ce et ’ we Aton ma FN Sage eat : “ aay ~! VEL MEY Ske ES SE ean She a) SDK a eee coogi peiet tow. - ‘ ? etm qe: . ¢ rary eee RAD ated ‘ : ‘ aati! Secnes oh : Sattederns PRS: ate ot Spat. te ee eh autores es evra tape ters 5 er | ris 4 | VOL. 104 JULY 2002 NO. 3 =f, 9 kK (ISSN 0013-8797) ; PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON > PUBLISHED 1S ay, \ QUARTERLY CONTENTS CONTENTS BLANK, STEPHAN M.—Taxonomic notes on Strongylogasterini (Hymenoptera: Tenthredinidae) 692 BUFFINGTON, MATTHEW L.—Description of Aegeseucoela Buffington, new name, with notes on the status of Gronotoma Forster (Hymenoptera: Figitidae: Eucoilinae) .................... 589 CHABOO, CAROLINE S.—Range extensions of New World tortoise beetles (Coleoptera: Ghrysomelidac2CassiGinaey ies cre acre le fete Serelels Hie zeus one SIS eine een st ec eres 716 FLORES, GUSTAVO E. and CHARLES A. TRIPLEHORN—Entomobalia, new genus, the first member of Nycteliini (Coleoptera: Tenebrionidae) from Brazil .....................-....00 ees 602 GOEDEN, RICHARD D.—Life history and description of immature stages of Oxyna palpalis (Coquillett) (Diptera: Tephritidae) on Artemisia tridentata Nuttall (Asteraceae) in southern GalitOrnii al rests ce ee erecta tis acs Annee mttete ms ROP arated ci vce ha stl ig een ne aM Cte nae eR eee ae 537 GOEDEN, RICHARD D.—Life history and description of immature stages of Goedenia rufipes (Curran) (Diptera: Tephritidae) on Jsocoma acradenia (E. Greene) E. Greene in southern (CPAIIFOVTINE es ee ad bes Sania bed te anaes ti A Ga SO ee Att Rae ROPE AES SRA ino RSE OuE Roos atone Broa. 576 GOEDEN, RICHARD D.—Life history and description of immature stages of Goedenia setosa (Foote) (Diptera: Tephritidae) on Ericameria brachylepis (A. Gray) H. M. Hall in southern @alifommi as pie acenet: eh oak ears sehes Gh ete ele aaa oe ete sates, be zieiae aye aS REE BS Ae Ce eats aR SERe hala eae 629 GOEDEN, RICHARD D.—Life history and description of adults and immature stages of Goedenia stenoparia (Steyskal) (Diptera: Tephritidae) on Gutierrezia californica (de Candolle) Torrey and A. Gray and Solidago californica Nuttall (Asteraceae) in southern (Bealeton ope hee a oor aaa olotesrak ig Gas tans biadielait Sidhe. Lalagate Beanie as. RETA EMM eRe aayorere eo a Rena Ee 702 GOEDEN, RICHARD D.—Life history and description of adults and immature stages of Goedenia steyskali, n. sp. (Diptera: Tephritidae) on Grindelia hirsutula Hooker and Arnott var. halli (Steyermark) M. A. Lane (Asteraceae) in southern California ...................... 785 HARRISON, B. A., P. B. WHITT, S. E. COPE, G. R. PAYNE, S. E. RANKIN, L. J. BOHN, F. M. STELL, and C. J. NEELY—Mosquitoes (Diptera: Culicidae) collected near the Great Dismal Swamp: New state records, notes on certain species, and a revised checklist for Virginia .... 655 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ) OFFICERS FOR 2002 GABRIELA CHAVARRIA, President MICHAEL G. PoGuE, Treasurer JONATHAN R. Mawps.Ley, President-Elect RONALD A. OcHoa, Program Chair Stuart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Ho us B. WILLIAMS, Corresponding Secretary JOHN W. Brown, Past President Jon A. Lewis, Custodian Davip R. Smirn, Editor Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. MaruHis Honorary President Louise M. RussELL Honorary Members KARL V. KROMBEIN RONALD W. HopGEs 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 7:00 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $25.00 (U.S. currency). PROCEEDINGS.—tThe 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. 