the dots erroneously reversed for N. brunneostig- mata Doane and vernoniae, and omitted the valid record of vernoniae from Westport Factory, Massachusetts reported by Freid- berg and Mathis (1986). The Newfoundland record was based on the misidentified male of N. achilleae in the USNM listed above. Also in the USNM there is one male of N. vernoniae with the following data: KEN- TUCKY: Fayette Co., 15 Jul 1980, J. Mill- stein (USNMS51662). This record extends the known middle part of the range of this species southward by approximately 400 km. ACKNOWLEDGMENTS We are grateful to John Jenkins for do- nation of the Michigan paratype of N. re- ticulata, to Ed Lisowski for permission to publish the Illinois records of N. floridana, to S. A. Marshall for the loan of the Uni- versity of Guelph specimen of N. achilleae, and to W. N. Mathis, A. Freidberg, N. Van- denberg, and E C. Thompson for their re- views of this paper. L. Rodriguez produced the illustration of the wing and cladogram, and T. Litwak inked the other figures. LITERATURE CITED Foote, R. H., EF L. Blanc, and A. L. Norrbom. 1993. Handbook of the fruit flies (Diptera: Tephritidae) of America north of Mexico. Comstock Publish- ing Associates, Ithaca, xii + 571 pp. Freidberg, A. and J. Kugler. 1989. Fauna Palaestina. Insecta IV. Diptera: Tephritidae. Israel Academy of Sciences and Humanities, Jerusalem, 212 pp. Freidberg, A. and W. N. Mathis. 1986. Studies of Ter- elliinae (Diptera: Tephritidae): A revision of the genus Neaspilota Osten Sacken. Smithsonian Contributions to Zoology 439: iv + 75 p. Goeden, R. D. 1989. Host plants of Neaspilota in Cal- ifornia (Diptera: Tephritidae). Proceedings of the Entomological Society of Washington 91: 164— 168. Korneyev, V. A. 1985. Fruit flies of the tribe Terelliini Hendel, 1927 (Diptera, Tephritidae) of the fauna of the USSR. Entomologicheskoe Obozrenie 64: 626—644. [In Russian; English translation in En- tomological Review 64: 35—55 (1986).] . In press. Phylogeny of the subfamily Tephri- tinae: Relationships of the tribes and subtribes. /n Aluja, M. and A. L. Norrbom, eds., Fruit Flies (Tephritidae): Phylogeny and Evolution of Behav- ior. CRC Press, Boca Raton. Norrbom, A. L., L. E. Carroll, and A. Freidberg. 1999a. Status of Knowledge, pp. 9-47. Jn Thomp- son, FE C., ed., Fruit Fly Expert Identification Sys- tem and Biosystematic Information Database. Myia (1998) 9: 524 pp. Norrbom, A. L., L. E. Carroll, E C. Thompson, I. M. White, and A. Freidberg. 1999b. Systematic Da- tabase of Names, pp. 65-251. Jn Thompson, E C., ed., Fruit Fly Expert Identification System and Biosystematic Information Database. Myia (1998) 9: 524 pp. White, I. M. 1989. A new species of Terellia Robi- neau-Desvoidy associated with Centaurea solsti- tialis L. and a revision of the Terellia virens (Loew) species group (Diptera: Tephritidae). En- tomologists’ Monthly Magazine 125: 53-61. White, I. M. and K. Marquardt. 1989. A revision of the genus Chaetorellia Hendel (Diptera: Tephri- tidae) including a new species associated with spotted knapweed, Centaurea maculosa Lam. (Asteraceae). Bulletin of Entomological Research 79: 453-487. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 151-161 ISCHNOPTERAPION (CHLORAPION) VIRENS (HERBST) (COLEOPTERA: CURCULIONOIDEA: BRENTIDAE: APIONINAE), A PALEARCTIC CLOVER PEST NEW TO NORTH AMERICA: RECOGNITION FEATURES, DISTRIBUTION, AND BIONOMICS E. RICHARD HOEBEKE, ROBERT A. BYERS, MIGUEL A. ALONSO-ZARAZAGA, AND JAMES EF. STIMMEL (ERH) Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853, U.S.A. (e-mail: erh2@cornell.edu); (RAB) Pasture and Watershed Management Research Unit, ARS, U.S. Department of Agriculture, U.S. Regional Pasture Research Laboratory, University Park, PA 16802, U.S.A.; (MAAZ) Depto. de Biodiversidad y Biol- ogia Evolutiva, Museo Nacional de Ciencias ‘Naturales, José Gutiérrez Abascal, 2. 28006 Madrid, Spain; (JFS) Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, PA 17110, U.S.A. Abstract.—Ischnopterapion (Chlorapion) virens (Herbst) is reported for the first time in North America based on collections in 22 counties in Pennsylvania, 5 counties in New York, 3 counties each in Maryland and New Jersey, and 1 county each in western Con- necticut, northern Delaware, and northern Virginia. This immigrant weevil is a pest of clover (Trifolium spp.), with adults injuring the foliage, and larvae mining in the petioles, stems (stolons), root-crowns, and roots. A detailed redescription and a thorough diagnosis of the adult are given to allow its identification and separation from similar Nearctic species, and its biology and seasonal history in the Palearctic region are summarized from the European literature. Key Words: Ischnopterapion (Chlorapion) virens, immigrant weevil, clover pest, system- atics, bionomics The first North American collections of | subfamily Apioninae! [currently placed in the Palearctic apionine weevil /schnopter- apion virens (Herbst), a pest of clover, were made during an inventory to: (1) determine invertebrate species richness and abundance in dairy farm paddocks under different grazing regimes in the northeastern United States; and, (2) to identify invertebrate spe- cies that may potentially affect sustainabil- ity of pasture systems. Soil sampling for this invertebrate survey was conducted by RAB and Gary M. Barker (Agresearch, Hamilton, New Zealand) on 21 Pennsylva- nia farms from 1994—1996. Most of the world’s described taxa in the the Brentidae, see Kuschel (1995) and Lawrence and Newton (1995)] historically have been assigned to the exceptionally large and difficult genus Apion, which in- cludes nearly 1,600 species. In America north of Mexico, more than 150 species of Apion have been recorded (O’Brien and Wibmer 1982). In this paper, we recognize the supraspecific taxa proposed by Alonso- Zarazaga (1990) in which many of the for- ' The classification presented in Kuschel (1995) and Lawrence and Newton (1995) is followed here, with the Apioninae recognized as a subfamily of the Bren- tidae (ERH). 152 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mer subgenera of Apion are elevated to ge- neric rank. Here, we list and map the known distri- butional records of J. virens in the eastern United States; provide characters to identify this newly detected immigrant and to allow it to be distinguished from similar native apionine weevils found in eastern North America; and summarize information on the biology, habits, food plants, and feeding damage of /. virens in its native range. INITIAL DETECTION, ADDITIONAL U.S. RECORDS, AND SPECIMEN DEPOSITION The first specimens of J. virens were tak- en in pitfall traps on a farm near Robesonia (Heidelberg Twp.), Pennsylvania (Berks Co.) in May 1994. During 1994-1997, nu- merous additional specimens of /. virens were collected in pitfall traps set in pad- docks of grazed pastures of 5 other Penn- sylvania counties. Other locality records, many from 1998 and 1999, have become available for Connecticut, Delaware, Mary- land, New York, New Jersey, Pennsylvania, and Virginia. The collection of specimens of I. virens from Connecticut, Maryland, and Virginia was the direct result of those states’ Cooperative Agricultural Pest Sur- vey-Eastern Region (CAPS) activities for 1999. All known eastern U.S. collections of I. virens (mostly by pitfall trapping, and sweep-net or vacuum sampling) are listed below and mapped in Fig. 1. CONNECTICUT: Litchfield Co., East Canaan, 15-VI-1999; Norfolk, 15-VI-1999. DELAWARE: New Castle Co., Newark, 21 & 22-IV-1998; Smyrna, 25-VII-1999. MARYLAND: Cecil Co., Calvert, 22-IV- 1999; Chesapeake City, 22-IV-1999; Con- owingo, 22-IV-1999; Cowentown, Fair Hill NRMA, 22-IV-1999; Elkton, 22-IV-1999; Rising Sun, 22-IV-1999; Warwick, 22-IV- 1999. Harford Co., Dublin, 26-IV-1999. Kent Co., Chestertown, 3-V-1999; Golts, 22-IV-1999; Kennedyville, 3-V-1999; Mill- ington, 22-IV-1999. NEW JERSEY: Hun- terdon Co., Snyder research farm, 30-VI- 1992 and 14 & 21-VII-1992; no precise lo- cality, 3-X-1997. Salem Co., Woodstown, 8-X-1997. Warren Co., 6-X-1997. NEW YORK: Chemung Co., Lowman, 16-VII- 1997. Chenango Co., Earlville, 15-VII- 1997. Orange Co., Westtown, 29-VII-1998. Tompkins Co., Ithaca, 28-VII-1998. Westchester Co., Yorktown Heights, 23- XII-1996. PENNSYLVANIA: Berks Co., Robesonia, V-1994, 1996 and VII, [X-1996, 6-VI-1997. Bucks Co., Quakertown, 1-VI- 1998; Nockamixon State Park, 1-VI-1998. Carbon Co., Normal Square, 20-V-1998; Lehighton, 20-V-1998; Jim Thorpe, 20-V- 1998. Chester Co., Honey Brook, 30-VII- 1997. Columbia Co., Centralia, 8-VI-1998; Numidia, 8-VI-1998. Cumberland Co., Car- lisle, 14-VI-1999; Summerdale, 10-VII- 1998. Dauphin Co., Harrisburg, 18-V-1998. Delaware Co., Swarthmore, 18-VI-1998. Lancaster Co., Kirkwood, 28-VII-1997. Lebanon Co., Myerstown, 6-VIII-1997. Le- high Co., Coopersburg, 1-VI-1998; Fogles- ville, 7-VIII-1997. Luzerne Co., Freeland, 20-VII-1998; Hazelton, 20-VII-1998; White Haven, 20-VII-1998. Lycoming Co., Montgomery, 25-VI-1998. Monroe Co., Appenzell, 7-VII-1998; Brodheadsville, 7- VII-1998; Saylorsburg, 7-VII-1998; Stroudsburg, 7-VII-1998. Montgomery Co., W. of Boyertown, [X-1997. Montour Co., Danville, 26-V-1998. Northampton Co., Bethlehem, 1-VI-1998. Northumberland Co., Dalmatia, 22-V-1998; Malta, 8-VI- 1998. Perry Co., Marysville, 27-VII-1998; Roseglen, 7-VI-1999. Philadelphia Co., Philadelphia, Fairmont Park, 14-VII-1998. Schuylkill Co., Lavelle, 8-VI-1998; New- town, 18-V-1998; Pitman, 8-VI-1998; Schuylkill Haven, 18-V-1998. York Co., Longlevel, 28-V-1998. VIRGINIA: Lou- doun Co., Rte. 15, nr. Gilberts Corner, 30- VI-1999. The first specimens of J. virens, repre- senting the initial detection of this Palearc- tic species in North America, were initially identified by ERH and later confirmed by MAAZ. Voucher specimens are deposited in collections of the following individual and institutions: Cornell University (Ithaca, VOLUME 102, NUMBER 1 =i vewancoy f ‘ SCLARION | - =o cLEARFIELD § CENTRE m ae / JANA j eiANA HANS INO! / = uM coc { prt | ae r _ x - = e | = / = = J é LALLEGHENY } JEFFERSON( z fe} Z ,2vor™ | TUCKER & 8 RANDOLPH 4 | é \ mS ? { geoFORO / SO [eaannei™| ADA” | = e Rigel: NY), USDA-Beneficial Insects Research Laboratory (Newark, DE), Pasture Systems and Watershed Management Research Lab- oratory (University Park, PA), Maryland Department of Agriculture (Annapolis), Pennsylvania Department of Agriculture (Harrisburg), Florida State Collection of Arthropods (Gainesville), Virginia Tech (Blacksburg), National Museum of Natural History, Smithsonian Institution (Washing- ton, DC), University of Connecticut Known geographic range of /schnopterapion virens in the eastern United States. (Storrs), and Miguel A. Alonso-Zarazaga (Museo Nacional de Ciencias Naturales, Madrid, Spain). NATIVE GEOGRAPHIC RANGE Ischnopterapion virens, widely distributed through most of the Palearctic Region, is common in temperate Europe and the Med- iterranean subregion, including the Iberian Peninsula, and in the Nordic countries of northern Europe (Denmark, Finland, Nor- 154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-6. 4, Head and pronotum, female, lateral aspect. 5, Head and pronotum, male, dorsal aspect, showing punctation in part. 6, Elytra, male, dorsal aspect. Scale line for Fig. 2 = 0.5 mm. (Figs. 3—4 after Morris 1990; Figs. 5—6 after Gonget 1997.) way, Sweden) (Gonget 1997). Occurring commonly and abundantly throughout Eng- land, Wales, Ireland, and locally in Scotland (Fowler 1891, Morris 1990), it is recorded also from Siberia, Syria, and northern Africa (Algeria) (Hoffmann 1958, Brito-Castro and Oromi-Masoliver 1986), and the Canary Is- lands (Tenerife and La Palma) (Brito-Castro and Oromi-Masoliver 1986). ADULT REDESCRIPTION The following redescription is adapted from Gonget (1997). Length 1.8—2.6 mm, oblong, slightly convex (Fig. 2). Body Ischnopterapion virens. 2, Adult, male, dorsal aspect. 3, Head and pronotum, male, lateral aspect. black, with aeneous or metallic greenish re- flection, elytron distinctly metallic greenish or greenish-blue, shining, vestiture rather sparse. Male rostrum (Fig. 3) shorter, slight- ly curved, about 1% times as long as pro- notum, pubescent at apex. Female rostrum (Fig. 4) longer, curved, about 1% times as long as pronotum, pubescent only at base. Frons slightly convex, finely and sparsely punctured. Vertex finely and sparsely punc- tured apically and transversely striate and shining basally. Eye prominent, medium- sized to large and almost round. Pronotum (Fig. 5) broader than long, broadest behind VOLUME 102, NUMBER 1 middle, slightly convex longitudinally, dif- fusely, finely and shallowly punctured. Bas- al fovea of pronotum (Fig. 5) rather distinct but short. Elytron (Fig. 6) oblong to elon- gate with sides rounded, broadest just be- hind middle, slightly convex longitudinally and somewhat depressed on disc, separately rounded at apex. Elytral striae moderately strongly punctured, first stria deeper on disc than others. Striae joined at apex as fol- lows: 1-F2--9; 3+4+-7; and. 5+8+6: Fine vestiture arranged in single row on each in- terstice, with one specialized seta at apical third of 7th interstice. Legs black with fem- ora having greenish or bluish metallic re- flection. Flight wings fully developed or ru- dimentary. Male genitalia illustrated by Alonso-Zarazaga (1990: 120). DIAGNOSIS Ischnopterapion virens can be separated from the majority of North American apionine weevils by the following combi- nation of characters: rostrum curved in both sexes, ventral surface of head with low sub- ocular ridges, dorsal margin of scrobe not dentiform, prothorax moderately constrict- ed behind middle, lacking basal flange, el- ytra apically neither expanded nor with deep pits, male metasternum lacks median tubercle near posterior margin, male tibiae without hooks (non-mucronate), legs dark, tarsal segments | longer than wide, tarsal segments 3 moderately large, strongly bi- lobed, tarsal claws with acute basal tooth. Using these characters, in Kissinger’s (1968) key to North and Central American apionid weevils, this newly detected species keys to couplet 27. It differs from the mem- bers of Pseudapion sensu Kissinger [Apion disparatum and A. varicorne species groups, most of the species transferred to genus Kissingeria by Alonso-Zarazaga (1990)] by the black legs and antennae (legs in part and/or antennae yellow in both groups of species in Kissingeria), only ely- tral interstria 7 with one specialized seta (one specialized seta on elytral interstria 7 and 9 in Kissingeria), striae at apex joining 155 7 with 3+4 and 8 with 5+6 (joining 7+8 in Kissingeria). It differs from Apion pan- amense Sharp (still placed in Apion sensu lato) by the shorter rostrum in both sexes, ca. 1.25 < length of pronotum in the male and ca. 1.50 in the female (ca. 1.42 in the male and ca. 1.70 in the female in Apion panamense) and by the latter having only one specialized seta on interstria 9. From the group of species assigned to Ceratapion by Kissinger (but not belonging to this ge- nus, cf. Alonso-Zarazaga 1990) and which, being absent from his key, should key also to the same couplet, J. virens is distinguish- able by the union of striae at the apex (as above), and the lack of a spine on the Ist mesotarsomere in the male (present in the so-called Ceratapion) and genital features. Following the criteria explained in Alon- so-Zarazaga (1990), Kissingeria and the so- called Ceratapion sensu Kissinger belong to the tribe Oxystomatini subtribe Trichap- iina. Ischnopterapion virens belongs to the subtribe Synapiina of the same tribe, which includes three other Palearctic genera, not yet known to be in the Nearctic. The two subtribes formerly present in the Nearctic Region were: Trichapiina (including the large and heterogeneous genus Trichapion Wagner, Kissingeria Alonso-Zarazaga, and several species groups still placed in Apion sensu lato, which will probably need genera of their own after a thorough revision and study of their relationships) and Oxysto- matina (represented in the Nearctic Region by two genera: Mesotrichapion Gyorffy (with the single species Mesotrichapion (Loborhynchapion) cyanitinctum (Fall)) and Eutrichapion Reitter, embracing three species in two different subgenera, Eutri- chapion s. str. and Leconteapion Alonso- Zarazaga, namely: Eutrichapion (Eutri- chapion) viciae (Paykull) (= Apion alas- kanum Fall), E. (Leconteapion) cavifrons (LeConte) and E. (L.) huron (Fall). All the Nearctic members of Oxystomatina have a median tubercle on the male metasternum and key separately at couplet 13 in Kissin- ger’s key. Even if the median tubercle in 156 these species is overlooked, members of Eutrichapion have black elytra and one spe- cialized seta on interstriae 7 and 9, and members of Mesotrichapion subgenus Lo- borhynchapion, although having metallic elytra, have simple claws, the frons is strio- late and the male mesorostrum is strongly lobed in dorsal view. In addition, /. virens is different from all these species by its first elytral stria being more deepened than the rest on the disc. Mesotrichapion (L.) cyan- itinctum is recorded only from Manitoba and Quebec, and its host plant appears to be Astragalus (Fabaceae). Host PLANTS AND DAMAGE Plants of the family Fabaceae (= Legu- minosae) are the principal hosts of adult and larval J. virens, with a decided prefer- ence shown for species of clover (Trifoli- um). These weevils are responsible for two types of feeding injury to clover: adults feed on the foliage (leaflets), whereas lar- vae feed or mine in the petioles, stems (sto- lons), root-crown, and possibly taproots. Ischnopterapion virens larvae mine the stems of Trifolium pratense and generally bore downward towards the root-crown and roots during development (Stein 1965). In one instance, root damage was observed in red clover, 7. pratense var. spontaneum (Stein 1965). Adult feeding creates small (ca. 0.5 mm/), elongate punctures in the leaf tissue (Wiech and Clements 1992: 438). Feeding sites vary depending on the age of the wee- vil. For example, newly emerged (first sea- son) adults (in July) exhibit a preference for feeding on the apical portion of clover leaf- lets, while ‘‘older’’ adults (14—21 days min- imum since emergence) use the basal por- tion of leaflets (see Stein 1965: 393, fig. 3). Adult feeding damage to white clover fo- liage is illustrated in Fig. 7. Although J. virens is generally consid- ered to be a minor pest of clover crops in Europe (Balachowsky 1963), it can be a major pest causing significant damage. Be- cause white clover (T. repens) is grown PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON widely in mixture with grasses for animal feed, it is one of the most important herbage legumes in Britain (Wiech and Clements 1992). Chief among the invertebrates dam- aging white clover in southeastern England are apionine weevils, especially J. virens (Wiech and Clements 1992). In Bavaria in 1930 and 1931, a serious local and heavy infestation by J. virens occurred in red clo- ver (T. pratense); larvae, by tunnelling in the taproot and root-crown, reduced plant growth so that fodder and seed yields were much decreased (Andersen 1932). In April and May 1926 in upper Austria, fields of red clover were heavily damaged by I. vi- rens, with as much as 80% of this forage crop damaged (Werneck 1930). The following species of Trifolium have been recorded as host plants of J. virens: T. campestre Schreb. (Brito-Castro and Oromi-Masoliver 1986); 7. pratense L. (Fowler 1891, Werneck 1930, Markkula and Myllymaki 1957, Stein 1965, 1968); T. alexandrinum L. (Frauenfeld 1866, Delas- sus 1936, Dieckmann 1977); T. arvense L. (Hoffmann 1958); T. incarnatum L. and T. fragiferum L. (Dieckmann 1977); and T. re- pens L. (Dieckmann 1977, Stein 1968, Wiech and Clements 1992). BIOLOGY, SEASONAL HISTORY, AND HABITAT The following account of the biology and seasonal history of /. virens is a summary drawn from field and laboratory observa- tions of various investigators: Werneck (1930), Andersen (1932), Bovien and Jor- gensen (1934, 1936), Markkula and Myl- lymaki (1957, 1958), Kokorin (1964), Scherf (1964), and Stein (1965, 1968, 1972a, b). Detailed studies on the mor- phology, biology, seasonal history, and be- havior of J. virens are provided by Stein (1965, 1968). Adult weevils overwinter in the soil near host plants or among plant debris at the base of their hosts (Trifolium spp.) and re- appear in the spring, generally as early as late February—early March. After weevils VOLUME 102, NUMBER 1 Figs. 7—10. 7, Adult feeding damage to white clover leaflet. 8, Larva tunneling in white clover stem. 9, Pupa exposed in white clover stem. 10, Pupa in white clover stem, enlarged. emerge from their overwintering sites, in- tensive feeding begins, followed by a spring mating period, generally from April to June. Oviposition generally begins in mid- April to early May and continues into late May-early June. The female uses her man- dibles to chew a cavity in the petiole or stem, lays a single egg, and seals off the egg cavity with a mixture of excrement and abdominal secretions, probably to protect the egg and to prevent its desiccation. After about 7-8 days, first-instar larvae hatch and chew long mines in the stems, mostly downward. Larvae move downward into the stem where actual feeding will begin (Fig. 8). After about four weeks, a larva completes its development and pupates in a cavity in the lower part of the clover stem (Figs. 9, 10), just under the epidermis, di- rectly above the root. The pupal stage lasts about 7 days, and first generation adults Ischnopterapion virens, adult and larval feeding damage to white clover and immature stages. emerge, harden and become fully colored in about 2—3 days. Preimaginal (egg, larval and pupal) development takes place from May through much of August. Adults emerge throughout the summer, but gener- ally in July and August. Newly emerged weevils feed intensively on the leaves for about four weeks, followed again by a non- feeding period of about 40 days, and then a dispersal, or migratory phase. Adults fly mainly during late summer and autumn (August-November), with a lower temper- ature threshold of 10°C (Stein 1972a), after which they feed until the beginning of win- ter. Mating takes place in early to mid-au- tumn. Thus, there are two periods of mat- ing, one in autumn and one in spring. Dur- ing a weevil’s lifespan, at least three feed- ing periods occur: one directly after adult emergence in summer, a second from Sep- tember to the beginning of the overwinter- 158 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ing period, and a third after the overwin- tering period in spring (Stein 1965). There is one generation annually. Egg development requires approximately 8 days, with a range of 6—12 days (Stein 1968). Bovien and Jorgensen (1936) re- ported a range of 14-18 days for egg de- velopment. Three larval instars occur in I. virens. Larval development takes about 27— 28 days (Stein 1968), whereas Bovien and Jorgensen (1936) noted that 28-30 days were necessary for larval development. The average duration from egg to the pupal stage is approximately 34 days, with little or no difference between males and fe- males, and a range of 30—39 days. The av- erage length of the pupal stage is 7 days. Adults can survive as long as 360 days. Un- der laboratory conditions, each mated fe- male is capable of laying a maximum of 90 to 166 eggs; however, for the majority of beetles in the field the number is much less, approximately 40—45 eggs per female (Stein 1968). The seasonality and habits of 7. virens in the Old World vary according to the inves- tigator. In upper Austria, for example, eggs are laid on the root-crown or stems of red clover, with first instar larvae found in Sep- tember; early instars overwinter in the tap- roots, make their way into the heart of the plant in spring, and pupate in July (Werneck 1930). In Germany, larvae of J. virens live in the tap root or root collar and overwinter as adults in the soil (Andersen 1932). Based on laboratory rearings in Denmark, Bovien and Jorgensen (1934) found that field-col- lected adults laid eggs in May in the mid- ribs of leaves, that eggs hatched in early June, and that larvae later mined in the leaf petioles, eventually penetrating into the roots of smaller plants, but continuing to feed in the stems of larger plants. Markkula and Myllymaki (1957) found that /. virens females oviposited in the stems of red clo- ver from late May to early August. Ischnopterapion virens is associated gen- erally with its leguminous host plants on slopes, along roadsides, in coastal pastures and meadows, and in grassy fields and cul- tivated clover fields (Gonget 1997). In Brit- ain, this weevil is found “‘in a wide variety of open and grassy biotopes, including sand dunes and agricultural land” (Morris 1990). SAMPLING AND NATURAL HISTORY OBSERVATIONS IN PENNSYLVANIA To gain a better understanding of popu- lation numbers of /. virens at the original collection site in Robesonia (Heidelberg Twp.), PA, one of us (JFS) conducted a sampling regime of white and red clover. Weekly, 1 m’, vacuum samples, taken from April 1998 through mid-July 1999, have yielded adult weevils nearly constant- ly. Even during the winter months (Decem- ber 1998 through March 1999), adults were taken in vacuum samples; the 1998—99 win- ter was unusually mild, with temperatures seldom falling below freezing and with lit- tle to no snow cover for the majority of this period. Numbers of adults taken in the weekly samples actually increased during the winter months, as compared to those collected during the dry, warm months of summer and early autumn of 1998. From the surprisingly high numbers taken weekly during the winter, the population climbed somewhat during the spring, until new gen- eration adults began to emerge. At this point, sample sizes increased dramatically to a high of 180 adults from a 1 m? area on June 15, 1999. Winter and early spring sample numbers averaged ca. 32 adults/m?. From the June 15 peak, the number of adults present in the weekly samples de- clined drastically to a low of O in the July 13, 1999 sample. The extreme drought of the 1999 summer has all but eliminated suc- culent host plant material in the sampling area. The July 13 sample is the first such sample to be devoid of adults since sam- pling began in April 1998. On numerous occasions in 1998 and 1999, we have observed the feeding habits of larvae of J. virens on white and red clo- ver at Robesonia, PA. Mature white clover plants spread by creeping, above ground, VOLUME 102, NUMBER 1 prostrate stems (stolons) that also root at the nodes (Uva et al. 1997). On May 21, 1998, we (ERH, JFS) split open, by means of a razor-blade knife, many stems (stolons) of white clover plants growing adjacent to grazed pastures and found that the majority were mined by larvae of /. virens (Fig. 8). Cut plant stems with larvae were brought into the lab and adults were reared to con- firm their identity. In addition to the dis- covery of many mature larvae, we also ob- served pupae (Figs. 9, 10) inside these stems (stolons). Random observations of larval feeding in the stems of white clover revealed extensive consumption of plant tissue. Larvae feed upon the central, pithy areas of the lower prostrate stems (stolons). During peak lar- val incidence (late May to early June), many stolons exhibited evidence of feeding by larvae over the entire length of the sto- lon. In such instances, larvae were found at approximately 2.5 cm intervals over the length of the infested stolon, and those sto- lons with late-stage larvae and pupae were often in a state of semi-collapse, contained large areas of browned and dessicating tis- sue, and often exhibited small holes through the epidermis of the plant. On 26 May 1998, RAB found larvae and pupae in the stems of red clover and noted nearly 100% infestation of these plants, with many dying and browning stems. In spite of extensive larval feeding dam- age to the stems (stolons) of white clover in grazed pastures at Robesonia, PA, these plants nonetheless appeared vigorous and healthy. In sharp contrast, the red clover plants appeared to be more heavily impact- ed by the larval feeding of J. virens, espe- cially the above ground stems (RAB obser- vations). On white clover, adults feed mostly in interveinal areas of the foliage, producing oval to elongate-oval holes. In the summer of 1999, JFS attempted to quantify the leaf area fed upon in relation to the total leaf surface area, using computer imaging and measurement packages. Percentage of leaf 159 surface devoured ranged from 0% to a high of 21.25%, and averaged 2.64% of leaf sur- face sampled during a 5 week period (sam- ples collected weekly from June 15 through July 13;.1999),. The pest status of /. virens and its long- term impact on white and red clover in the eastern United States are equivocal and will remain so until further field studies are con- ducted. ACKNOWLEDGMENTS We are grateful to the following individ- uals for providing us with supplemental collections of /. virens in the Northeast: W. H. Day (USDA, Beneficial Insects Research Laboratory, Newark, DE) for Delaware, J. Ingerson-Mahar (Rutgers University, New Brunswick, NJ) for New Jersey, and M. C. Thomas (Florida State Collection of Arthro- pods, Gainesville, FL) for New York. Dis- tributional data acquired through the Co- operative Agricultural Pest Survey (CAPS) activities in the following states is grateful- ly acknowledged: Connecticut (Donna E]- lis), Maryland (Dick Bean), and Virginia (Eric Day). We also thank the following in- dividuals for their assistance in the field and for collecting pitfall trap samples in Penn- sylvania, New York, and Vermont during the past 4 years: Steve LaMar, John Ever- hart, Ruth Haldeman, Andy Paolini, Kevin Nelson, Jeremy Everhart, Karen Jackson, Allison Henry, Chrissa Rose, Sharyn Leach, Andy Davis, Mark Giacomin, Amy Kellerman, Chante Gordon, Cindy Long, Brian Dougherty, Mike Hutchinson, Jim Gavlik, Kanesha Williams, Michelle Lorek, Heather Lagoda, Matthew Hartman, and Claudia Violette. Chad M. Petrovay and Justin Newell are acknowledged for their extensive survey findings of J. virens in Pennsylvania. We greatly appreciate the critical review of an earlier draft of this pa- per by A. G. Wheeler, Jr. (Clemson Uni- versity), and also the comments provided by two anonymous reviewers. 160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Alonso-Zarazaga, M. A. 1990. Revision of the su- praspecific taxa in the Palaearctic Apionidae Schoenherr, 1823 (Coleoptera, Curculionoidea), 2. Subfamily Apioninae Schoenherr, 1823: Intro- duction, keys and description. Graellsia 46: 19— 156. Andersen, K. T. 1932. Spitzmauschen (Apion virens Hbst. und Apion seniculum Kirby) als Kleescha- dlinge. Zeitschrift fiir Pflanzenkrankheiten 42: 18— 28. Balachowsky, A. S. 1963. Entomologie appliquée a Vagriculture. Tome I, Coléopteres. Second Vol- ume—Phytophagoidea (suite et fin) (Chrysomeli- dae—Curculionidae—Attelabidae—Scolytidae et Platypodidae). Masson et C'* Editeurs, Paris. 1391 Pp. Bovien, P. and M. Jorgensen. 1934. Orienterende Un- dersogelser over Angreb af Snudebiller (Apion) 1 Kloverhoveder. Tidsskrift Planteavl 40: 375—404. . 1936. 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Kissinger, D. G. 1968. Curculionidae subfamily Apioninae of North and Central America with reviews of the world genera of Apioninae and world subgenera of Apion Herbst (Coleoptera). Taxonomic Publications, South Lancaster. IN. 555 pp. Kokorin, A. N. 1964. The pests of seed clover (in Russian). Trudy Vsesoyuznogo Nauchno-Issle- dovtel’skogo Instituta Zashchity Rastenii 20: 61— 63. Kuschel, G. 1995. A phylogenetic classification of Curculionoidea to families and subfamilies, pp. 5— 33. In Anderson R. S. and C. H. C. Lyal, eds. Biology and phylogeny of Curculionoidea: Pro- ceedings of a symposium convened at the XVIII International Congress of Entomology, Vancouver, Canada, July 3—9, 1988. Memoirs of the Ento- mological Society of Washington 14. Lawrence, J. EF and A. EK Newton, Jr. 1995. Families and subfamilies of Coleoptera (with selected gen- era, notes, references and data on family-group names), pp. 779-1006. /n Pakaluk J. and S. A. Slipinski, eds. Biology, phylogeny, and classifi- cation of Coleoptera: Papers celebrating the 80th birthday of Roy A. Crowson. Museum 1 Instytut Zoologii PAN, Warszawa. Markkula, M. and S. Myllymaki. 1957. Investigation into the oviposition on red and alsike clover and alfalfa of Apion apricans Herbst, A. assimile Kir- by, A. flavipes Payk., A. seniculus Kirby and A. virens Herbst (Col., Curculionidae). Annales En- tomologici Fennici 23: 203-207. . 1958. On the size and location of the eggs of Apion apricans Herbst, A. assimile Kirby, A. fla- vipes Payk., A. seniculus Kirby, and A. virens Herbst (Col., Curculionidae). Annales Entomolo- gici Fennici 24: 1-11. Morris, M. G. 1990. Orthocerous weevils. Coleoptera, Curculionoidea (Nemonychidae, Anthribidae, Urodontidae, Attelabidae and Apionidae). Hand- book for the Identification of British Insects 5: 1— 108. O’Brien, C. W. and G. J. Wibmer. 1982. Annotated checklist of the weevils (Curculionidae sensu lato) of North America, Central America, and the West Indies (Coleoptera: Curculionoidea). Mem- oirs of the American Entomological Institute 34: 1-382. Scherf, H. 1964. Die Entwicklungsstadien der mitte- leuropdischen Curculioniden (Morphologie, Bionomie, Okologie). Abhandlungen. Sencken- bergische Naturforschende Gesellschaft 505: 1— 335: Stein, W. 1965. Das Frassverhalten von Apion virens Herbst. (Col., Curculionidae) an Rotklee (Trifoli- um pratense). Zeitschrift fiir Angewandte Ento- mologie 55: 389-399. . 1968. Zur Biologie, Morphologie und Ver- haltensweise von Apion virens Hrbst. (Col., Cur- culionidae). Oecologia 1: 49-86. . 1972a. Untersuchungen zum Flug und Flu- gverhalten von Curculioniden. Zeitschrift fiir An- gewandte Entomologie 71: 368—375. . 1972b. Tages- und jahresperiodische Schwan- kungen der Frassaktivitat von Apion virens Hrbst. (Col. Curculionidae). Zeitschrift fiir Angewandte Entomologie 70: 296-302. Uva, R. H., J. C. Neal and J. M. DiTomaso. 1997. Weeds of the Northeast. Cornell University Press, Ithaca, New York. 397 pp. Werneck, H. L. 1930. Das griinliche Spitzmauschen VOLUME 102, NUMBER 1 161 (Apion virens Hbst.). Neu als verheerender Sché- | Wiech, K. and R. O. Clements. 1992. Studies on the dling der Rotkleebestande (Wurtzelhals und Sitona spp. and Apion spp. weevils feeding on Herttzteil). Zeitschrift fir Angewandte Entomo- white clover foliage at a site in S. E. England. logie 16: 585-591. Journal of Applied Entomology 113: 437—440. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 162-169 THE NEW WORLD HAIRSTREAK GENUS ARAWACUS KAYE (LEPIDOPTERA: LYCAENIDAE: THECLINAE: EUMAEINI) ROBERT K. ROBBINS Department of Entomology, National Museum of Natural History, Smithsonian Insti- tution, Washington, DC 20560-0127 U.S.A. (e-mail robbins.robert@nmnh.:si.edu) Abstract.