247-248 OF THE JANUARY 2002 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 26 June 2002 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 537-553 LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF OXYNA PALPALIS (COQUILLETT) (DIPTERA: TEPHRITIDAE) ON ARTEMISIA TRIDENTATA NUTTALL (ASTERACEAE) IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e- mail: richard.goeden @ucr.edu) Abstract.—Oxyna palpalis (Coquillett) is a univoltine, circumnatal tephritid uniquely reproducing as an inquiline in rosette galls of Rhopalomyia florella Gagné (Diptera: Ce- cidomyiidae) of terminal buds on branches of Artemisia tridentata Nuttall. Its larvae also routinely function as facultative predators of R. florella larvae, novel behavior for Te- phritidae. The egg, first-, second-, and third-instar larvae, and puparia are described and figured for the first time. The egg is distinguished by a pedicel circumscribed by one complete and a partial second ring of irregularly shaped micropyles. All three larval instars of O. palpalis are compared to and distinguished from those of O. aterrima (Doane), its only other known congener in North America. Oviposition occurs in spring (June) in southern California in nearly fully formed galls of R. florella containing young larvae of this cecidomyiid. The young larvae of O. palpalis pass the summer (June—September) as first instars singly in small, central, ovoidal cells basad of the cecidomyiid larvae. Second instars occupy their still-small, separate chambers until late fall/early winter (September— October), when some begin to molt to third instars. By mid-winter (February) all larvae are third instars, which continue to overwinter and grow slowly until the resumption of the spring flush of new plant growth (March). At this time, one to six or more third instars enlarge the central gall chamber to accommodate their faster growth and feed gregariously. Cecidomyiid larvae encountered during this third stadium are killed and devoured; sur- viving immature gall midges usually occupy the periphery of the galls. Pupariation follows in early April, and adults emerge by mid-April. Eurytoma sp. (Hymenoptera: Eurytomi- dae) and Eupelmus sp. (Hymenoptera: Eupelmidae) were individually reared from puparia of O. palpalis as primary, solitary, probably larval-pupal endoparasitoids. Lyrcus sp. (Hy- menoptera: Pteromalidae) was reared from individual puparia as a gregarious, primary endoparasitoid. Key Words: Insecta, Oxyna, Asteraceae, Artemisia, nonfrugivorous Tephritidae, Rho- palomyia, Cecidomyiidae, biology, taxonomy of immature stages, galls, inquiline, circumnatal life cycle, parasitoids, insect predation Oxyna palpalis (Coquillett) (Diptera: Te- O. aterrima (Doane) was reviewed by phritidae) is one of two species of Oxyna Foote et al. (1993) and studied by Goeden now known from North America (Foote et (2002b), who synonymized it with O. utah- al. 1993, Goeden 2002b). The other species, ensis Quisenberry. MATERIALS AND METHODS The present study was based in large part on dissections of samples of galls of the gall midge, Rhopalomyia florella Gagné (Diptera: Cecidomyiidae), on Artemisia tri- dentata Nuttall prob. ssp. parishii (A. Gray) H. M. Hall and Clements (Asteraceae) col- lected mainly 0.2 km north of the hamlet of Mile High and just south of the hamlet of Largo Vista; 1580-m elevation; Township 4N, Range 9W, Section 4; Angeles National Forest, Los Angeles Co., during 1996 and 1997. Excised R. florella galls, containing eggs and early-instar larvae of O. palpalis, and later, overwintered galls containing third instars and puparia of O. palpalis were sampled mid-monthly from gall-bearing plants during 1996 and 1997. Samples were transported in cold-chests in an air-condi- tioned vehicle to the laboratory and stored under