—Even though Arawacus Kaye is widely used in works on North American butterflies, its characterization has been inconsistent and its included species have not been listed. Subterminally constricted tips of the papillae anales are proposed to be the best way to characterize Arawacus. They are consistent with the higher classification of the Thereus Section of the Eumaeini, to which Arawacus belongs. They are consistent with patterns of oviposition specificity. This characterization re-confirms that Polyniphes Kaye and Dolymorpha Holland are junior synonyms of Arawacus. Tigrinota Johnson is synonymized with Arawacus, new synonym. All nomenclaturally available specific names that belong to Arawacus are listed. Key Words: Thereus, Rekoa, Contrafacia, ceae The generic name Arawacus Kaye was rarely used until 1981. The original descrip- tion was based on superficial wing pattern elements of the South American type spe- cies, A. aetolus (Sulzer) (Kaye 1904). As a result, besides being cited in nomenclatural lists (e.g., Comstock and Huntington 1959— 1964, Eliot 1973), Arawacus was used only for species with wing patterns similar to the type (Brown and Mielke 1967; Robbins 1980, 1981). But after Miller and Brown (1981) made Arawacus a senior synonym of Dolymorpha Holland, whose type spe- cies ranges into the United States, the use of Arawacus in general butterfly works mushroomed (e.g., Scott 1986, Bailowitz and Brock 1991, Stanford and Opler 1993, Cassie et al. 1995). Arawacus was characterized by mor- phology of its male genitalia. Clench (1961: 212) had delimited Dolymorpha primarily by a “‘ventro-lateral conical, acuminate pro- cess at middle” of the valves and by the Dolymorpha, Polyniphes, Tigrinota, Solana- two terminal cornuti, “‘one of which has an abrupt, larger disc-like terminal expansion with peripheral teeth” (Fig. 1). He later made Dolymorpha a junior synonym of Ar- awacus using the same two character states (an unpublished manuscript that is being deposited in the Archives of the Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, U.S.A.). Miller and Brown (1981), who received a copy of the manu- script after Clench’s death, published this synonymy in their catalog without expla- nation. Arawacus has been characterized in dif- ferent ways since 1991. Because Clench mistakenly lumped five distinct species in the A. aetolus complex (Robbins, Hudson, and Clench, in preparation), he did not re- alize that the type species of Arawacus lacks both male genitalia traits that he had proposed (Fig. 2). Consequently, when Robbins (1991: 3) briefly treated Arawacus as an outgroup of Rekoa Kaye, he provi- VOLUME 102, NUMBER 1 Figs. 1-2. ry er ENT Titel TER ase | a f ¥ Male genitalia in ventral aspect with penis in lateral aspect, posterior end towards top of the page. 1, Arawacus jada, with arrows pointing to the posterior pointing processes on the valves and to the flattened cornutus with terminal teeth. 2, A. aetolus, which lacks the posterior pointing processes and has an arrowhead shaped cornutus. sionally re-characterized it by “*(1) papillae anales constricted subterminally, and (2) larvae that feed on the foliage of Solanum (Solanaceae),’’ but did not illustrate the first character or list the included species. He confirmed the synonymy with Dolymorpha, added Polyniphes Kaye as a second junior synonym of Arawacus, and placed Arawa- cus in the Thereus Section of the Eumaeini. Johnson (1992, 1993) described and re- vised Tigrinota for species that possess both traits that Robbins had proposed for characterizing Arawacus, but Johnson did not note either trait or mention that Tigri- nota might be closely related to Arawacus. Further, he treated Dolymorpha as a distinct genus without mentioning the previous syn- onymy with Arawacus. (Note: Some copies of Johnson [1993] have Johnson and Kroenlein listed as the authors. I follow the citation in Lamas et al. [1995] with Johnson as the sole author). The purpose of this paper is to stabilize usage of Arawacus and to provide a generic taxonomy for revision at the species level. Specifically, this paper shows that the con- 164 stricted papillae anales occur in the type species of Arawacus, Dolymorpha, Polyni- phes, and Tigrinota, but not in other genera of the Thereus Section, to which Arawacus belongs. It is further shown that Tigrinota was delimited by characters that are incor- rectly described or that occur in other gen- era of the Thereus Section. Finaily, the spe- cific taxa that belong to Arawacus are listed for the first time. MATERIALS AND METHODS This study was based upon the approxi- mately 3,250 specimens belonging to the Thereus Section genera Arawacus, Rekoa, Thereus Hiibner, Contrafacia Johnson, and the ‘‘Thecla”’ ligurina species group (an un- described genus whose species are placed, following convention, in Thecla FE, a genus that does not belong to the Eumaeini, Eliot 1973) in the National Museum of Natural History, Smithsonian Institution, Washing- ton, DC, USA (USNM). Included were all species that belong to these genera as de- limited by Robbins (1991). Preparation of genitalia for examination using light and scanning electron microscopes follows the procedures in Robbins (1991). The male and female genitalia of all species in the Thereus Section were dissected (except for those few species known from only one sex), including more than 120 genitalic dis- sections of those species placed in Arawa- cus in this paper. PREVIOUS RESULTS The Thereus section.—Robbins (1991) proposed two phylogenetic hypotheses in- volving Arawacus. First, Arawacus, Rekoa, Thereus, Contrafacia, and the ‘“‘Thecla”’ ligurina group form the Thereus Section of the Eumaeini because they share a unique process of the vinculum abutting the brush organs, when present (illustrated in Robbins 1991). Second, the first three genera form a monophyletic group. The ductus semin- alis arises from a pouch of the posterior corpus bursae. This pouch is dorsal of the ductus bursae, sclerotized laterally, and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON membranous dorsally (again illustrated in Robbins 1991). This classification is a framework within which the monophyly of Arawacus can be assessed. Type species nomenclature.—Synonymy of the type species of Arawacus, Contra- facia, and Tigrinota is a bit confused. The type of Arawacus is Papilio linus Fabricius, which is a junior synonym of P. aetolus Sulzer (Comstock and Huntington 1959— 1964). The type of Contrafacia is C. mex- icana Johnson 1989, a species that is indis- tinguishable from C. imma (Prittwitz 1865) (Robbins 1991). The type of Tigrinota is Thecla ellida Hewitson, which occurs from northern Venezuela to central Argentina. Its wing pattern varies slightly over this wide range with virtually no geographical varia- tion in genitalic structures (Robbins, in prep.), contrary to the treatments in John- song(i9972, 1993): RESULTS Monophyly and Arawacus.—The papil- lae anales (“‘ovipositor valves’’) were ex- amined for 57 of the approximately 63 spe- cies that belong to the Thereus Section. Fe- males are not known for the other 6 species. The tips of the papillae anales (ventral as- pect) for the type species of Arawacus (Fig. 3), Dolymorpha (Fig. 4), Tigrinota (Fig. 5), and Polyniphes (Fig. 6) have a subterminal constriction, which might alternately be de- scribed as terminally expanded. The tips of the papillae anales of Rekoa (Figs. 7—9) are notched, not subterminally constricted. The papillae anales of Thereus (Figs. 10—11) are similar to those of Rekoa, but contain a sclerotized patch that is otherwise unre- ported in the Eumaeini (Robbins 1991). The tips of the papillae anales of Contra- facia (Fig. 12) and the ‘‘Thecla” ligurina group (Fig. 13) are regularly tapered. Al- though the papillae anales in the Thereus Section vary in shape, only those of Ara- wacus have a conspicuous subterminal con- striction. Larval food plant specificity—Among the eumaeine genera, larval feeding on the VOLUME 102, NUMBER 1 165 Figs. 3-13. Papillae anales in ventral aspect, posterior end towards top of the page. 3, Arawacus aetolus; arrow points to the subterminal constriction. 4, A. jada. 5, A. ellida. 6, A. dumenilii. 7, Rekoa meton. 8, R. palegon. 9, R. marius. 10, Thereus lausus; arrow points to sclerotized patch. 11, 7. praxis; arrow points to sclerotized patch. 12, Contrafacia imma. 13, ““Thecla” lyde. leaves of Solanum (as opposed to flowers) occurs only in Arawacus, but the larvae of three Arawacus species have recently been reported to eat plants in the Compositae and Leguminosae (Robbins, in press). Leaf feeding on Solanum has been recorded for 11 of the 17 Arawacus species (noted in the list at the end of this paper), including the 166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON type species of Arawacus, Dolymorpha, Ti- grinota, and Polyniphes (Guppy 1904, 1914; Bourquin 1945; Kendall 1975; Rob- bins, in press). However, larvae of A. ellida (Hewitson), which have been reared re- peatedly from the leaves of Solanum, and A. binangula (Schaus) were both recently recorded eating flowers of Compositae—a larval food plant used in the related genus Rekoa—and the larvae of A. tarania were recorded on legumes (Brown 1993). Con- sequently, although larval feeding on the leaves of Solanum is widespread in Ara- wacus and is unreported in other Eumaeini, it cannot be used to delimit the genus. The larvae of other Thereus Section spe- cies primarily use plants in families other than the Solanaceae as larval food (Robbins 1991, in press). Rekoa larvae are extremely polyphagous on buds, flowers, and shoots of plants in 16 families, including Solana- ceae, but most records are in the Compos- itae, Leguminosae, and Malvaceae. In con- trast, larvae of Thereus specialize on Lor- anthaceae, with one species eating Mal- pighiaceae and Chrysobalanaceae. Contrafacia larvae are recorded eating plants in the Compositae and Leguminosae, and the only food plant record in the “‘The- cla”’ ligurina group is in the Erythroxyla- ceae. Monophyly and Tigrinota.—The only al- ternative to the proposed classification in this paper is that of Johnson (1992, 1993). It is the purpose of this section to assess the evidence supporting the monophyly of 77- grinota. The diagnosis of Tigrinota (Johnson 1992, 1993) is similar in both works, but difficult to interpret. The diagnosis of the wings refers to the hindwing “with concen- tric bands,”’ but bands on the wings of these species are not concentric. It refers to the ‘“‘ovate brand,’ but the type species lacks an ovate brand or other scent patch. It men- tions alternating bands on the hindwing, but these also occur in Rekoa palegon (Cramer) and R. malina (Hewitson). Since Rekoa is a close relative of Arawacus (Robbins Fig. 14. valves of Arawacus ellida in ventral aspect, posterior end towards the top of the page. Note the lack of rims, Scanning electron micrograph of the microtrichia, and ‘‘caudo-lateral sculptures.” 1991), these bands would appear to be a symplesiomorphy and are questionably di- agnostic. The diagnosis of the male genitalia of 77- grinota in Johnson (1992: 186) is cited in its entirety because it is confusing. “*Ven- trally, valvae appearing as paired and rather smoothly sclerotized oblongate lobes sepa- rated by a thin transparent fissure which at each end, shows sclerotized ridges forming (1) prominent rims about the bilobed area and (2) caudo-lateral sculptures of the val- val terminus from which emerge clusters of robust microtrichia.” The valves of the type species of Tigrinota are illustrated in ven- tral aspect (Fig. 14). There are no rims. There are no microtrichia, robust or not. Perhaps setae were mistaken for microtri- chia, but if so, their occurrence on the ven- tral valves is true for virtually all eumaeines (e.g., Clench [1961] characterized Callo- VOLUME 102, NUMBER 1 phrys Westwood by the unusual lack of se- tae on the valve tips). It is unclear to what the ‘“‘caudo-lateral sculptures” refer (Fig. 14). Johnson (1993: 2) repeats these char- acters and adds “‘brush organ occurrences diffential (sic)”’, but it is unclear what this means or how it might be diagnostic. As best I can tell, those parts of this diagnosis that are accurate do not differentiate 7igri- nota from most eumaeines. The diagnosis of the female genitalia is similarly confusing (Johnson 1992, 1993). The description (I presume of the ductus bursae) with a “‘transparent neck”’ refers to the ductus bursae of most Rekoa and Ara- wacus (Robbins 1991; Robbins, Hudson, and Clench, in preparation) and is not di- agnostic. The description of a simple “‘sub- cordate incised posterior cavity (abbreviat- ed sipc in the diagnosis) of the 8" tergite”’ (probably referring to the shape of the 8" abdominal tergum) is difficult to interpret. However, the shape of this tergum is similar in Tirgrinota and some Rekoa, such as R. zebina (Hewitson), so it is unlikely to be diagnostic. DISCUSSION Taxonomy.—The genus Arawacus ap- pears to be monophyletic, characterized by the shape of the papillae anales. Although it is not desirable to delimit Arawacus by only one character, this classification is con- sistent with the hierarchical classification of the Thereus Section, and other morpholog- ical traits support the monophyly of Ther- eus and Rekoa (Robbins 1991). These char- acterizations of Arawacus, Thereus, and Re- koa are reasonably consistent with patterns of larval plant use. For these reasons, this classification appears to be the best option for promoting a stable generic nomencla- ture in the Thereus Section. Recognition of Tigrinota is unlikely to be a reasonable alternative. Its use leaves Ar- awacus and Polyniphes uncharacterized and would consequently destabilize the generic taxonomy of the Thereus Section. There is little, if any, evidence to support the mono- 167 phyly of Tigrinota. Either its diagnostic traits are incorrect, such as the concentric hindwing bands and rims on the valves, or appear to be symplesiomorphic, such as the alternating hindwing bands and “‘transpar- ent neck”’ of the ductus bursae. Provisional classification of Arawacus.— The status of Polyniphes as a synonym (Robbins 1991) is confirmed. The status of Dolymorpha as a synonym (Miller and Brown 1981), which was not recognized in Johnson (1992), is also confirmed. Tigri- nota is synonymized with Arawacus, new synonym. Johnson (1992, 1993) presented a clas- sification of almost half the specific names listed below. He did not assess intraspecific geographical variation and reported “‘diag- nostic’’ morphological traits, such as con- centric hindwing bands and rims on the valves, that do not exist. Further, my ex- amination of more than 120 genitalic dis- sections and detailed comparison of andro- conia and wing patterns from throughout the range of each species failed to confirm most of his conclusions. Rather than await detailed species level revisions, I present an alternate classification below that is intend- ed to be a working hypothesis for species level revisions. Although it is unusual to present a tentative classification, such as this one, the extensive reporting of charac- ters that do not exist (Johnson 1992, 1993) is perhaps a more unusual circumstance that creates the need for a more reasonable working classification. As characterized in this paper, Arawacus contains 39 available specific names repre- senting 17 biological species. There is also one undescribed species (Robbins, Hudson, and Clench, in preparation). The female genitalia of all have been examined except for A. euptychia, for which no females are known. However, A. euptychia is placed in Arawacus because its male genitalia are barely distinguishable from those of A. du- menilti and A. tadita. Those species that have been reared from plants in the Solanaceae, Compositae, or 168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Leguminosae are designated respectively in bold with S, C, or L. 1. Arawacus togarna (Hewitson, 1867) 2. Arawacus lincoides (Draudt, 1917)—S 3. Arawacus aetolus (Sulzer, 1776)—S Papilio linus Fabricius, 1776 Papilio amelia Herbst, 1804 4. Arawacus separata (Lathy, 1926)—S Thecla paraguayensis Lathy,1926 5. Arawacus aethesa (Hewitson, 1867) 6. Arawacus sito (Boisduval, 1836)—S Thecla phaenna Godman and Salvin, 1887 Arawacus mexicana D’ Abrera, 1995 7. Arawacus leucogyna (Felder and Felder, 1865)—S Thecla phaea Godman and Salvin, 1887 8. Arawacus meliboeus (Fabricius, 1793)—S Jolaus eurisides Hiibner, 1823 Thecla barrensis Rosa, 1936 9. Arawacus jada (Hewitson, 1867)—S 10. Arawacus ellida (Hewitson, 1867)—S,C Thecla toba Hayward, 1949 Tigrinota perinota Johnson, 1992 Tigrinota jennifera Johnson, 1992 Tigrinota catamarciana Johnson, 1993 Tigrinota chaosa Johnson, 1993 11. Arawacus hypocrita (Schaus, 1913) 12. Arawacus dolylas (Cramer, 1777)—S Pseudolycaena spurius Felder and Felder, 1865 Thecla dolosa Staudinger, 1888 Thecla pallida Lathy, 1930, 13. Arawacus dumenilii (Godart, 1824)—S Thecla argiva Hewitson, 1877 Thecla obscura Staudinger, 1888 Thecla carteri Weeks, 1906 14. Arawacus euptychia (Draudt, 1921) 15. Arawacus tadita (Hewitson, 1877)—S Thecla datitia Jones, 1912 16. Arawacus binangula (Schaus, 1902)—C Thecla bolima Schaus, 1902 17. Arawacus tarania (Hewitson, 1868)—L Thecla atrana Schaus, 1902. ACKNOWLEDGMENTS Mary Clench kindly sent me her late hus- band’s unpublished manuscript on part of Arawacus. Some information reported in this paper was discovered during a joint project with P. Hudson on species level sys- tematics of the A. aetolus complex. For help with the illustrations, I thank S. Braden, W. Brown, V. Malikul, and G. Venable. For reading and commenting on the manuscript, I am grateful to J. Glassberg, G. Lamas, J. Scott, and A. Warren. LITERATURE CITED Bailowitz, R. A. and J. P. Brock. 1991. Butterflies of Southeastern Arizona. Sonoran Arthropod Stud- ies, Tucson, 342 pp. Brown, K. S. Jr. and O. H. H. Mielke. 1967. Lepidop- tera of the central Brazil plateau. I. Preliminary list of Rhopalocera (continued): Lycaenidae, Pier- idae, Papilionidae, Hesperiidae. Journal of the Lepidopterists’ Society 21: 145-168. Brown, Jr., K. S. 1993. Theclinae endemic to the cer- rado vegetation (central Brazil), p. 152. In New, T. R, ed., Conservation Biology of Lycaenidae (Butterflies), IUCN, Gland, Switzerland. Bourquin, F 1945. Mariposas Argentinas. Edicion del autor, Buenos Aires, 211 pp., 192 photographs, 3 maps. Cassie, B, J. Glassberg, P. Opler, R. Robbins, and G. Tudor. 1995. North American Butterfly Associa- tion (NABA) Checklist & English Names of North American Butterflies, First Edition. North American Butterfly Association, Morristown, New Jersey, 43 pp. Clench, H. K. 1961. Tribe Theclini, pp. 177—220, In Ehrlich, P. R., and A. H. Ehrlich, How to Know the Butterflies, Brown Company, Dubuque, Iowa, 262 pp., 525 figs. Comstock, W. P. and E. I. Huntington. 1959-1964. An annotated list of the Lycaenidae (Lepidoptera: Rhopalocera) of the Western Hemisphere. Journal of the New York Entomological Society 66: 103— 118, 67: 59-95, 163-212, 68: 105-122, 176-186, 232-240, 69: 54-58, 105-118, 157-176, 191— 200, 70: 39—46, 100-118, 177-179, 71: 45-57, 115-119, 189-197, 262-264, 72: 120—130, 173- 192. Eliot, J. N. 1973. The higher classification of the Ly- caenidae (Lepidoptera): A tentative arrangement. Bulletin of the British Museum (Natural History) Entomology 28(No.6): 371—S05. Guppy, J. 1904. Notes on the habits and early stages of some Trinidad butterflies, pp. 225-228, 2 plates. Appendix to: W. J. Kaye. A catalogue of the Lepidoptera Rhopalocera of Trinidad. Trans- actions of the Entomological Society of London, pp. 159-228, 2 pl. . 1914. Notes on the habits and early stages of VOLUME 102, NUMBER 1 some Trinidad butterflies, pp. 76-79. Appendix to: W. J. Kaye. Fauna of Trinidad. Part 1. A catalogue of the Lepidoptera Rhopalocera of Trinidad. (But- terflies of Trinidad). The Agricultural Society of Trinidad and Tobago, Paper No. 558, 79 pp. Johnson, K. 1992. Genera and species of the Neotrop- ical ‘‘elfin’’-like hairstreak butterflies (Lepidop- tera, Lycaenidae, Theclinae). Reports of the Mu- seum of Natural History, University of Wisconsin (Stevens Point) 22: 1—279. . 1993. Hairstreak butterflies of the genus 77- grinota (Lepidoptera, Lycaenidae, Theclinae). Re- ports of the Museum of Natural History, Univer- sity of Wisconsin (Stevens Point) 37: 1—21. Kaye, W. J. 1904. A catalogue of the Lepidoptera Rho- palocera of Trinidad. Transactions of the Ento- mological Society of London, pp. 159-228, 2 pl. Kendall, R. O. 1975. Larval foodplants for seven spe- cies of hairstreaks (Lycaenidae) from Mexico. Bulletin of the Allyn Museum of Entomology 24: 1-4. Lamas, G., R. G. Robbins, and W. D. Field. 1995. Bibliography of butterflies. An annotated bibli- ography of the neotropical butterflies and skippers (Lepidoptera: Papilionoidea and Hesperioidea). Atlas of Neotropical Lepidoptera. 124: 463 pp. 169 Miller, L. D. and E M. Brown. 1981. A catalogue/ checklist of the butterflies of America north of Mexico. Memoirs of the Lepidopterists’ Society, No. 2. 280 pp. Robbins, R. K. 1980. The lycaenid “false head” hy- pothesis: historical review and quantitative anal- ysis. Journal of the Lepidopterists’ Society 34: 194-208. . 1981. The “false head” hypothesis: predation and wing pattern variation of lycaenid butterflies. The American Naturalist 118: 770-775. 1991. Evolution, comparative morphology, and identification of the eumaeine butterfly genus Rekoa Kaye (Lycaenidae: Theclinae). Smithsoni- an Contributions to Zoology No. 498, 64 pp. . In press. Larval Food Plants of the Neotrop- ical Eumaeine Hairstreaks (Lycaenidae: Thecli- nae). Memoir No. 5, Lepidopterists’ Society. 246 Pp. Scott, J. A. 1986. The Butterflies of North America, A Natural History and Field Guide. Stanford Uni- versity Press, Stanford, California, 583 pp., 64 pl. Stanford, R. E. and P. A. Opler 1993. Atlas of Western USA Butterflies, Including Adjacent Parts of Can- ada and Mexico. Published by the authors, Denver and Fort Collins, Colorado, 275 pp. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 170-177 NEW MEGOPHTHALMINE LEAFHOPPERS (HOMOPTERA: CICADELLIDAE) FROM MEXICO, WITH A KEY TO NEW WORLD SPECIES D. V. NovIKOV AND C. H. DIETRICH Center for Biodiversity, Illinois Natural History Survey, 607 E. Peabody Dr., Champaign, IL 61821, U.S.A. (e-mail: dietrich @ mail.inhs.uiuc.edu) Abstract.—Four new Mexican species of the leafhopper subfamily Megophthalminae are described and illustrated: Brenda gracilicauda, B. bidentata, B. licrocerca, and Tiaja leonensis. These species extend the known range of both New World megophthalmine genera considerably southeast. A key to New World Megophthalminae is provided. Key Words: The subfamily Megophthalminae com- prises 44 species of small, beetlelike leaf- hoppers with facial ocelli, carinate frontal sutures, coarsely punctate integument and, often, vestigial hind wings. Species are re- corded from Europe, Africa, the Middle East, and western North America (Metcalf 1962, Oman et al. 1990). Most megophthal- mine species are apparently restricted to coastal or montane habitats, where they oc- cur in dense, low-growing vegetation, and are seldom collected (Curtis 1833, Edwards 1894, Ribaut 1952, Oman 1941, 1972, Lin- navuori 1972, 1973, Sawbridge 1975, Lo- dos and Kalkandelen 1981, Gill and Oman 1982, Van Stalle 1983a, b). Recent vacuum collecting (MaCleod et al. 1994) by C. H. Dietrich on low-growing vegetation among rock outcrops in the Si- erra Madre Occidental of Sinaloa and Du- rango states, Mexico, yielded specimens of three undescribed species of Brenda Oman, a previously monotypic genus endemic to North America. Specimens of an undescri- bed Mexican species of Tiaja Oman, the only other known North American mego- phthalmine genus, were found in the insect collection of the Field Museum of Natural History. Cicadellidae, leafhopper, morphology, distribution In this paper, we provide a revised key to the New World Megophthalminae and describe and illustrate the four new Mexi- can species. Methods for preparing genita- lia follow Oman (1949). Nomenclature for leg chaetotaxy follows Rakitov (1998: AV = anteroventral; AD = anterodorsal; PV = posteroventral; PD = posterodorsal). Spec- imens are deposited in the insect collections of the Illinois Natural History Survey, Champaign, IL, USA [INHS] and Field Museum of Natural History, Chicago, IL, USA [FMNH]. Types of previously de- scribed species were not examined. KEY TO GENERA AND SPECIES OF NEW WORLD MEGOPHTHALMINAE (modified from Oman 1941) 1. Hind wing fully developed. Head with crown broad and short, median length not greater than length next to eye (Fig. 1) .... Brenda Oman, 2 — Hind wing vestigial. Head with crown dis- tinctly longer medially than next to eye (Fig. IO) Fo at:.. Sheree presser Gh ae Tiaja Oman, 5 Forewing with pits inconspicuous, visible only along veins on basal half, membrane densely shagreen. Male anal appendage with T-shaped preapical hook; style without elon- gate lateral arm. Female sternum VII shallow- ly emarginate B. arborea (Ball) N VOLUME 102, NUMBER 1 171 Figs. 1-13. 1-3, Brenda bidentata. 1, Habitus. 2, Head, anterior view. 3, Distal portion of female abdomen, ventral view. 4—7, B. gracilicauda, male genitalia. 4, Genital capsule, lateral view. 5, Style and connective, ventral view. 6-7, Aedeagus, posterodorsal and lateral views, respectively. 8-12, B. bidentata. 8, Female second valvula. 9-10, Aedeagus, posterior and lateral views, respectively. 11, Male genital capsule, lateral view. 12, Style and connective, ventral view. 13, Tiaja leonensis, female second valvula. 7/2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Forewing with circular pits numerous, con- spicuous throughout length, at least along veins, membrane glabrous distally. Male anal appendage without T-shaped hook; style (Fig. 5) with lateral arm at least half as long as apophysis. Female sternum VII distinctly pro- duced mediallya (Rigs 3) eens cr eie 3 Aedeagal shaft compressed, in lateral view (Fig. 7) more than 2X as wide as when viewed posteriorly, in posteroventral view (Fig. 6) attenuated, not wider than %4 maximal width of base, without paired projections api- Cally; 3.25 5 eas ees B. gracilicauda, n. sp. Aedeagal shaft flattened anteroposteriorly, in posteroventral view maximal width more than Ys; maximal width of base, apex with paired jarmoreresyes) (IRIS. 8), IS))Saloc ob clo Bo oo 9 0c 4 Margins of aedeagal shaft in posteroventral view (Fig. 9) parallel, apex truncate, with pair of shont. acute Jaterall pProceSSes\ 9.7.) 1) Pin Seis Pere Lag PECL eeu B. bidentata, n. sp. Aedeagal shaft in posteroventral view (Fig. 15) oval, distad of gonopore forming slender stem branching into pair of triangular pro- cesses B. licrocerca, n. sp. Forewing veins Cu and A1 not distinctly el- evated, claval suture not sulcate opposite fork Ol ReandgN lt reese ee eta et Suen ome rake 6 Forewing veins Cu and A1 distinctly elevat- ed, claval suture sulcate opposite fork of R PIO E C pgnetass cleat Gokenel a mrdon Cunia © uaged once. 7 Posterior margin of female sternum VI sub- laterally produced into pair of bluntly pointed divergent triangular projections, thus forming broad V-shaped medial emargination; sternite VII evident only as broad flap medially, al- most completely covered by sternum VI ... T. mexicana (Ball) Female sternum VI truncate, subequal in length to sternum VII; sternum VII with pos- terior margin narrowly emarginate medially (Fig. 18) T. leonensis, n. sp. Length > 4 mm. Crown short, ratio of width to middle length greater than 3.0 ........ 8 Length < 4 mm. Crown long, ratio of width tommiddlelensthilesssthany2385" 5 -)-1- ara 10 Style with apophysis acuminate. Female ster- num VII without U-shaped emargination... 9 Style with apophysis expanded preapically. Female sternum VII with broad U-shaped emargination T. montara Oman Style very broad at base of apophysis, with well-developed preapical lateral lobe; anal ap- pendage tridentate. Female sternum VII with wide V-shaped emargination comprising en- tire posterior margin T. arenaria Oman Style slender throughout length, without dis- tinct preapical lateral lobe; anal appendage bi- dentate. Female sternum VII with posterior margin truncate and usually slightly deflexed medially T. friscana (Ball) 10. Anal hook with either 2 or 4 points apically. Female sternum VII with pair of rounded lobes laterad of narrow U-shaped medial GUAIGTITMAMON oo peuos ove ob bencsenne 11 — Anal hook with 3 points apically. Female ster- num VII without U-shaped emargination and lobes, often partially concealed ........ 12 11. Anal hook with 2 points apically ........ T. insula Sawbridge — Anal hook with 4 points apically T. californica (Ball) 12. Aedeagus in lateral view with preatrium elon- gate; style with width just proximad of preap- ical lateral lobe much less than length of apophysis. Female sternum VII various... 13 — Aedeagus in lateral view with preatrium short; style with width proximad of preapical lateral lobe subequal to length of apophysis. Female sternum VII very short, often partially concealedie eae eee T. interrupta (Ball) 13. Male with both preapical points on anal ap- pendage recurved in lateral view. Distribu- tion: Coastal Southern California T. ventura (Oman) — Male anal appendage with only first ventral preapical point recurved. Distribution: Santa Cruz Island, CA ... 7. cruzensis (Gill & Oman) Brenda gracilicauda Novikov and Dietrich, new species (Figs. 4—7) Description.—Measurements (mm): Male: length 2.8, pronotum width 1.03, head width 1.12. Female: length 2.8, pron- otum width 1.03, head width 1.17. Color. Variable, mottled with shades of reddish brown to black; mesoscutum dark brown with pale posterior corners; scutellum dark brown; forewing translucent, darker basally, with whitish areas at apex of each anal vein and clavus, ground color forming reticulate pattern on membrane. Texture. Circular pits numerous and conspicuous on head, pron- otum and forewing in basal half, only bor- dering veins distally. Head (as in Figs. 1— 2): Beak attaining anterior margins of hind coxae; clypellus flat, parallel sided medial- ly, rounded basally, apex slightly emargin- ate, exceeding apical margin of gena. Face weakly convex, with 5 shallow depressions: VOLUME 102, NUMBER 1 1 pair mesiad of antennal pits, 1 pair be- tween eyes and ocelli, | between ocelli. Ca- rina formed by frontal sutures obsolete me- dially; gena broad, deeply emarginate ven- trad of eye, section between antennal pit and lorum short and protruding; antennal pit moderately deep, ledge well developed, concealing antennal base in anterior view; antennal flagellum as long as width of fron- toclypeus between antennal pits; ocelli on face closer to eyes than to each other; crown very short, broadly depressed be- tween parallel anterior and posterior mar- gins; transition to face prominent, at about right angle. Thorax: Pronotum (Fig. 1) slightly convex, with 2 shallow depressions anterolaterally; posterior corners of mesos- cutum slightly elevated; scutellum weakly convex. Forewing vein R 3-branched, R, originating slightly distad of initial fork of R, 2 r-m and 3 m-cu crossveins present; hindwing as illustrated by Oman (1949) for B. arborea, except r-m crossvein present. Prothoracic femur with anteroventral preap- ical gibbosity; metathoracic femur with 2 apical macrosetae (ADI, PD1); tibial ma- crosetal formula (PD:AD:AV) 4—5:3—4:4; row PV with one large preapical macroseta and several smaller setae more proximad. Male genitalia: Subgenital plate in ventral view parallel sided, abruptly tapered api- cally. Style (Fig. 5) with lateral arm more than half as long as apophysis; apophysis with toothlike, transverse, ventral ridge at midlenth. Connective (Fig. 5) slightly short- er than maximum width. Aedeagus (Figs. 6—7) U-shaped, with long base; shaft mod- erately compressed, slender, sides parallel, gonopore dorsal subapical, apex in lateral view tapered evenly and acutely distad of gonopore, curved slightly anteroventrad. Segment X ventrolateral appendage (Fig. 4) with subapical projection ventrally, apex in lateral view tapered and slightly curved posterodorsad. Female: Sternum VII (Fig. 3) subequal to other sterna in length, trans- versely rugose, rugae becoming less distinct basally, posterior margin medially with convex lobe comprising % width of ster- 173 num. Second valvula (Fig. 8) slender, dorsal and ventral margins almost parallel basally, apex tapered symmetrically, bearing smaller teeth and having longer dentate section on ventral margin than that of B. arborea (Da- vis: 1975: Fig. 9D): Material examined.—Holotype 6 la- beled: ““MEXICO: Sinaloa/rt. 40, 4 km W. El Palmito/el. 2,000 m, 24 Oct 1995, C.H. Dietrich, lot #95-042; HOLOTYPE/Brenda gracilicauda/Novikov & Dietrich.”’ Other material: 12, same data, lot #95-046 [both INHS]. Note.—The name of the species is an ad- jective formed by combining two Latin words, gracilis (slender) and cauda (tail), referring to the shape of the aedeagus. Brenda bidentata Novikov and Dietrich, new species (Figs. 1-3, 8-12) Description.—Measurements: Male: length 3.0 + 0.1; pronotum width 1.07 + 0.04; head width 1.21 + 0.04. Female: length 3.1 + 0.1; pronotum width 1.22 + 0.08; head width 1.09 + 0.05. Characters as described for B. gracilicauda, except male genitalia as follows: aedeagus (Figs. 9—10) with shaft flattened anteroposteriorly, with two short oblique apical projections, gonopore apical; connective (Fig. 12) rela- tively narrow; segment X ventrolateral ap- pendage (Fig. 11) without preapical projec- tion ventrally, acutely tapered apically. Fe- male as described for B. gracilicauda. Material examined.—Holotype 4d la- beled: ““MEXICO: Durango/rt. 40, 6 km NE El Salto/26 Oct 1995/C.H. Dietrich, lot #95-063; HOLOTYPE/Brenda/bidentata/ Novikov & Dietrich.”” Other material: 2d, 62, same data, lots #95-059, 95-061, 95- 062, 95-063 [all INHS]. Notes.—One male specimen lacks cross- vein m-cu, on both forewings. The name of the species refers to the toothlike apical projections on the aedeagus. 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Brenda licrocerca Novikov and Dietrich, new species (Figs. 14-17) Description.—Measurements: Length 3.0, pronotum width 1.12, head width 1.27. General appearance like that of the 2 pre- viously described species, but differing in the male genitalia as follows: aedeagus (Figs. 15-16) with shaft flattened antero- posteriorly, with pair of triangular apical processes on slender stem, shaft in posterior aspect oval, at widest point slightly narrow- er than base, gonopore dorsal subapical; connective (Fig. 17) almost identical to that of B. bidentata; segment X ventrolateral ap- pendage (Fig. 14) similar to that of B. bi- dentata, but curved posterad at right angle preapically. Material examined.—Holotype d la- beled: “MEXICO: Durango/E La Escondi- da/el. 2680 m, 8 Nov 1995/C.H. Dietrich, lot #95-105”’ [INHS]. Note.—The left forewing of the holotype has vein R 4-branched and 2 s-crossveins. The name of the species combines two Greek words, likros (antler) and kerkos (tail), and refers to the antlerlike processes at the tip of the aedeagus. Tiaja leonensis Novikov and Dietrich, new species (Figs. 13, 18—25) Description.—Measurements: Male: length 3.0 + 0.2; pronotum width 0.95 + 0.02; head width 1.18 + 0.04. Female: length 3.7; pronotum width 1.11; head width 1.35. General appearance character- istic of the genus. Circular pits numerous and conspicuous on dorsum and face. Col- oration variable, the following areas more darkly pigmented, in order of decreasing in- tensity: antennal pit, anterolateral corner of scutellum, venter of thorax and abdomen, dorsum under wing, pronotum mesiad of eye anteriorly, frontoclypeus posterad of ca- rina, crown mesad of eye, center of crown apically, apex of frontoclypeus, ocellus pit, forewing towards costal area. Head (Figs. 19-20): Beak attaining hind coxae; clypel- lus slightly convex, oval, distal margin emarginate, considerably exceeding margin of gena; frontoclypeus weakly convex pos- teriorly, flat adjacent to the edge of frontal carina, carina prominent laterally, obsolete medially; face dorsad of frontal carina with 2 deep lateral and 1 shallow medial depres- sions; gena strongly emarginate ventrad of eye, elevated between lorum and antennal pit; lorum slightly convex; antennal pit deep, ledge fully developed, concealing an- tennal base in anterior view; antennal fla- gellum longer than width of frontoclypeus between pits; ocelli on face in deep pits, slightly closer to eyes than to each other; crown produced medially, ratio of width to length 2.8, with medial shallow depression; transition to face prominent, forming 60° angle. Thorax: Pronotum (Fig. 19) consid- erably narrower than head, shallowly con- cave laterally, lateral margins parallel; pos- terior corners of mesoscutum prolonged and slightly elevated, scutellum small, weakly convex. Forewing with claval suture more or less level with Cu and A, apical portion with inconsistent number of crossveins; anal veins obsolete distally. Hindwing ves- tigial. Metathoracic femur with 2 apical ma- crosetae (AD1, PD1); tibial macrosetal for- mula (PD:AD:AV) 6:6:5; row PV with 5 preapical macrosetae, distal seta larger than others. Male genitalia: Pygofer (Fig. 21) projected posterodorsally forming hoodlike structure with T-shaped process apically (Fig. 22); subgenital plate in ventral view slightly tapered, apex rounded; style (Fig. 23) curved laterad distally, apical section attenuated, tip slightly extended beyond plates, lateral subapical surface rugose; connective elongate, subrectangular. Aedea- gus (Figs. 23—24) with shaft compressed, length of shaft subequal length of base; gonopore dorsal, subapical; tip of shaft acutely tapered distad of gonopore. Seg- ment X ventrolateral appendage (Fig. 21) with transverse double-pointed apical pro- cess, similar to that of 7. montara Oman, but with dorsal tooth comparatively smaller VOLUME 102, NUMBER 1 a ara temen mus ERR AARC Pennie na STN Su Zl 24 Figs. 14-25. 