refrigeration for subsequent dissec- tion, photography, description, and mea- surement. Eight eggs, 18 first-, seven sec- ond-, and 19 third-instar larvae and nine puparia dissected from galls were preserved in 70% EtOH for scanning electron micros- copy (SEM). Additional prepuparia and pu- paria in excised, opened galls were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temperature for adult and parasitoid emergence. Specimens for SEM were hydrated to distilled water in a de- creasing series of acidulated EtOH. They were osmicated for 24 h, dehydrated through an increasing series of acidulated EtOH and two, |-h immersions in hexa- methyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, and studied and photographed with a Philips XL-30 scanning electron micro- scope in the Central Facility for Advanced Microscopy and Microanalysis, University of California, Riverside. Most adults reared from isolated prepu- paria and puparia were individually caged in 850-ml, clear-plastic, screened-top cages with a cotton wick and basal water reser- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON voir and provisioned with a strip of paper toweling impregnated with yeast hydroly- zate and sucrose. These cages mainly were used for studies of longevity in the insec- tary of the Department of Entomology, University of California, Riverside, at 25 + 1°C, and 14/10 (L/D) photoperiod. Two pairs of virgin flies, each consisting of a male and a female obtained from emer- gence cages also were held in a clear-plas- tic, petri dish provisioned with a flattened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1994) for observations of their courtship and copulation behavior. Plant names used in this paper follow Hickman (1993) and Bremer (1994); te- phritid names and adult terminology follow Foote et al. (1993); cecidomyiid names and gall terminology follow Gagné (1989). Ter- minology and telegraphic format used to describe the immature stages follow Goe- den (2001, 2002a, b), Goeden and Norrbom (2001), Goeden and Teerink (1999), and earlier works cited therein. Means + SE are used throughout this paper. All remaining voucher specimens and reared parasitoids of this tephritid reside in my research col- lections. Digital photographs used to con- struct text figures were processed with Ado- be Photoshop® Version 6. RESULTS AND DISCUSSION Adult.—Oxyna_ palpalis originally was described as a Tephritis by Coquillett in Baker (1904) from a single male specimen from Ormsby County, Nevada. Quisenberry (1949) in his revision of Oxyna redescribed this species from three females and two males variously from California, Idaho, and Nevada. Foote and Blanc (1963) and Foote et al. (1993) pictured the right wing. Immature stages.—The egg, first-, sec- ond-, and third-instar larvae, and puparium of O. palpalis are described below. Egg: Thirteen eggs of O. palpalis dis- sected from field-collected galls of R. flo- rella galls also bearing cecidomyliid larvae were white, opaque, smooth, ellipsoidal, VOLUME 104, NUMBER 3 Fig. 1. Egg of Oxyna palpalis: (A) habitus, anterior to left; (B) pedicel 1, (C) pedicel 2, (D) pedicel 3. (The three pedicels show variation in size, shape, and placement of micropyles.) 