14-17, Brenda licrocerca, male genitalia. 14, Genital capsule, lateral view. 15—16, Aedeagus, posterior and lateral views, respectively. 17, Style and connective, ventral view. 18—24, Tiaja leonensis. 18, Distal portion of female abdomen, ventral view. 19, Habitus. 20, Head, anteroventral view. 21, Male genital capsule, lateral view. 22, Detail of posterodorsal process of pygofer, dorsal view. 23-24, Aedeagus, lateral and posterior views, respectively. 25, Style and connective, ventral view. 176 and ventral tooth longer and narrower. Fe- male: Sternum VII (Fig. 18) with posterior margin narrowly emarginate to half sternum length. Second valvula (Fig. 13) similar to that of B. bidentata but with dorsal preap- ical angle more pronounced. Material examined.—Holotype ¢ la- beled: ““MEX.:N.L.; Galeana/ Cerro Potosi, 10,500ft./VI:26-28:1969/leg. S. & J. Peck; FM(HD) #69-60/pine forest/carrion traps” [FMNH]. Other material: 23, 192, same data [FMNH, INHS]. Note.—The name of the species refers to the type locality in Nuevo Leon, Mexico. DISCUSSION The three new Brenda species closely re- semble each other in shape and color pat- tern and are also very similar externally to B. arborea, but differ from B. arborea in their smaller size, more conspicuously punctate, yet less densely granulose (sha- greened) integument, and the more strongly emarginate margin of the gena below the eye. The male genitalia of the new species are very distinctive, differing markedly from those of B. arborea in the shape of the style, segment X appendages and ae- deagus. Examined females of the new spe- cies differ from those of B. arborea in the shape of sternum VII and the second val- vula, but no discrete morphological char- acters were found to distinguish them from each other. The new Brenda species are unique among New World Megophthalmi- nae in having cheliform styles, a possible synapomorphy. However, such styles are common among species of certain genera of Agalliinae (e.g., Agallia Curtis, Agalliopsis Kirkaldy) and, in addition to the similarities in head morphology, leg chaetotaxy, and the segment X appendages, suggest a close relationship between the two subfamilies. Tiaja leonensis appears to be related to T. mexicana, known only from two females collected in the Sierra Madre Occidental, Chihuahua, Mexico. Unlike other species of Tiaja, T. leonensis, and T. mexicana lack a distinct groove near the base of the claval PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON suture in the forewing. The new species dif- fers from other Tiaja for which males are known in the shape of posterodorsal pro- jection of male pygofer, style, apex of anal hook, and female sternum VII. Of those species, T. leonensis most closely resembles T. insula Sawbridge, recorded from Santa Barbara, Channel Islands, California. Discovery of the new species described herein extends the known range of both New World megophthalmine genera con- siderably to the southeast. Brenda arborea (Ball), the only previously known species in the genus, is apparently restricted to the coastal hills of California, west of the Sierra Nevada. Of the nine previously known spe- cies of Tiaja, seven are apparently restricted to Pacific coastal habitats ranging from southern British Columbia to northern Mexico. Because they occur in dense, low- growing vegetation, these leafhoppers are difficult to collect by conventional methods (i.e., sweeping). The remarkable success of preliminary vacuum collecting in Mexico suggests that the Mexican megophthalmine fauna may be considerably richer than available specimens would indicate. ACKNOWLEDGMENTS We are grateful to Paul Peterson, De- partment of Botany, Smithsonian Institu- tion, for facilitating the collection of spec- imens of the new Brenda species, to Phil Parillo for the loan of specimens from the Field Museum, and to G. Levin, S. Mc- Kamey, R. Rakitov, D. Voegtlin, and an anonymous referee for contructive criticism of the manuscript. This work was support- ed, in part, by National Science Foundation Grant DEB-9870187. LITERATURE CITED Curtis, J. 1833. Characters of some undescribed genera and species indicated in the “Guide to an arrange- ment of British Insects.”” Entomologist Magazine 1: 186-199. Edwards, J. 1894. The Hemiptera-Homoptera of the British Islands 3: 65—96, pls. 9-12. Gill, R. J. and P. Oman. 1982. A new species and new distributional records for megophthalmine leaf- VOLUME 102, NUMBER 1 hoppers, genus Tiaja (Homoptera: Cicadellidae). Entomography 1: 281—288. Linnavuori, R. 1972. Revision of the Ethiopian Cica- dellidae (Hom.), Ulopinae and Megophthalminae. Suomen Hyonteistieteellinen Aikakaukirja 38: 126-149. . 1973. Two new species of the genus Odomas Jac. (Homoptera Cicadellidae, Megophthalminae) from Cameroon. Journal of Natural History 7: 121-123. Lodos, N., and A. Kalkandelen. 1981. Preliminary list of Auchenorrhycha with notes on distribution and importance of species in Turkey. VII. Family Ci- cadellidae: Ulopinae, Megopthalminae [sic], Led- rinae, Macropsinae and Agallinae [sic]. Tiirkiye Bitki Koruma Dergisi 5: 215—230. Metcalf, Z. P. 1962. General catalogue of the Homop- tera. Fascicle VI. Cicadelloidea. Part 5. Ulopidae. United States Department of Agriculture, Agri- cultural Research Service, Washington, D.C., vil = Ol pp: Oman, P. W. 1941. Revision of the Nearctic Mego- phthalminae (Homoptera: Cicadellidae). Journal of the Washington Academy of Sciences 31: 203— 210. . 1949. The Nearctic leafhoppers (Homoptera: Cicadellidae), a generic classification and check 177 list. Memoirs of the Entomological Society of Washington 3, 253 pp. 1972. A new megophthalmine leafhopper from Oregon, with notes on its biology and be- haviour (Homoptera: Cicadellidae). Journal of En- tomology, (Series B) 41: 69-76. Oman, P. W., W. J. Knight, and M. W. Nielson. 1990. Leafhoppers (Cicadellidae): A bibliography, ge- neric check-list, and index to the world literature 1956-1985. C.A.B. International Institute of En- tomology, Wallingford, U.K., 368 pp. Rakitov, R. A. 1998. On differentiation of cicadellid leg chaetotaxy (Homoptera: Auchenorrhyncha: Membracoidea). Russian Entomological Journal 6: 7-27. Ribaut, H. 1952. Homopteres Auchenorhynques. I (Jassidae). Faune de France 57: 1—474. Sawbridge, J. R. 1975. Tiaja insula, a new mego- phthalmine leafhopper from the Santa Barbara Channel Islands (Homoptera: Cicadellidae). Pan- Pacific Entomologist 51: 268-270. Van Stalle, J. 1983a. On some new and interesting West African Ulopinae and Megophthalminae (Homoptera, Cicadellidae). Suomen Hyonteistie- teellinen Aikakauskirja 49: 75-78. . 1983b. On some new and interesting East Af- rican Megophthalminae. Revue de Zoologie Af- ricaine 97: 304-312. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 178-182 NUCLEOTIDE SEQUENCE AND RESTRICTION SITE ANALYSES IN THREE ISOLATES OF KASHMIR BEE VIRUS FROM APIS MELLIFERA L. (HYMENOPTERA: APIDAE)'” AKEY C. KF HUNG AND HACHIRO SHIMANUKI Bee Research Laboratory, U.S. Department of Agriculture, Beltsville, MD 20705, U.S.A. (e-mail: ahung @asrr.arsusda.gov) Abstract.—The 393bp nucleotide sequences of RNA polymerase genes in three isolates of Kashmir bee virus were compared. There was a 97.4% similarity between Canadian and U.S. isolates and an 80.4% or 81.2% similarity between the Australian strain and the two North American isolates. The amino acid sequence similarity between the two North American isolates was 99.2%. The similarities between the Australian isolate and Cana- dian and U.S. isolates were 97.7% and 96.9%, respectively. These three KBV isolates could also be differentiated by differences in the cleavage sites of the restriction endo- nuclease Maell. Key Words: Apis mellifera, honey bees, Kashmir bee virus, amino acid sequence, nu- cleotide sequence, restriction endonuclease, RT-PCR Kashmir bee virus (KBV) was first iso- lated from a diseased adult bee of Apis cer- ana Fab. (Bailey and Woods 1977). Strains of KBV have been found in adult bees of Apis mellifera L. in Canada, Spain, India, Australia and New Zealand (see Allen and Ball 1995 for further details), Fiji (Ander- son 1990) and the U.S. (Bruce et al. 1995, Hung et al. 1995). According to Anderson (1991), KBV is not a virulent pathogen of A. mellifera. Bailey and Ball (1991) and Al- len and Ball (1995) considered KBV the most virulent virus of all known honey bee viruses. Recently, Hung et al. (1996) re- ported for the first time that a strain of KBV was found in dead honey bees in the U.S. They also reported the first case of simul- ' The nucleotide sequence data reported here have been registered with GenBank under Accession Nos. AF027125, AF034541, and AFO34542. * Mention of any proprietary product in this publi- cation does not constitute an endorsement for use of this product by the USDA. taneous infection of acute paralysis virus (APV) and KBV in the same honey bee col- ony. The reverse transcription-polymerase chain reaction (RT-PCR) was first used for detecting KBV by Stoltz et al. (1995). However, their paper contains no sequence information of KBV genome other than the two ““KBV specific’’ primers they used in PCR. In this paper, we report the results of our nucleotide sequence and restriction en- zyme analyses in three KBV isolates. MATERIALS AND METHODS Freeze-dried samples of pupae injected with a South Australian isolate of KBV were supplied by Dr. Brenda V. Ball. A crude KBV extract of 30 pooled bees from Leamington, Ontario, Canada (identified by Dr. Ball) was supplied by Dr. Cynthia Scott- Dupree. One KBV-infected adult worker bee from the same Maryland bee colony used in Hung and Shimanuki (in press a) VOLUME 102, NUMBER 1 was also included in this study. Freeze- dried pupae were rehydrated and homoge- nized in 0.45 mL of 0.01 M potassium phosphate buffer (pH 6.7). The same amount of buffer was used to homogenize the adult worker bee. After centrifugation at ca. 4,000 g for 15 minutes, direct RT- PCR was carried out with 2 wL of super- natant from either the homogenized sample or the crude extract as described by Hung and Shimanuki (in press b). The primer pair 5'-GATGAACGTCGACCTATTGA-3’ and 5'TGTGGGTTGGCTATGAGTCA-3’ (Stoltz et al. 1995) was used in PCR with each primer at a final concentration of 0.2 pM. The RT-PCR reaction product was puri- fied with the High Pure PCR Product Pu- rification Kit (Boehringer Mannheim). The dye-terminator sequencing of the purified PCR product was carried out on an ABI DNA sequencer (model 373a) at the DNA Sequencing Facility of the University of Maryland Center for Agricultural Biotech- nology. Both DNA strands were sequenced three times to confirm the accuracy of the sequence data. Sequence data were edited and analyzed using DNASIS® for Win- dows® Version 2.5 (Hitachi Software, South San Francisco, CA). DNASIS® was also used in the multiple alignment and the re- striction site search. Restriction endonuclease (RE) digestion was carried out with 3uL of purified PCR products, 5 wL of 2X RE buffer, 1wL of sterile water and 1.5 wL (= 3 units) of the RE, Maell (Boehringer Mannheim), at 50°C for 24h. The digested DNA was analyzed by electrophoresis through a 1.5% (w/v) agarose gel and stained in ethidium bro- mide (0.5 wg/mL). After destaining, DNA bands were visualized with a UV transil- luminator. RESULTS The RT-PCR of both the Australian and Canadian KBV isolates generated a 393 bp product which is the same size as the U.S. KBV reported by Hung and Shimanuki (in 179 press a). Fig. 1 shows the edited sequences of the 393 bp PCR product of both isolates and the sequences of these two segments aligned with that of the U.S. KBV. Multiple alignment of these sequences showed a 97.4% similarity between Canadian and U.S. isolates. The similarities between the Australian isolate and Canadian and U.S. isolates were 80.4% and 81.1%, respective- ly. All three sequences contain a single large open reading frame (deduced amino acid sequence not shown). Although there are 43 miss-matched triplet groups, only four of them with miss-match resulted in amino acid coding differences (Fig. 1, triplet groups in bold face and shaded). These are at positions 123 (ACT for threonine; GCA for alanine), 303 (GCT for alanine; CCA for proline), 312 (TGC for cysteine; GCT for alanine), and 336 (ATG for methionine; GTG for valine). Therefore, the amino acid sequence similarity between the two North American isolates was 99.2%. The similar- ities between the Australian isolate and Ca- nadian and U.S. isolates were 97.7% and 96.9%, respectively. The DNAS1S® database was used in the search for restriction enzymes with one cut- ting site at these 393bp amplicons that will result in a restriction fragment length poly- morphism (RFLP). We found that these three isolates could also be differentiated by the differences in Mael/ restriction endo- nuclease cleavage sites. No cleavage site was found in the Australian isolate. As shown in Fig. 1, the U.S. and Canadian iso- lates each possessed a different Mael/ re- striction site: in the U.S. isolate at triple #83 and in the Canadian at triplet #107. Fig. 2 shows the RFLP of Maell/. The “‘extra”’ low molecular weight fragment (about 100bp) in the U.S. sample was a nonspecific PCR product from the homogenate of the adult worker bee. DISCUSSION The size of the 417 bp product reported for Canadian KBV by Stoltz et al. (1995) 180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON iat 20 2g 38 47 KBVAUO1 --GA CCT ATT GAA AAG GTT AAT CAA TTG AAA ACA CGA GTA TTC TCA AAT KBVECAOM eS — A KBVUSO1 TC S06 5X one eee O00 SOC 56 65 74 83 OZ KBVAUIOIS GGAVCEAVALG IGAT LIC Ler ALTA CCLALLENCCATALG LAL LAT (iG riGG@e asian IGEWIGNOIL 465 coc ooo oo 5JN ood bo © KBVUSO1 50 5G ooo colt oodb 5 oG 506 BOC 101 110 NES) yrs uSy7/ KBVAUO] ATA GCT CAT TTA ATG GAA AAT CGA ATT ACT AAT GAG GTA TCT ATT GGA KEV CRO ® tienen ene MG ers. con G.A ie AG KE VUSOI er. ..G sGW « G.A SAE vee 146 55 164 uIs 182 UI KBVAUOQ1 ACG AAT GTG TAT TCT CAG GAC TGG AGT AAA ACT GTT CGC AAA TTG ACC MEWIC/AON Soo coo oct ao Son 5 Git A.G A KBVUSO1 nig dl sala tl aig lvn Soe 50 ¢ S06 Nal os olN 200 209 218 22a) 236 KBVAUO1 AAA TTT GGA AAT AAA GTT ATT GCG GGT GAT TTT TCA ACT TTT GAT GGA KBVCAO1 A gilt KBVUSO1 50.0 S00 6 Of 50 tt 500 200 245 254 263 272 281 KBVAUO1 TCA CTG AAT GTA TGT ATT ATG GAA AAA TTT GCG GAT TTA GCG AAT GAG KBVCAO1 dN <0 oll no bab C 6U Nec A KBVUSO1 ... ..A .AG@eGPtT Sc en S06) bu G S06 au ene 290 299 308 SN 326 SS KBVAUO1 TTT TAC GAT GAT GGT GET GAG AAT FEC CTG ATT AGG CAT GTG TTA TTG REVCAOI So an orate ACen ee ee Sor T.. . A eee ee KBVUSO1 Sie IRIS OV eee prov a GCT T 5G Gort g/N Wao 344 353 362 Sia 380 KBVAUO1 ATG GAT GTG TAT AAT TCT GTA CAC ATT TGC AAT GAC TCT GTC TAT ATG KBVCAO1l Ae, 2c A at ey Me Gages oo 74s € KBVUSOlL Gey... IC G Tek . Te Cire SAA. thE 389 KBVAUO1L ATG ACT CAT A IMEWC/AOIL ooodecanoo MEWUSOIL concn domc = Fig. 1. Multiple nucleotide sequence alignment of the 393bp RT-PCR product of Australian (KBVAUO1, GenBank AF034541), Canadian (KBVCAO1, GenBank AF034542) and U.S. (KBVUSO1, GenBank AF027125) KBV isolates. Dots represent bases the same as in KBVAUO1. The recognition sequence of the restriction enzyme Maell (ALCGT) is underlined and shaded. Triplets that coded for different amino acids between the samples are in bold face and shaded. was based on their estimation using the mo- lecular weight DNA markers. Although we also used their two primers in PCR, the 393 bp length of the sequenced amplicon re- ported here represents the actual count of bases. Stoltz et al. (1995) did not report the nucleotide sequence of the estimated 417 bp amplicon of their Canadian KBV sam- ple. Our search in the available databases has failed to find any honey bee virus se- quence data deposited by them. Therefore, no comparison can be made between our Canadian sample and theirs. We have shown that the “traditional” RT-PCR protocols used in Hung and Shi- manuki (in press a) and the direct RT-PCR protocol of Hung and Shimanuki (in press b) used in this report produced similar re- sults, both in terms of the size of amplicon and the nucleotide sequence (Fig. 1). The latter protocol can circumvent the time-con- suming virus and RNA purification steps. According to Bruce et al. (1995, Table 1) there are three “serological” strains of KBV in each of the seven states they sur- veyed. Fig. 1 from their paper, however, VOLUME 102, NUMBER 1 1234 Restriction fragment length polymorphism of the 393bp PCR products in three isolates of KBV. DNA products were digested with the restriction en- zyme Maell. Lane 1, 1O0bp DNA ladder; lane 2, Aus- tralian KBV; lane 3, Canadian KBY; lane 4, U.S. KBV. Fig. 2. does not demonstrate that “‘serologically distinct strains of the virus exist in the U.S.” Allen and Ball (1995) divided strains of KBV into two serological groups: the Canadian and Spanish strains formed one group and the Indian, South Australian and New Zealand strains formed another group. As Allen and Ball (1995) further pointed out, “studies on the genomes of both APV and KBV should help to clarify these rela- tionships.’’ Sequence data presented here will serve as reference in further nucleotide sequence analyses of multiple APV and KBV isolates from different localities throughout North America. Restriction enzyme analysis of PCR-am- plified fragments has been used in differ- entiating isolates of cucumber mosaic virus isolates (Rizos et al. 1992) and ovine len- tiviruses (Leroux et al. 1995). As shown in Fig. 2, these three isolates of KBV can be differentiated from each other by the RFLP of Maell. RT-PCR and RFLP analysis of 181 PCR-amplified fragments, therefore, should provide a quick and easy tool to distinguish these three KBV isolates without carrying out sequence analysis. ACKNOWLEDGMENTS We thank Sean Davison, Jay Evans, Neil Leat and Ing-Ming Lee for reviewing the manuscript; Brenda V. Ball and Cynthia Scott-Dupree for providing KBV samples; and Michele Logan for technical assis- tance. LITERATURE CITED Allen, M. FE and B. V. Ball. 1995. Characterisation and serological relationships of strains of Kashmir bee virus. Annals of Applied Biology 126: 471—484. Anderson, D. L. 1990. Pests and pathogens of the hon- eybee (Apis mellifera) in Fiji. Journal of Apicul- tural Research 29: 53-59. . 1991. Kashmir bee virus- a relatively harm- less virus of honey bee colonies. American Bee Journal 131: 767-770. Bailey, L. and B. V. Ball. 1991. Honey bee pathology. Academic Press, London, 193 pp. Bailey, L. and R. D. Woods. 1977. Two more small RNA viruses from honey bees and further obser- vations on sacbrood and acute bee-paralysis virus. Journal of general Virology 37: 175-182. Bruce, W. A., D. L. Anderson, N. W. Calderone, and H. Shimanuki. 1995. A survey for Kashmir bee virus in honey bee colonies in the United States. American Bee Journal 135: 352-354. Hung, A. C. F, J. R. Adams, and H. Shimanuki. 1995. Bee parasitic mite syndrome (II): The role of Var- roa and viruses. American Bee Journal 135: 702— 704. Hung, A. C. F, B. V. Ball, J. R. Adams, H. Shimanuki, and D. A. Knox. 1996. A scientific note on the detection of American strains of acute paralysis virus and Kashmir bee virus in dead bees in one U.S. honey bee (Apis mellifera L.) colony. Api- dologie, 27: 55-56. Hung, A. C. F and H. Shimanuki. In press a. A sci- entific note on the partial nucleotide sequence of a U.S. strain of Kashmir bee virus isolated from Apis mellifera L. Apidologie. . In press b. A scientific note on the detection of Kashmir bee virus in individual honey bees and Varroa mites. Apidologie. Leroux, (€5 4S) Vuillermoz, JF. Morex,and> T- Greenland. 1995. Genomic heterogeneity in the pol region of ovine lentiviruses obtained from bronchoalveolar cells of infected sheep from France. Journal of General Virology 76: 1533-— 1537/5 182 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Rizos, H., L. V. Gunn, R. D. Pares, and R. Gillings. Stoltz, D., X. E. Shen, C. Boggis, and G. Sisson. 1995, 1992. Differentiation of cucumber mosaic virus Molecular diagnosis of Kashmir bee virus infec- isolates using the polymerase chain reaction. Jour- tion. Journal of Apicultural Research 34: 153— nal of General Virology 73: 2099-2103. 160. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 183-187 BIOLOGY AND IMMATURE STAGES OF OCHROTRICHIA QUADRISPINA DENNING AND BLICKLE (TRICHOPTERA: HYDROPTILIDAE), A SPRING- INHABITING SCRAPER J. B. KEIPER AND W. E. WALTON Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e—mail: jkeiper@mail.ucr.edu) Abstract.—Larvae of Ochrotrichia quadrispina Denning and Blickle were obtained from a spring adjacent to the high tide line of a southern California beach. The spring created a madicole over exposed bedrock within which larvae grazed periphyton. Al- though the filamentous chlorophyte Cladophora was present, no larvae consumed this alga during observations of all instars in the laboratory. Gut contents analysis showed that unicellular algae, detritus, and mineral materials were consumed in the field. Final instars constructed cases of a single convex dome with a flexible silken ventral valve that sealed the case when larvae were withdrawn. Morphological descriptions of the five instars are given. Key Words: Microcaddisflies (Trichoptera: Hydroptil- idae) are known from lentic and lotic fresh- water habitats (Wiggins 1996a). Larvae of the genus Ochrotrichia Mosely appear to be associated with flowing water (Ross 1944). Most species of this genus feed by scraping periphyton, although some species, such as O. xena Ross, consume the liquid contents of cells within filaments of green algae (Chlorophyta) (Wiggins 1996b, Keiper and Foote 1998). Vaillant (1984) described the morphology and gave biological observa- tions of the mature larvae of O. confusa found in a madicolous (i.e., dripping rock) habitat; he mentioned that O. quadrispina inhabits these shallow waters as well. Al- though the larva of the latter species was not described, Vaillant (1984) mentioned that slight morphological differences be- tween the two exist. Herein, we describe the early and mature larvae of O. quadrispina and give biological observations acquired during laboratory rearings. Ochrotrichia, microcaddisflies, aquatic insects, larvae, madicoles MATERIALS AND METHODS Larvae of O. quadrispina were collected in June from a spring flowing over a bed- rock outcropping at the ocean-side base of sand dunes approximately 500 m south of South Carlsbad Beach (CA, San Diego Co.). This freshwater habitat was situated 1—2 m vertically from the debris wrack cre- ated by the Pacific Ocean high tides. The water emanated from a seepage area within a dense stand of cattails (Typha sp.), was less than 0.5 cm deep, and formed a few pools of water in shallow concavities. Examination of the substrate revealed di- atoms, unidentified unicellular green algae (Chlorophyta), long filaments of Cladopho- ra sp. (Chlorophyta), short cyanobacterial filaments, and miscellaneous detrital and mineral particles. Odontomyia sp. (Diptera: Stratiomyidae) larvae were abundant throughout the spring, and Hydropsyche (Trichoptera: Hydropsychidae) larvae and Hydrophilidae (Coleoptera) larvae and 184 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON adults were occasionally encountered. Lar- val and pupal O. quadrispina were placed in jars of field-collected spring water, and put in a cooler along with Cladophora and pieces of diatom-covered bedrock for trans- port back to the laboratory. Adult speci- mens were taken from dead Typha stems hanging over the bedrock. Immature stages were separated into pu- pae, fifth instars, and early instars (1-4), and each group was placed separately into small (2.5 cm diameter) Petri dishes, along with water and field-collected materials. Early instars were determined based on qualitative appearance recognizable from experience working with Hydroptilidae, and confirmed using Dyar’s Rule (Dyar 1890, Richards and Davies 1977). Dishes were placed near a window to maintain a natural photoperiod, and the temperature in the lab- oratory was maintained at ~20° C. Water was changed every 3—4 days. Some specimens were killed in near-boil- ing water, fixed with Kahle’s fluid, and pre- served in 70% ethanol following the meth- ods of Wiggins (1996a). Gut contents ana- lyses of field-collected fifth instars (n = 6) were performed by dissecting the guts from fixed specimens and mounting on slides. Larval behavior was observed with a Wild MZ8 dissecting microscope, with the light source set on the lowest intensity needed for observation to reduce the possibility of disturbance. Morphological description of larvae fol- lowed the methods described by Keiper and Foote (1999). Only field-collected larvae were used for description, as efforts to ob- tain viable eggs from field-collected fe- males were unsuccessful, and few early in- stars were collected in nature. Therefore, only one larva for each early instar was de- scribed; measurements and descriptions of fifth instars are based on six larvae. RESULTS Final (5th) instar—Length 2.39—2.63 (x = 2.48 + 0.13) mm. Head: width 0.21—0.22 (x = 0.21 + 0.01) mm; uniformly dark brown to black, without muscle scar pat- tern, with pale ring around black eye spot (Fig. 1); position of primary setae as in Fig. 2; long seta near eye approximately 1.5 length of head; anterior edge of labrum slightly convex in dorsal view; antenna short, with broad base, bearing long seta 5X length of antenna, flagellum 3 segmented; each mandible bearing one seta on postero- lateral corners (Fig. 3), asymmetrical, with right pointed and left bearing coarse teeth and fine setae. Thorax: uniformly dark brown; anterolateral lobes not well devel- oped; two prosternal sclerites, posterior margin of each with a short pointed lobe. Prothoracic leg as in Fig. 4; tarsal claw somewhat curved, basal seta approximately half the length of claw; tarsus with one dor- sal seta approximately the length of claw; venter of tibia with strong projection bear- ing four stout setae (only three visible in lateral view); femur with two strong setae ventrally and one dorsally; minute second- ary setae present on all segments. Abdo- men: milky white and translucent, primary setae as in Fig. 1; dorsomedial sclerite of first segment roughly rectangular, with 3 well-developed setae laterally (Fig. 5); dor- somedial ring-like sclerites on segments 2— 8 dark brown (Fig. 5); sclerite sa2 elliptical, bearing one strong seta; sclerite sa3 roughly circular, small, bearing one small seta; dor- sal sclerite of segment 9 rectangular, with anterior margin convex, bearing 3 strong posterior setae, middle pair approximately 2—3 times length of others; sclerites of ab- dominal segment 10 and prolegs typical of Ochrotrichia (Wiggins 1996a). Case.—Tortoise-shell type (Wiggins 1996a), constructed from tiny mineral par- ticles; dorsal peak of upper valve skewed slightly in transverse cross section; lower valve lacking particles; resembles dome case illustrated by Wiggins (1996a). Early instars flattened dorsoventrally, with uniformly brown sclerites on thoracic and abdominal segments, darkening with age; non-sclerotized areas milky white; se- tae well developed. Anal prolegs with long VOLUME 102, NUMBER 1 Figs. 1-6. Mandibles of fifth instar, ventral view. 4, Right foreleg of fifth instar. 5, Dorsomedial sclerite, first abdominal segment (top), and dorsomedial ring sclerite of second abdominal segment (bottom), fifth instar. 6, Third instar, lateral view. apical claw. Differences among early in- stars mainly a matter of proportion. Instars 1—4 free-living, never constructing cases. First instar—Length 0.42 mm. Head: width 0.05 mm; uniformly light brown, translucent, no evidence of ring around black eye spot. Thorax same color as head capsule. Abdomen sclerites same color as head capsule. Second instar—Length 0.55 mm. Head: width 0.09 mm; uniformly light brown, slightly darker than first instar, translucent, no evidence of ring around Ochrotrichia quadrispina. |, Fifth instar, lateral view. 2, Head of fifth instar, dorsal view. 3, black eye spot. Third instar—Length 0.72 mm. Head: width 0.12 mm; uniformly light brown, slightly darker than second instar, with inconspicuous pale ring around black eye spot (Fig. 6). Fourth instar—Length 1.04 mm. Head: width 0.19 mm; uniformly brown (somewhat lighter than fifth instar), with definite pale ring around black eye spot. Early instars were found almost exclu- sively in the masses of Cladophora col- lected from the madicole. The algal micro- 186 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON habitat probably represents refugia and a foraging area for the small larvae. During observations of feeding in the laboratory, larvae crawled across the rock substrate, and rapidly abducted and adducted their mandibles in an apparent effort to graze the periphyton present; their guts became dark brown. Some larvae, particularly later in- stars (3—4), moved among filaments of Cla- dophora and grazed the epiphytes growing on the cell walls. At 50X, we observed lar- vae scraping diatoms and unicellular chlo- rophytes scattered on the cell walls with the rapid action of the mandibles. The fifth instars fed in a manner similar to that of the early instars. Larvae grazed the substrates with their mandibles held perpendicular to, or nearly parallel with, the cell walls or rocks. When perpendicular to the substrates, the distal tips of the mandi- bles macerated diatomaceous substrates or exploited small concavities. Larvae often shifted the position of their mandibles, and placed them nearly parallel to the substrate so the inner margins scraped the periphy- ton. The setation on the inner margins prob- ably aided in brushing the scraped materials into the mouth (cf., Cummins and Merritt 1996). Fifth instars were observed only on rock substrates in the field, and never among the filaments of algae. Gut contents analyses showed that diatoms, unicellular green algae, and miscellaneous detrital and mineral particles were consumed. Newly-molted fifth instars initiated case construction by fashioning small rings of material around their abdomens. After ap- proximately 24 hours, the cases resembled completed ones, except that they were roughly the same length as the larva; cases were completed in 3—4 days. Upon with- drawing into the case, mechanical tension caused the silk sheet of the lower valve to become contiguous with the upper valve occluding the opening; the flexibility of the lower sheet allowed larvae to extrude their head and thoracic segments for feeding or movement. Larvae that fully withdrew into their cases appeared to have a silken guy- line (cf., Nielsen 1948, Wiggins 1996a) at- tached to the substrate to help maintain their position in the current. In the field, pupal cases were often scat- tered individually on the bedrock surface. However, many aggregations were formed by up to 15 individuals situated side-by- side or end-to-end. Examination of these cases, as well as those individuals which pupated in the laboratory, showed that only the upstream end of the case was attached to the substrate; cases were therefore posi- tioned at a shallow angle relative to the sub- strate when viewed laterally, and were ori- ented parallel to the flow of the madicole. The duration of the pupal stage was 9-10 days under laboratory conditions (n = 3). DISCUSSION To our knowledge, there have been no published descriptions of the early instars of Ochrotrichia spp. The early instars of O. quadrispina have dark sclerites, whereas larval O. wojcickyi Blickle are colorless during the first two stadia, and only the tho- racic sclerites and head capsule darkened slightly during the third and fourth stadia (Keiper 1998). More information on early instars is needed before further morpholog- ical comparisons between species can be made. Ross (1944) provided brief descriptions of and a key to seven Ochrotrichia spp. fi- nal instars; O. quadrispina keys to O. riesi based on the presence of dark dorsomedial sclerites with a transverse membranous cen- ter on abdominal segments 2—8, as does O. confusa (Vaillant 1984). These three species can be separated based on the shape of the dorsomedial sclerites; the anterior edge is produced forming a triangular point in O. confusa contrasting the pointed posterior edge of O. quadrispina, and the anterior and posterior edges are truncated on O. rie- si. The dorsomedial sclerites of other spe- cies form a small, membranous ring (Ross 1944, Flint and Herrmann 1976, English and Hamilton 1986, Keiper, in press). Man- dible morphology also appears to be a use- VOLUME 102, NUMBER 1 ful character for separating species because dentition, presence or absence of bristles on the inner edges, and number of setae on the posterolateral corners varies among those species for which this information is avail- able (Vaillant 1984, English and Hamilton 1986, Keiper and Foote 1998, Keiper 1998, in press). Morphology of dorsomedial ab- dominal sclerites and mandibles appear dis- tinct among species suggesting that the eventual compilation of larval keys to spe- cies is possible, and preliminary efforts in- dicate that other characters such as protho- racic sclerite shape and sclerite coloration may prove useful. Gut contents analysis showed that a wide variety of periphytic inaterials were con- sumed by larvae. Although species of the hydroptilid genera Dibusa Ross (Resh and Houp 1986), Hydroptila Dalman, and Ox- yethira Eaton (Keiper et al. 1998) appear quite specialized in their food preferences, O. quadrispina is a generalist consumer of periphyton. ACKNOWLEDGMENTS We thank R. T. Pachon and P. D. Work- man (University of California-Riverside) for critically reviewing the manuscript. O. S. Flint (USNM) and an anonymous re- viewer offered helpful suggestions to im- prove this work. LITERATURE CITED Cummins, K. W. and R. W. Merritt. 1996. Ecology and distribution of aquatic insects, pp. 74—86. /n Mer- ritt, R. W. and K. W. Cummins, eds., Introduction to the Aquatic Insects of North America. Kendall/ Hunt Publishing Company, Dubuque. Dyar, H. G. 1890. The number of molts of lepidopter- ous larvae. Psyche 5: 420—422. English, W. R. and S. W. Hamilton. 1986. The larva of Ochrotrichia arizonica (Trichoptera: Hydrop- tilidae) with notes on distribution and geographic variation. Journal of the Kansas Entomological Society 59: 474-479. 187 Flint, O. S. and S. J. Herrmann. 1976. The description of, and environmental characterization for, a new species of Ochrotrichia from Colorado (Trichop- tera: Hydroptilidae). Annals of the Entomological Society of America 69: 894-898. Keiper, J. B. 1998. Biology, larval feeding habits, and resource partitioning by microcaddisflies (Tri- choptera: Hydroptilidae). Ph.D. Thesis, Kent State University. 146 pp. . In press. Morphology of final instar Ochro- trichia xena (Trichoptera: Hydroptilidae). Ento- mological News. Keiper, J. B., D. A. Casamatta, and B. A. Foote. 1998. Use of algal monocultures by larvae of Hydroptila waubesiana and Oxyethira pallida (Trichoptera: Hydroptilidae). Hydrobiologia 380: 87-91. Keiper, J. B. and B. A. Foote. 1998. Notes on the bi- ology of Ochrotrichia xena Ross (Trichoptera: Hydroptilidae), a species newly recorded for Ohio. Proceedings of the Entomological Society of Washington 100: 594-595. . 1999. Biology and immature stages of two species of Hydroptila Dalman (Trichoptera: Hy- droptilidae) which consume Cladophora (Chlo- rophyta). Proceedings of the Entomological So- ciety of Washington 101: 514-521. Nielsen, A. 1948. Postembryonic development and bi- ology of the Hydroptilidae. Det Kongelige Danske videnskabernes selskabs Biologiske Skrifter 5: 1— 200. Resh, V. H. and R. E. Houp. 1986. Life history of the microcaddisfly Dibusa angata and its association with the red alga Lemanea australis. Journal of the North American Benthological Society 5: 28— 40. Richards, O. W. and R. G. Davies. 1977. Imms’ gen- eral textbook of entomology: Vol. 1, structure, physiology, and development, 12th ed. Chapman and Hall Publishers, London. Ross, H. H. 1944. The caddis flies, or Trichoptera, of Illinois. Bulletin of the Hlinois Natural History Survey 23: 1-326. Vaillant, EF 1984. The hydroptilid larvae of dripping rocks, pp. 407—412. In Morse, J. C., ed., Proceed- ings of the 4th International Symposium on Tri- choptera. Series Entomologica 30, 486 pp. Wiggins, G. B. 1996a. Larvae of the North American caddisfly genera (Trichoptera), 2nd ed. University of Toronto Press, Toronto. . 