0.65 + 0.02 (range, 0.62—0.68) mm long, 0.20 + 0.00 (range, 0.20—0.20) mm wide, smoothly rounded at basal end, with a 0.2- mm, buttonlike pedicel at anterior end (Fig. 1A). The pedicel was circumscribed suba- pically by one complete and a partial sec- ond ring of irregularly shaped micropyles (Bigs IBC LD) This is the first Oxyna egg pictured at high magnification (Goeden 2002b). On av- erage, the egg of O. palpalis is slightly lon- ger and slightly narrower than that of O. aterrima (Goeden 2002b). First instar larva: White, ellipsoidal, flat- tened anteriorly and posteriorly (Fig. 2A); body segments with hemispherical or pos- teriorly-directed, short-spinose, minute acanthae on intersegmental areas of meta- thorax and abdominal segments Al through A6 as well as pleura and lateroventrum of Al through A6; prothorax and gnathoce- phalon smooth, the latter conical (Fig. 2B), both circumscribed by verruciform sensilla (Figs. 2B-1, -2); dorsal sensory organ, well- defined, flat pad (Fig. 2C-1); anterior sen- sory lobe (Fig. 2C-2) bears terminal sen- sory organ (Fig. 2C-3), lateral sensory or- gan (Fig. 2C-4), supralateral sensory organ (Fig. 2C-5), and pit sensory organ (Fig. 2C- 6); stomal sense organ reduced to two ver- ruciform sensilla ventrolaterad of terminal sensory organ (Figs. 2C-7, D-1), not fused with flattened, protrudent, lateral integu- mental petal (Figs. 2C-8, D-2) above each mouthhook, one medial, papillate, integu- mental petal (Figs. 2C-9, D-3) between an- terior sensory lobes and lateral integumen- tal petals; mouthhook bidentate (Figs. 2B- 3, C-10, D-4); median oral lobe laterally compressed, apically rounded (Figs. 2B-4, 540 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. trolateral view, 1—verruciform sensilla on gnathocephalon, 2—verruciform sensilla on prothorax, 3—mouth- First instar of Oxyna palpalis: (A) habitus, anterior to left, (B) gnathocephalon and prothorax, ven- hook, 4—median oral lobe; (C) gnathocephalon, frontal, close-up view, 1—dorsal sensory organ, 2—anterior sensory lobe, 3—terminal sensory organ, 4 lateral sensory organ, 5—supralateral sensory organ, 6—pit sensory organ, 7—stomal sense organ, 8—lateral integumental petal, 9—medial integumental petal, 10—mouthhook; (D) oral cavity, ventral view, 1—stomal sense organ, —lateral integumental petal, 3—medial integumental petal, 4—mouthhook, 5—median oral lobe, 6—labial lobe, 7—pores. D-5); labial lobe (Fig. 2D-6) broad, sepa- rated from median oral lobe, and with two pores ventrally (Fig. 2D-7); anterior spira- cle absent; lateral spiracular complexes not seen; caudal segment (Fig. 3A) with a ste- lex sensillum dorsolaterad (Fig. 3A-1), lat- (Figs 3A-2, B), ventrolaterad erad and (Figs. 3A-3, C-1) of posterior spiracular plate (Figs. 3A-4, D); posterior spiracular plate bears two rimae (Fig. 3D-1), ca. 0.005 mm long, and four, unbranched or bifurcate, spinose or apically toothed (some bifurcate) interspiracular processes (Fig. 3D-2), the longest process measuring 0.007 mm; in- termediate sensory complex (Figs. 3A-5, C- 2, D-3) consists of stelex sensillum (Fig. 3C-3) and medusoid sensillum (Fig. 3C-4). The habitus of the first instar of O. pal- palis is similar to the first instar of O. ater- however, as with Trupanea_ spp. (Goeden and Teerink 1999), Neaspilota spp. (Goeden 2001), and Tephritis spp. (Goeden 2002a), the incidence and patterns of minute acanthae on the thorax and ab- domen show interspecific differences. Ox- yna palpalis has fewer minute acanthae in the intersegmental areas of the thorax and rima; abdominal segments Al and A2, and unlike O. aterrima, none on or between abdominal segments A3 through A6 (Goeden 2002b). VOLUME 104, NUMBER 3 Figs 3: lateral verruciform sensillum, 3—ventrolateral stelex sensillum, 4—posterior spiracular plate, 5—intermediate First instar of Oxyna palpalis, continued: (A) caudal segment, 1—dorsolateral stelex sensilla, 2— sensory complexes; (B) lateral stelex sensillum (arrow); (C) 1—ventrolateral verruciform sensillum, 2—inter- mediate sensory complex (composed of), 3—stelex sensillum, 4—medusoid sensillum; (D) posterior spiracular plate with 1—two