1996b. Trichoptera families, pp. 309-349. In Merritt, R. W. and K. W. Cummins, eds., Intro- duction to the Aquatic Insects of North America. Kendall/Hunt Publishing Company, Dubuque. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 188-197 MACROLEPIDOPTERAN LARVAE SAMPLED BY TREE BANDS IN TEMPERATE MESIC AND XERIC FORESTS IN EASTERN UNITED STATES LINDA BUTLER AND JOHN STRAZANAC Division of Plant and Soil Sciences, PO. Box 6108, West Virginia University, Mor- gantown, WV 26506-6108, U.S.A Abstract.—We studied macrolepidopteran larval richness and abundance under canvas tree bands in 18 oak-dominated plots in the George Washington National Forest in Vir- ginia and the Monongahela National Forest in West Virginia. In each plot, 12 canvas- banded trees were monitored from early May through mid-August in 1995 and 1996, totalling 216 trees. The bands trapped a total of 6,347 larvae representing 77 species in 8 macrolepidopteran families. Seventy-five percent of all larvae (4,773) were Lymantria dispar (L.) (Lymantriidae). Of the remaining 1,574 larvae, 62% (981) were noctuids; 22% (348) arctiids, and 8% (129) lasiocampids. The most abundant species after Lymantria dispar, were Morrisonia latex (Guenée) (Noctuidae) (5%), Hypoprepia fucosa Hiibner (Arctiidae) (4%), Phoberia atomaris Hiibner (Noctuidae) (2%), and Abagrotis alternata (Grote) (Noctuidae) (2%). Differences in species distribution and abundance occurred between forests and between years. Key Words: moth, diversity Relatively few studies have been con- ducted on communities of forest macrole- pidopteran larvae in North America. These studies include Lepidoptera associated with an outbreak species (Stevens et al. 1984), seasonal diversity and abundance on spe- cific host species (Marquis and Passoa 1989; Butler 1992; Wagner et al. 1995), and larval species and abundance in areas being treated with insecticides for gypsy moth suppression (Miller 1990; Butler and Kon- do 1993; Butler et al. 1995a, b; Wagner et al 1995291996): In most of these studies, macrolepidop- teran larvae were sampled only from fo- liage. However, studies by Butler and Kon- do (1993), Butler et al. (1995a), Butler et al. (1997), and Wagner et al. (1995, 1996) sampled them from under burlap or canvas bands, thus establishing tree banding as a Macrolepidopteran larvae, canvas tree bands, forest caterpillars, gypsy viable method for assessing lepidopteran larval diversity or nontarget spray impact. In 1994, we began a study of impact of the biological spray Bacillus thuringiensis kurstaki Berliner and defoliation by gypsy moth, Lymantria dispar L., on nontarget arthropods in large plots in two adjacent national forests in the central Appala- chians. Pretreatment sampling of macro- lepidopteran larvae from foliage and from canvas bands and adult macrolepidopter- ans by light trap was conducted in 1995 and 1996. Here we present the seasonal di- versity of species in these forests as as- sessed by canvas tree bands. Other papers will cover results of the other sampling methods and a final paper will present spe- cies diversity indices and species accu- mulation curves for the lepidopteran com- munity. VOLUME 102, NUMBER 1 189 Rigel. in Virginia (VA) and plots 10—18 in the Monongahela National Forest (MNF) in West Virginia (WV). MATERIALS AND METHODS In 1994 18, 200-ha study plots were des- ignated and flagged in gypsy moth suscep- tible oak-dominated forests. Plots 1 through 9 are located primarily on the eastern slope of Great North Mountain in the Deerfield Ranger District of the George Washington National Forest (GWNP), Augusta County, Virginia (Fig. 1). The GWNF plots are in a xeric forest of mixed oak and pine with a range in elevation of 586 through 791 m in the rain shadow of the Appalachians. Plots 10 through 18 are in the Monongahela Na- tional Forest (MNF), Pocahontas County, West Virginia (Fig. 1). Plots 10 through 12 are in the southern Greenbrier Ranger Dis- trict on Chestnut Ridge (Paddy Knob North); plots 13 through 18 are in the Mar- linton Ranger District, with plots 13 through 15 near Sugar Camp Run (Paddy Knob South) and plots 16 through 18 on Marlin Mountain. The MNF plots are in a more mesic forest with a range in elevation of 860 through 1,070 m. In June 1995, rain gauges and maximum- Study area showing arrangements of plots 1—9 in the George Washington National Forest (GWNF) minimum thermometers were installed in all 18 plots. Additional weather data was obtained from N.O.A.A. weather stations. Thirty-cm wide, 18 oz. OD green canvas bands were stapled around the circumfer- ence of trees. The top edges of the bands were 1.4 m above the ground. Twelve dom- inant or codominant trees were banded in each of the 18 plots making a total of 216 trees. Generally in each plot, 6 banded trees were at a higher elevation near a ridge top and 6 were at a lower elevation within the watershed. Of the 12 trees in each plot, 10 were various oak species (Quercus spp.), one a red maple (Acer rubrum (L.) and one a hickory (Carya spp.). Among the 180 banded oak trees, 69 were in the red oak group (Quercus coccinea Muenchh, Q. rub- ra L., and Q. velutina Lambert), 57 were chestnut oak (Q. prinus L.), and 54 were white oak (Q. alba L.). Caterpillars were removed from under bands weekly from early May through mid- August (9 May through 15 August; 7 May through 13 August, for 1995 and 1996, re- 190 spectively). Larvae were placed in labelled, plastic 8-dram vials with snap lids and stored in a cooler in the field until they were returned to the laboratory in Morgantown, WV the following day. In the laboratory, vials were stored in a refrigerator at 4° C until larvae were identified the following day. Voucher specimens from the study are deposited in the WVU Arthropod Collec- tion. Species are listed in Table 1 according to the checklist given by Hodges et al. (1983), with modifications from Poole and Gentili (1996). RESULTS AND DISCUSSION During the sampling seasons of 1995 and 1996, a total of 77 species of macrolepi- dopteran larvae were collected from canvas bands, 59 species from the GWNEF and 52 species from the MNF (Table 1). These lar- vae represented 8 families, with most spe- cies being Noctuidae (40), followed by Geometridae (16), Arctiidae (9), and Ly- mantriidae (4). The remaining 4 families were represented by one to three species. Lymantriid larvae were the most abundant, with 4,792 individuals; 4,773 being gypsy moth, Lymantria dispar. Noctuid total abundance was 981 followed by Arctiidae (348), Lasiocampidae (129), Geometridae (52), and Saturniidae (36) (Table 1). The more abundant species following the gypsy moth were Morrisonia latex (Gue- née) (315), Hypoprepia fucosa Htibner (270), Phoberia atomaris Hiibner (135), Abagrotis alternata (Grote) (117), Mala- cosoma americana (FE) (107), Catocala amica (Hiibner) (79), Orthosia rubescens (Walker) (71), Catocala ilia (Cramer) (55), Acronicta modica (Walker) (51), Epiglaea decliva (Grote) (41), Hemileuca maia (Dru- ry) (35), Malacosoma disstria Hiibner (21), and Lambdina fervidaria (Hiibner) (20) (Table 1). All of these most abundant spe- cies were taken from both forests with the exception of H. maia which was collected only from the GWNE This species is col- lected more typically in dry oak-pine for- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ests such as the GWNF than in more mesic mixed oak forests such as the MNF (Fer- guson 1971; Butler et al. 1995b). Of other more abundant species, several were col- lected at a much higher percentage in the GWNE: Morrisonia latex (79%), Phoberia atomaris (82%), Catocala amica (72%), Orthosia rubescens (79%), and Epiglaea decliva (98%). Malacosoma americana was in similar abundance in both forests. The preferred host of M. americana in our study area is black cherry (Prunus serotina Ehrh.); the larvae do not feed on any of our banded tree species. In the last instar the larvae tend to wander in search of a pupa- tion site (Franclemont 1973). Several species were collected exclusive- ly from a single forest. Listed in the order in which they appear in Table 1, these in- clude: Besma quercivoraria (Guenée), Na- data gibbosa (J.E. Smith), Zanclognatha li- turalis (Hiibner), Acronicta lobeliae (Gue- née), Acronicta haesitata (Grote), and Hi- mella intractata (Morrison) from the MNE and P. titea (Cramer), Phigalia denticulata Hulst, P. strigataria (Minot), Erannis tili- aria (Harris), Campaea perlata (Guenée), Dryocampa rubicunda (F.), Meganola spo- dia Franclemont, Lithophane querquera Grote, L. antennata (Walker), Psaphida re- sumens Walker and Achatia distincta Htib- ner from the GWNE However, we collected only one individual of each of these species (Table 1). An overall decline in larval abundance occurred from 1995 (5,129 larvae) to 1996 (1,218 larvae). Lymantria dispar had the greatest influence on that decline as its numbers under bands fell from 4,114 in 1995 to 659 in 1996. Exclusive of gypsy moth, larval numbers were 1,015 and 559 for 1995 and 1996, respectively (Table 1). Among the 7 more abundant larval spe- cies (exclusive of gypsy moth), 5 declined from 1995 to 1996. Hypoprepia fucosa fell from 189 to 81; Catocala ilia, 43 to 12; Catocala amica, 69 to 10; Orthosia rubes- cens, 57 to 14; and Abagrotis alternata, 83 to 34. During that period, numbers of Phob- VOLUME 102, NUMBER 1 eria atomaris larvae remained stable (60 in 1995 and 75 in 1996), while numbers of Morrisonia latex increased from 94 in 1995 to) 221 in 1996; We suggest that the decline in abundance of gypsy moth larvae from 1995 to 1996 was due to movement into the study areas of the gypsy moth fungus, Entomophaga maimaiga Humber, Shimazu, and Soper. Declines of other species may have been influenced by weather, changes in numbers of their natural enemies, or both. The plots were drier and warmer in 1995 than in 1996 (Table 2). Numerous studies have shown di- rect and indirect effects of weather trends and atypical catastrophic temperature and rain events on insect populations (Martinat 1987). The increase in abundance of Mor- risonia latex may likely represent a cyclical buildup because numbers of this species have continued to climb in subsequent years of this study (unpublished data). Tree bands mimic naturally occurring bark flaps and provide a refuge for cater- pillars that move up and down trunks (But- ler et al. 1995a). Thus, larvae under bands may be feeding on forest floor plants, li- chens, or other organic matter on tree trunks (Wagner et al. 1995), or band col- lections may represent actual hosts. The highest number of larval species was col- lected from the red oak group (50), fol- lowed by white oak (39), chestnut oak (38), and hickory and maple (23 each). We note that tree species groups were not sampled equally; of the 216 total banded trees, 32% were red oak; 26%, chestnut oak; 25%, white oak; and maple and hickory, 8% each. The highest larval abundance was taken from red oak (2,438), followed by chestnut oak (2,250), white oak (885), hick- ory (484), and maple (290). Eighty-eight percent of the gypsy moth larvae were tak- en from Quercus spp., its preferred host; Quercus spp. were 83% of the banded trees. Among the banded oaks, 44% of the larvae were on red oak; 41%, on chestnut oak; and 15%, on white oak. Other species for which oaks are a preferred host include Phoberia 19 atomaris, Catocala amica, and C. ilia (Forbes 1954). Morrisonia latex is consid- ered to be a general feeder (Wood and But- ler 1989), and it was collected from all host species in the current study; 85% were col- lected from the oak species. A similar pat- tern was seen for Abagrotis alternata, a general feeder on many tree species (Butler et al. 1997, Forbes 1954); it frequently feeds on blueberry (Vaccinium spp.) and other low woody plants, and climbs trees to shelter under bark flaps. Hypoprepia fucosa feeds on algae, lichens, and other organic material on tree trunks (Forbes 1960). While we might expect to find it randomly distributed among the tree species in our study, 87% of the larvae were on trunks of oaks; 74% of the larvae on oaks were on red and chestnut oaks, the species with the roughest bark that harbors notable quanti- ties of lichens in our study plots. Those spe- cies groups represented 70% of the banded oaks. During our 2 years of sampling, various species of larvae were taken from canvas bands throughout the season, from the ear- liest date (8 May) through the latest (14 Au- gust) (Table 1). Among the more abundant species were the spring defoliators such as Malacosoma spp., Lymantria dispar, Phob- eria atomaris, Catocala spp., Epiglaea de- cliva, Orthosia rubescens, and Abagrotis alternata, that generally were collected from one of the earliest sample dates into June or July. The abundant lichen feeder Hypoprepia fucosa was sampled from 8 May through 22 July. The most abundant larva taken from bands during the summer was Morrisonia latex, collected from 10 July through 12 August (Table 1). While collection of 77 species of macro- lepidopteran larvae produces some useful information on species richness in the plots, information on seasonal diversity and forest differences can be evaluated for only a few of these species. Fifty-three species were represented by 5 or fewer specimens (Table 1). Only species that are in relatively high numbers and more typically shelter under 192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Species and abundance of macrolepidopteran larvae collected from canvas bands of five tree groups in the George Washington (G) and Monongahela National Forest (M) in eastern United States during 1995 and 1996. Maple Hickory Red Oak Family —————— ee eee Species G M G M G M Lycaenidae Satyrium falacer (Godart) O 0) 0) 0) 3 1 Geometridae Alsophila pometaria (Harris) 10) O 0) O O O Tridopsis larvaria (Guenée) 0) 0) 0) O O 1 Melanolophia canadaria (Guenée) 0) 0) 0) 0) 1 l Hypagyrtis unipunctata (Haworth) 1 0) 0) 0) 0) 0) Phigalia titea (Cramer) 0) 0) 0) 0) 0) 0) Phigalia denticulata Hulst 0) 0 0) 0) 1 0) Phigalia strigataria (Minot) 0) 0) 0) 0) 1 0) Erannis tiliaria (Harris) 0) 0) 0) 0) 1 0) Lomographa vestaliata (Guenée) 0) 0 0 O 0) 8 Campaea perlata (Guenée) 0) 0) 0) 10) 0 0) Probole amicaria (Herrich-Schiaffer) 0) 0) 0) O 1 10) Besma quercivoraria (Guenée) 0 0) 0) 0) 0) 0 Lambdina fiscellaria (Guenée) 0 0) 0) O 0 0) Lambdina fervidaria (Htibner) 0) 0 0) 0) =) 2 Hydrelia condensata (Walker) 0 0) 0 0) 0) ] Eupithecia herefordaria Cassino & Swett 0) 0) 0) 0) 0) 0) Lasiocampidae Tolype velleda (Stoll) 0) 0) 0) 0) 0) 0) Malacosoma disstria Hiibner 0) 0) 0 5 y 9 Malacosoma americana (F.) | 0) 5 36 28 Saturniidae Dryocampa rubicunda (F.) | O 0) 0) 0) 0) Hemileuca maia (Drury) 0 0 l 0 6 0) Notodontidae Nadata gibbosa (J.E. Smith) 0) 0) 0) 0) 0) 0) Heterocampa guttivitta (Walker) 0) O 0 0) O O Arctiidae Crambidia pallida Packard 0) 0) 0) 0) 1 0) Hypoprepia miniata (Kirby) 3 l 7 1 8 10 Hypoprepia fucosa Hiibner 6 13 8 9 39 41 Clemensia albata Packard O 0) 0) 0) 1 1 Holomelina opella (Grote) 0) 0) 0) 0) 2 0) Holomelina nigricans (Reakirt) 0) 0) 0) 0) 3) 0) Hyphantria cunea (Drury) 0) O 0) 0) 0) O Ecpantheria scribonia (Stoll) 0) 0) 0) 0) O Halysidota tessellaris (J.E. Smith) 0) 0) 1 0) 0) O Lymantriidae Dasychira dorsipennata 0) 1 O O 1 4 (Barnes & McDunnough) 0) 0) O 0) 0) 0) Dasychira basiflava (Packard) O 10) | 0 1 O Dasychira obliquata (Grote & Robinson) 0) 0) l 0) 0) 1 Lymantria dispar (L.) 134 80 279 2 1,185 679 Noctuidae Idia aemula Hiibner 2, 1 1 0) O 2 Idia lubricalis (Geyer) O 0) 0) 0) 1 0) VOLUME 102, NUMBER 1 193 Table 1. Extended. eR SSS SS ES Chestnut White Years Oak Oak G M Combined Neen ee SS ee, jf Grand Seasonal G M G M 1995 1996 1995 1996 G M Total Range eeeeeeeeeeeeeEeEeEeEeEeEeEeEeEyEy————————EEEEeEeEE—E——E——EEeE>E>y~»~L__EEEE 0) 2 0) 0) 3 0) 3 0) 3 3 6 22 May-—12 June O 0) 1 1 | 0) 1 0) ] 1 2 29 May-12 June 0) 0) 0) 0) 0) 0) 0) l 0) 1 1 1 July—1 July | 3 0) ] 2 0) 4 1 2, 5 7 29 May-3 June 0) 0) 0) 1 1 0) 0) l l l 2 15 May—24 June 1 0) 0) 0) 1 0) 0) 0) 1 0) 1 22 May—22 May 0) 0) 0) 0) ] 0) 0) 0) 0) 1 22 May—22 May 0) 0) 0) 0) l 0) 0) 0) 1 0) 1 22 May—22 May 0) 0) 0) 0) | 0) 0 0 1 0 1 29 May—29 May 0) 0) 0) ] 0) 0 9 0) 0) 9 9 7 Aug.—14 Aug. l 0) 0) 0) 1 O 0) 0) l 0) ] 17 July—17 July 0) 0) 0 0) 0 | 0 0) | 0) 5 Aug.—5 Aug. 0) 0) 0) 1 0) 0 1 0) 0) l l 7 Aug.—7 Aug. D 0) 0) 0) 2 0) 0) 0) 2 0) 2 19 June—31 July 4 1 2, 6 4 7 3 6 11 9 20 19 June—12 Aug. 0) 0) 0) 0) 0) 0) 0) l 0 l 1 29 July—29 July 0) 0) 0) | 0) ) 0) l 0 | 1 15 July—15 July 1 0) 0) O l 0 0) 0) l 0) l 17 July—17 July 1 2 1 1 3 1 Ivy) 0) 4 LF 21 29 May-—10 June 0) 16 12 8 49 0) 58 0) 49 58 107. 15 May—19 June 0 10) 0 0) | 0) 0) O l 0) | 7 Aug.—7 Aug. 20 0) 8 0) 30 5 0 0) 35 0) 35. 29 May-29 July 0) 0) 0) ] 0) 10) 1 0) 0) ] l 7 Aug.—7 Aug. 2 0) 0) 0) 2 0) 0 0) 2 0 2 7 Aug.—7 Aug. ] 0) 1 0 2 1 O 0) 5} 0 } 3 July—5 Aug. 10 5 5 7 21 12 23 1 33 24 57 22 May-17 June 62 Si 23 38 84 54 105 27 138 132 270 8 May—22 July 0) 0) 0) 1 1 0) 0 2 1 Z: 3. 24 July—5 Aug. 0) 0) 0) 0) 2 0 0) 0) y) 0) 2 15 May—22 May 1 0) 1 1 l 4 | 0 5 | 6 22 May-17 June 0) 0) 0) 1 0) 0) ! 0) 0 | l 7 Aug.—7 Aug. 0) 10) 0) 0) 0) l 0) 0) 1 0 l 3 June—3 June 0) 1 2 l 1 2 0) 2 3 2 5 14 Aug.—12 Aug. 0 0) 0) 1 1 0 6 0) ] 6 7 29 May-3 July 0) 0) 0) 0) 0) 0) 0) 0) 0) 0) 0) 2 0) 0) 0) 3 | 0) 0) 4 0) 4 8 May-—5 Aug. 2 7) 0) 2 3 0) 3 2 3 5 8 22 May-10 June 1077 646 383 238 2S 9283 1,339 376 3,058, Auliloy 445773 8 May—22 July 2 2 2 0) 6 1 4 1 7 5 12. 15 May-1 July 10) O 10) 0) l 0) 0) 10) 1 0) | 5 June—5 June 194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Maple Hickory Red Oak Family Species G M G M G M Zanclognatha lituralis (Hubner) 0) 0) 0) 0) 0) 0) Zanclognatha laevigata (Grote) 0) 0) 0) 0) 1 0) Zanclognatha jacchusalis (Walker) 0) 0) 0) 0) 1 2 Renia sobrialis (Walker) 0) 0) 0) 0) O 2) Bomolocha baltimoralis (Guenée) 1 0) 0) 0) 0) 0) Phoberia atomaris Hiibner l 0) DD l 47 8 Zale lunata (Drury) 0) 0) 0) 0) 0) 0) Zale undularis (Drury) 0) 1 0) 0) O 0) Zale minerea (Guenée) 0) 1 0) 0 1 0) Parallelia bistriaris Htibner Dy 0) 0) 0) 0) 0) Catocala epione (Drury) O 0) 19 1 0) 0) Catocala flebilis Grote 0) 0) 3 2 0) 0) Catocala ilia (Cramer) 0) 0) 0) 0) 15 3 Catocala similis W. H. Edwards 0) 0) 0) O 5 0) Catocala micronympha Guenée O 0) | O I 0) Catocala amica (Hitibner) 3 4 O 26 0) Meganola spodia Franclemont 0) 0) 0) 0) 1 0) Nola triquetrana (Fitch) 0) 0) 0) 0) 0) 0) Acronicta lobeliae Guenée 0) 0) 0) 0) O 1 Acronicta ovata Grote 0) 0) 0) 0) 0) 1 Acronicta modica Walker ] 0) 5 0) 12 0) Acronicta haesitata (Grote) 0) 0) 0) O 0) 0) Acronicta afflicta Grote O 0) 0) 0) 1 0) Lithophane innominata (J.B. Smith) 4 0) 0) O ] 72 Lithophane querquera Grote 0) O l 0) 0) 0) Lithophane antennata (Walker) 0) O 0) 0) 1 0) Epiglaea decliva (Grote) l O 5 O 16 O Sunira bicolorago (Guenée) 0) 0) 0) O 0) l Psaphida resumens Walker l 0) 0) 0) 0) O Polia nimbosa (Guenée) 0) | 0) 0) 0) 3 Morrisonia latex (Guenée) 13 4 21 8 92 19 Orthosia rubescens (Walker) 6 0) 12 i 22 11 Orthosia hibisci (Guenée) 0) O | O 0) 1 Himella intractata (Morrison) 0) 0) 0) 0) 0) 1 Achatia distincta Hiibner 0) 0) 1 0) 0) 0) Spaelotis clandestina (Harris) 1 ] 0 0) 0) 0) Abagrotis alternata (Grote) 1 1 4 1 35) 16 Acronicta sp. 0) 0 0) 0) 0) 0) Totals by tree species/forest/years 183 107 378 106 1,578 860 Grand Total by Tree Species 290 484 2,438 bands lend themselves to diversity assess- ment by canvas bands. In our study, these species included in order of declining abun- dance, Lymantria dispar, Morrisonia latex, Hypoprepia fucosa, Phoberia atomaris, Abagrotis alternata, Catocala amica, C. ilia, Orthosia rubescens, and Acronicta modica. Several of the more abundant species un- der bands in our study were among the more numerous band species in previous re- ports. In 1990 and 1991, Butler and Kondo (1993) noted that the five more abundant band larvae at Coopers Rock (WV) were Lymantria dispar, Abagrotis alternata, Morrisonia latex, Orthosia hibisci (Gue- née), and Halysidota tesselaris (J. E. Smith). Hypoprepia fucosa and Phoberia VOLUME 102, NUMBER 1 a eS Table 1. Continued. Extended. Chestnut White Oak Oak G M ——TGa\ine) 2) Galen, 1 1995. 1996-1 1995. 1996 0) 0) 0 ] 0) 10) 0 0) 0 1 3 l 1 8 10) 10) 0) 0) 0) 1 0) 0) 10) 10) 0) 0) 0) 0) 0) 2, 6) ] 0 1 3 53 9 8 6 48 63 1. 0) 1 0 0) 0) 0) 1 10) 0 0) 0) 0 10) 1 1 0) 1 0 2 l 0) 10) 0 0 0) ] 1 0) 0) 0) 0) 0) 14 5 1 0 0) 10) 0) 3 0) 2 13 11 2, 1] PY, 3 16 0) 0) 0 0) 3 2 0) 8 1 2 0 9 3 10) 18 DD | 6 1) 49 8 20 0 10) 0 0) 0) 1 0) 10) 0) 0 ] 0 0) l 0 0 0 0) 0) 0) | 2 1 0 l 2 0) 1 1] 3 3 16 Og 5 7 O 0) 0) 1 0) 0 l 0 0) 6) 1) 10) l 0 0) 2) 0 | 5 0 5 10) 0) 0 0) | 0 0) 0) 0 0) 0 0) | 0 14 0) 4 l 33 ci l 0) 0) 0 0) 0) 0 ] 0 0 1) 0) 1 0) 0 0) 1 0) 0 10) 0) 2 96 16 28 18 68 182 26 15 2 1 | 46 10 11 1 6) 0) 10) 2 0) 10) 0 0) 0) 0) 0) 1) 0) 0 10) 0) 0 1 0) 0 0) 1 0) 0) ] 0) 2 32 13 a) TE Sl 28 32 l 10) 0) 10) 0) ] 0) 1455 795 504 381 3,400 698 1,729 2,250 885 Years Combined = es ee Grand Seasonal 1996 G M Total Range 1 0) 1 1 13 May—13 May 0) 2 3 5 8 May—6 May 2 l 2 3 3 June—17 June 2 0) 2 2 10 June—17 June 0) 1 3) 4 7 Aug.—15 July 12. 111 24 135. 15 May—17 June 0) 0) 1 l 13 July—1 July O 0) 1 1 24 July—24 July 1 3 1 4 12 June—15 July O 2 0) 2 10 July—15 July O 19 1 20 15 May-—3 June 0) 3 22 5 5 June—19 June 9 30 25 55 8 May-—3 June 0) 5 0) 5 15 May—20 May ] 12 1 13. 22 May-1 July 2 i, 22 79 22 May-12 Aug. 0) ] O ] 6 May—6 May 0 0 | l 12 June—12 June 0) O | 1 24 July—24 July 22 2 3 5) 5 June—12 Aug. 12 52 19 51 17 July—12 Aug. 0) 0) l l 14 Aug.—14 Aug. 0) l 0 l 15 July—15 July 0) 5 5 10 29 May-3 July 0) | 0 1 29 May—29 May 0) | 0) 1 24 June—24 June 0) 40 1 4] 8 May—27 May O O | 1 29 May—29 May 0) | 0) l 15 May—15 May 3 0 5 5 8 May—27 May 39 250 65 315 10 July—12 Aug. 4 56 1S 71 22 May—17 June 1 2 | 3. 22 May—6 May | O l | 10 June—10 June 0) l 0) I 10 July—10 July 0) I 2 3 5 June—19 June 6 79 38 ial 8 May—27 May 0) l 0) l 22 July—22 July 520 4,098 2,249 6,347 atomaris were not among the 40 species of _ band larvae reported for that study. In a 6- year study in the Fernow Experimental For- est, WV, the 5 more abundant species under burlap tree bands were Lymantria dispar, Morrisonia latex, Orthosia rubescens, Aba- grotis alternata, and Lithophane hemina Grote (Butler et al. 1995a). In 1992, Wag- ner et al. (1995) found the 5 more abundant macrolepidoptera at Goshen (Rockbridge Co., VA) to be Phoberia atomaris, Hemi- leuca maia, Dasychira basiflava (Packard), D. obliquata (Grote and Robinson), and Hypoprepia fucosa. In three studies in which macrolepidop- teran larvae were collected from both tree foliage and tree bands, more species of lar- vae were always taken from foliage. How- 196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Temperature and rainfall measurements from May through August, 1995 and 1996 for the George Washington National Forest (GWNF) and Mo- nongahela National Forest (MNF). Temperature (°C) Average Average Rainfall Year Forest Low High (cm) 1995 GWNEF 16.8 28.5 21.8 MNF 1329 23.8 16.8 1996 GWNEF 12.0 Dae BOs MNF 10.0 24.4 49.3 ever, in all three of these studies, several species were taken only from beneath can- vas bands: Butler and Kondo (1993) re- corded 10 species and Butler et al. (1995) recorded 21 species only under bands; Wag- ner et al. (1995) noted 19 species under bands but not on foliage. We do not con- sider any of the species we collected in this study to be rare within our study plots. Many of the larvae poorly represented un- der bands were relatively abundant as sam- pled by other methods (unpublished data). Previous studies have used canvas or burlap bands to evaluate the impact of for- est insecticides including diflubenzuron (Butler and Kondo 1993, Butler et al. 1997) and Bacillus thuringiensis on nontarget in- sects (Wagner et al. 1996), and to study Lepidoptera diversity (Butler et al. 1995a, b; Wagner et al. 1995). From results of our studies, we recommend that tree bands are useful also for evaluating seasonal differ- ences, including weather effects, and im- pact of Entomophaga maimaiga on popu- lations of the gypsy moth. In future publi- cations from this study, we will evaluate impact of Bacillus thuringiensis application to Lepidoptera as sampled by canvas bands, foliage pruning, and blacklight traps. ACKNOWLEDGMENTS This report represents a portion of a study funded by the USDA Forest Service as Cooperative Agreement 42-793. We are grateful for that support. We thank Richard Reardon, USDA Forest Service, and our staff of research assistants, and graduate and undergraduate students. We are partic- ularly grateful to Toby Petrice and Tina Roth Plaugher for their help. Published with the approval of the Director of the West Virginia Agricultural and Forestry Experi- ment Station as Scientific Article # 2711. LITERATURE CITED Butler, L. 1992. The community of macrolepidopter- ous larvae at Coopers Rock State Forest, West Virginia: A baseline study. Canadian Entomolo- gist 124: 1149-1156. Butler, L., G. Chrislip, and V. Kondo. 1995a. Canopy arthropods at Fernow Experimental Forest in West Virginia’s Allegheny Mountain Section: H. Ma- crolepidopterous larvae collected on foliage and under burlap bands. West Virginia University Ex- periment Station Bulletin 713. Butler, L., G. A. Chrislip, V. A. Kondo, and E. C. Townsend. 1997. Effect of diflubenzuron on non- target canopy arthropods in closed deciduous wa- tersheds in a central Appalachian forest. Journal of Economic Entomology 90: 784-794. Butler, L. and V. Kondo. 1993. Impact of Dimilin on non-target Lepidoptera: results of an operational gypsy moth suppression program at Coopers Rock State Forest, West Virginia. West Virginia Uni- versity Experiment Station Bulletin 710. Butler, L., C. Zivkovich, and B. E. Sample. 1995b. Richness and abundance of arthropods in the oak canopy of West Virginia’s Eastern Ridge and Val- ley section during a study of impact on Bacillus thuringiensis with emphasis on macrolepidoptera larvae. West Virginia University Experiment Sta- tion Bulletin 711. Ferguson, D. C. 1971. Bombycoidea (Saturniidae Part), Fascicle 20.2A. The Moths of America North of Mexico. E.W. Classey Ltd. and R.B.D. Publications Inc., London, 153 pp. Forbes, W. T. M. 1954. Lepidoptera of New York and Neighboring States, Part II. Cornell University Agricultural Experiment Station Memoir 329, 433 PP- . 1960. Lepidoptera of New York and Neigh- boring States, Part IV. Cornell University Agri- cultural Experiment Station Memoir 371, 188 pp. Franclemont, J. G. 1973. Mimallonoidea and Bomby- coidea, Fascicle 20.1. The Moths of America North of Mexico. E.W. Classey Ltd. and R.B.D. Publications, Inc., London, 86 pp. Hodges, R. W., T. Dominic, D. R. Davis, D. C. Fer- guson, J. G. Franclemont, E. G. Munroe, and J. A. Powell. 1983. Checklist of the Lepidoptera of America North of Mexico. London. EW. Classey, Ltd. and Washington, D.C. Wedge Entomological Research Foundation. VOLUME 102, NUMBER 1 Marquis, R. J. and S. Passoa. 1989. Seasonal diversity and abundance of the herbivore fauna of striped maple, Acer pensylvanicum L. (Aceraceae) in West Virginia. American Midland Naturalist 122: 313-320. Martinat, P. J. 1987. The role of climatic variation and weather in forest insect outbreaks, pp. 241—268. In Barbosa, P. and J. C. Shultz, eds., Insect Out- breaks. Academic Press, San Diego. Miller, J. C. 1990. Field assessment of a microbial pest control agent on nontarget Lepidoptera. American Entomologist 36: 135-139. Poole, R. W. and P. Gentili (eds.). 1986. Nomina In- secta Nearctica. A Checklist of the Insects of North America, Vol. 3. Diptera, Lepidoptera, Si- phonaptera. Entomological Information Services, Rockville, Maryland, 1143 pp. 197 Stevens, R. E., V. M. Carolin, and G. P. Markin. 1984. Lepidoptera associated with western spruce bud- worm. United States Department of Agriculture, Forest Service Agricultural Handbook No. 622. 63 Pp. Wagner, D. L., J. W. Peacock, J. L. Carter, and S. E. Talley. 1995. Spring caterpillar fauna of oak and blueberry in a Virginia deciduous forest. Annals of the Entomological Society of America 88: 416— 426. 1996. Field assessment of Bacillus thurin- giensis on nontarget Lepidoptera. Environmental Entomology 25: 1444-1454. Wood, P. S. and L. Butler. 1989. Biology and immature stages of Polia latex (Guenée) (Noctuidae). Jour- nal of the Lepidopterists’ Society 43: 299-304. PROC. ENTOMOL. SOC. WASH. 102(1), 2000, pp. 198-211 REVIEW OF THE CYNIPID GALL WASPS OF THE GENUS LOXAULUS MAYR (HYMENOPTERA: CYNIPIDAE) WITH DESCRIPTIONS OF NEW SPECIES GEORGE MELIKA AND WARREN G. ABRAHAMSON Department of Biology, Bucknell University, Lewisburg, PA 17837, U.S.A.; (GM) per- manent address: Systematic Parasitoid Laboratory, Kelcz-Adelffy St. 6, Koszeg 9700, Hungary (GM e-mail: chalcini@savaria.hu; WGA e-mail: abrahmsn @bucknell.edu) Abstract.—Diagnostic characters of the genus Loxaulus Mayr and a key to all fourteen species are given. Two species are newly synonymized: Compsodryoxenus humilis Weld is a new synonym of Loxaulus vaccinii (Ashmead) and Loxaulus ashmeadi Kieffer is a new synonym of Loxaulus quercusmammula (Bassett). Lecto- and paralectotypes of L. beutenmuelleri Weld, C. ferrugineus Gillette, C. quercusmammula Bassett, and C. tenuis Weld are designated. Two new species, L. huberi and L. masneri are described. Key Words: ology Burks (1979) listed 12 species and one variety for the genus Loxaulus Mayr. Sub- sequently, a new species, L. boharti Dailey and Sprenger, was described and species status was given to L. brunneus var. atrior (Kinsey) (Dailey 1977, Dailey and Spren- ger 1983). Thus, the current number of Lox- aulus species is 14. Herein we synonymize two species, describe 2 new species, L. hub- eri and L. masneri, and provide a key to all 14 species. The identification of the gall wasps of the North American genus Loxaulus Mayr (Hy- menoptera, Cynipidae, Cynipini) is difficult because the existing keys (Weld 1921, 1952) use a number of unsatisfactory and inadequate diagnostic characters. Further- more, the nomenclature used to describe taxonomically important structures in the publications prior to the 1960s is inconsis- tent with current terminology (Dailey and Menke 1980). Various authors have used different terminology to define the same morphological structure and employed the Cynipidae, gall wasps, Loxaulus, taxonomy, morphology, distribution, bi- same term for different structures. As a con- sequence, species have been misunderstood and misidentified. We follow the current terminology of morphological structures as given in Gibson (1985), Ronquist and Nordlander (1989), Menke (1993), and Fergusson (1995). Ab- breviations for fore wing venation follow Ronquist and Nordlander (1989). The mea- surements and abbreviations used herein in- clude: FI1—F12, Ist and subsequent flagel- lomeres; POD (post-ocellar distance), the distance between the inner margins of the posterior ocelli; and OOD (ocellar-ocular distance), the distance from the outer edge of a posterior ocellus to the inner margin of the compound eye. The width of the radial cell is measured along 2r. Drawings were made with the aid of a Wild™ drawing tube, scanned into a PC and modified in Adobe Photoshop 3.0. Material was examined from the follow institutions: National Museum of Natural History, Smithsonian Institution (USNM), VOLUME 102, NUMBER 1 Washington, DC, USA; Canadian National Insect Collection, Ottawa, Canada (CNCI); American Museum of Natural History, New York, NY, USA (AMNH); and from the G. Melika private cynipid collection. Loxaulus Mayr Loxaulus Mayr 1881: 9, 12, 33. Type spe- cies: Loxaulus mammula Mayr. Mono- typic. Solenozopheria Ashmead 1887: 149. Type species: Solenozopheria vaccinii Ash- mead. Monotypic; Weld 1951: 643 (syn- onym of Loxaulus Mayr). Compsodryoxenus Ashmead 1896: 128. Type species: Compsodryoxenus maculi- pennis Ashmead. Designated by Ash- mead 1903: 155; Weld 1921: 234 (key to species); Weld 1951: 643 (synonym of Loxaulus Mayr). Diagnosis.—Length 2.5 mm or less. Head in dorsal view broader than thorax; gena with malar sulcus (Figs. 6, 12). Scu- tum usually finely transversely coriaceous; scutellum without foveae, with a transverse shallow depression (Figs. 9, 15). Central portion of propodeum narrow, limited by parallel or slightly outward bent lateral ca- rinae and with a median longitudinal carina and/or longitudinal striae; median longitu- dinal carina in some species indistinct, frag- mented, but always present at least in an- terior half (Figs. 11, 17). Radial cell of fore- wing short and broad, not more than 2.5 times as long as broad (except L. quercus- mammula with radial cell 2.6—2.7 times as long as broad), forewing margin of female with or without cilia, with brown, smoky spots (or stripes) along areolet, 2r, Rs, and M (Figs. 1, 2, 10, 16). Tarsal claws without tooth. Ventral spine of hypopygium short, slender or needle-like; subapical setae short and sparse, not reaching beyond apex of spine and prominent part never more than 3.03.5 times as long as broad (Figs. 8, 14). Discussion.—The propodeum is similar to that of the Mediterranean genus Plagi- 199 otrochus Mayr. However, in the latter, the central portion of the propodeum is much broader, and is limited by the strongly out- ward-bending lateral carinae (Fig. 3). Distribution.—North America only. Biology.—Alternate unisexual and bisex- ual generations are known only for L. tri- zonalis Weld. Weld (1921) published a key to five spe- cies. However, a manuscript key to Loxau- lus species written by Weld includes all the known species and also a number of un- described species. Weld’s manuscript key is on file at the USNM. We credit Weld’s use- ful notes and manuscript keys to the Cyni- pidae. Herein, we offer a key to all known 14 species. KEY TO SPECIES OF LOXAULUS 1. Female; antenna 12-14 segmented ....... 3 — Male; antenna 15-segmented ........... 2 2. Forewing banded, with two extensive smoky brown spots, radial cell 2.3—2.4 times as long as) broad) (Big. .2) is cess stents aw eer trizonalis — Forewing without smoky spots, or with only a slight indication of very narrow darkish ar- eas along M and around areolet, radial cell 2.6—2.7 times as long as broad (Fig. 1) .... Aa. acy od epeus, Seon eeise uses os quercusmammula 3; ~Antennael4-seomented 52.25.25: oc o.c.5 oon 4 = Antenna l2—I3 sesmented 2. 0265205 anse 6 4. Forewing banded, with three extensive smoky brown spotsi(Fig2)) 4 22 bw. Pewantes Ss ok 5 — Forewing with narrow dark stripes along M, Rs erM and: 2r(Bi S51) ssc. quercusmammula 5. Scutum and scutellum same light brown color AaSUSIGES Of INESOSOMA: seaserog. 2c eep eee he Sart Aris Nee eae ae trizonalis, bisexual generation — Scutum and scutellum dark reddish-brown, much darker than light brown sides of me- SOSOMA if. sia. s trizonalis, unisexual generation 6. Antenna 13-segmented, with distinct suture between FIO and gh 5. Bake ens est © 2 qi — Antenna 12-segmented, indistinct suture in- dicates Fll whose length equals to F9+F10 RN AE RAM usa EN CHER SS 1c SPORE ROR eee tenuis 7. Forewing margin with cilia very short, or ab- sent, with dark spots along areolet, M, and Rs (Fig. 4) — Forewing margin with long cilia, with or without distinct darker spots, but with darker stripes along M and Rs present (Fig. 5) .. 11 8. F12 divided medially by an indistinct suture; F12 nearly equal to FIO+F11; disk of scutel- 200 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1 = Sess 3 Figs. 1-5. 1-2, Forewing. 1, Loxaulus quercusmammula. 2, L. trizonalis, unisexual female. 3, Plagiotrochus quercusilicis, propodeum. 4—5, Forewing. 4, L. maculipennis. 5, L. vaccinii. VOLUME 102, NUMBER 1 . Scutum longer than broad lum mostly coriaceous, rugose behind and on sides; length less than 2.0 mm F12 not divided by a suture; F11 longer than F9+F10; disk of scutellum rugose, edges co- miaceous; length 3:6%mm* =. -... . maculipennis Head more than 1.25 times as wide as high, occiput round; inner orbits ventrally diverg- ing; prominent part of ventral spine of hypo- pygium 1.5—2.0 times s long as broad .. . Head less than 1.25 times as wide as high, occiput flattened; inner orbits parallel; prom- inent part of ventral spine of hypopygium 2.0—3.0 times as long as broad ......... 10 . Notauli complete; prominent part of ventral spine of hypopygium 2.0—2.5 times as long AGHDTOAG eas a nS ates Rents digs 3 oe ys brunneus Notauli incomplete; prominent part of ventral spine of hypopygium 3.0 times as long as (araeral 325% eee ce ee ere = hee eee boharti . Forewing with dark smoky spot(s) around ar- ecoletwand/oraloms