rimae, 2—four interspiracular processes, and 3—two intermediate sensory complexes. The first instar of O. aterrima has two, medial, integumental petals (Goeden 2002b); whereas, O. palpalis has only one (Figs. 2C-9, D-3). The single, lateral inte- gumental petal above each mouthhook is fused with the stomal sense organ in O. aterrima (Goeden 2002b), but they are not fused in O. palpalis (Figs. 2C-8, D-2). The mouthhooks are tridentate in the first instar of O. aterrima (Goeden 2002b), but are bi- dentate in O. palpalis (Figs. 2B-3, C-10, D- 4). The lateral sensilla surrounding the pos- terior spiracular plate of O. aterrima are verruciform (Goeden 2002b; whereas, these are stelex sensilla in O. palpalis (Figs 3A- 2, B). The interspiracular processes on the caudal segment of O. aterrima are un- branched (Goeden 2002b); whereas, some processes of O. palpalis are two-branched (Fig. 3D-2). Second instar larva: Ovoidal, rounded anteriorly, truncated posteriorly (Fig 4A), body segments with short-spinose, posteri- orly-directed, minute acanthae (Fig. 4B-1) circumscribing intersegmental areas of tho- rax and abdomen; thorax and gnathoce- phalon smooth (Fig. 4B), the latter conical, the former circumscribed around middle with verruciform sensilla (Fig. 4B-1); dor- sal sensory organ, a well-defined, flat pad (Figs. 4C-1, D-1); anterior sensory lobe (Fig. 4C-2) with terminal sensory organ (Fig. 4C-3), lateral sensory organ (Fig. 4C- 4), supralateral sensory organ (Fig. 4C-5), and pit sensory organ (Fig. 4C-6); two pairs of foliose, protrudent, lateral integumental 542 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Second instar of Oxyna palpalis: (A) habitus, anterior to left; (B) prothorax and gnathocephalon, frontolateral view, 1—minute acanthae, 2—verruciform sensillum; (C) gnathocephalon, frontolateral view, 1— dorsal sensory organ, 2 anterior sensory lobe, 3—terminal sensory organ, 4—lateral sensory organ, 5—supra- lateral sensory organ, 6—pit sensory organ, 7—lateral integumental petals, 8—medial integumental petals; (D) gnathocephalon, ventrolateral view, 1—dorsal sensory organ, 2 integumental petals, 4—oral ridges, 5—-stomal sense organ, 6—mouthhook, 7—median oral lobe, 8 9—pores. petals (Figs 4C-7, D-2) above each mouth- hook, two pairs of papillate, medial inte- gumental petals (Fig. 4C-8) between each anterior sensory organ, the ventral pair elongate (Fig. 4D-3); at least four oral ridg- es (Fig. 4D-4) ventrolaterad of each anterior sensory lobe and stomal sense organ (Fig. 4D-5); mouthhook (Fig. 4D-6) with two teeth; median oral lobe laterally flattened (Fig. 4D-7); labial lobe (Fig. 4D-8) broad, separated from median oral lobe, with two pores ventrally (Fig. 4D-9); anterior thorac- ic spiracle (Fig. 5A) with five, rounded, wedge-shaped papillae; lateral spiracular complexes not seen; caudal segment (Fig. 5B) with stelex sensillum dorsolaterad lateral integumental petals, 3—ventral, medial labial lobe, (Figs. 5B-1), laterad (Fig 5B-2), and ven- trolaterad (Figs. 5B-3, C-1) of posterior spi- racular plate (Figs. 5B-4, D); posterior spi- racular plate with four, elongate, upright, foliose, interspiracular processes (Fig. 5D- 1), the longest process measuring 0.011 mm; intermediate sensory complex (Figs. 5B-5, C-2) consists of stelex sensillum (Fig. 5C-3) and medusoid sensillum (Fig. 5C-4). One major difference between the second instars of O. palpalis and O. aterrima is the absence of a black marking on the abdom- inal ventrum of the former species, where the ventrum and pleura of the latter species are densely covered with knoblike minute acanthae (Goeden 2002b). In O. palpalis VOLUME 104, NUMBER 3 Fig. 5. 543 i Wh u (UGE Se Ne aS 4) ae ‘i A aj \\ ie, ¥ f az ‘“ ES a iff aS fh) - ‘ i a \ Second instar of Oxyna palpalis, continued: (A) anterior spiracle; (B) caudal segment; caudal seg- ment, 1—dorsolateral stelex sensilla, 2—lateral stelex sensilla, 3—ventrolateral stelex sensilla, 4—posterior spiracular plate, 5—intermediate sensory complexes; (C) 1—ventrolateral stelex sensillum, 2—intermediate sen- sory complex (composed of), 3—stelex sensillum, 4—medusoid sensillum; (D) posterior spiracular plate, 1— four interspiracular processes. the minute acanthae are short-spinose and confined mainly to the intersegmental areas of the thorax and abdomen. Oxyna palpalis has two pairs of medial integumental petals between each anterior sensory lobe (Fig. 4C-8); whereas, O. aterrima has only one such pair (Goeden 2002b). The inner, ven- tral oral ridge among the four such ridges of O. aterrima is ventrally toothed and fused with the stomal sense organ (Goeden 2002b); whereas, none of the four oral ridg- es of O. palpalis is toothed nor fused with the stomal sense organ (Figs. 4D-3, 4). Third instar larva: Pale yellow or white, ovoidal, tapering anteriorly, truncated pos- teriorly, distinctly segmented (Fig. 6A), short-spinose, posteriorly-directed, minute acanthae in transverse bands on dorsopos- terior fifth of gnathocephalon (Figs. 6A, B- 1), minute acanthae also circumscribe an- terior fourth of thoracic segments and an- terior fourth of abdominal segment A1, dor- sum, ventrum and anterior third of pleura of A2-A7, and all but posterior of spiracular plate A-8 (Fig. 7D-1); gnathocephalon smooth, conical (Fig. 6-B); dorsal sensory organ well-defined, hemispherical (Figs. 6B-2, C-1); anterior sensory lobe bears ter- minal sensory organ (Figs. 6B-3, C-2), lat- eral sensory organ (Fig. 6C-3), supralateral sensory organ (Fig. 6C-4), and pit sensory organ (Fig. 6C-5); nine or 10 oral ridges (Fig. 6D-1) laterad and ventrolaterad of an- terior sensory lobe and stomal sense organ, at least six lobes ventrally toothed and sep- arate from prominent stomal sense organ 544 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Third instar of Oxyna palpalis: (A) habitus, anterior to left; (B) gnathocephalon, ventrolateral view, I—minute acanthae, 2—dorsal sensory organ, 3—terminal sensory organ, 4—stomal sense organ, 5—mouth- hook, 6—median oral lobe, 7—lateral integumental petal, 8—inner, lateral integumental petals; (C) gnatho- cephalon, dorsolateral view, 1—dorsal sensory organ, 2—terminal sensory organ, 3—lateral sensory organ, 4— supralateral sensory organ, 5—pit sensory organ, 6—stomal sense organ, 7—medial integumental petals, 8— lateral integumental petals, 9—inner, lateral integumental petals; (D) gnathocephalon and oral cavity, ventrolat- eral view, 1—oral ridges, 2—three-toothed mouthhook, 3—median oral lobe, 4 5—inner, lateral integumental petal, 6 lateral integumental petals, labial lobe, 7—pores; (E) anterior spiracle with three papillae; (F) anterior spiracle with four papillae. VOLUME 104, NUMBER 3 (Fig. 6B-4, C-6); mouthhook (Fig. 6B-5, D- 2) tridentate (Fig. 5D-2); median oral lobe laterally flattened, apically rounded (Figs. 6B-6, D-3); three pairs of medial integu- mental petals in vertical row between an- terior sensory lobes (Fig. 6C-7); five, lateral integumental petals between each mouth- hook and anterior sensory lobe, including four foliose, lateral petals (Fig. 6B-7, C-8, D-4) and single, inner, elongate, papillate, lateral petal (Figs. 6B-8, C-9, D-5); labial lobe (Fig. 6D-6) broad, separated from me- dian oral lobe, and with two pores ventrally (Fig. 6D-8); anterior thoracic spiracle with three (Fig. 6E, 7A-1) or four, rounded, wedge-shaped papillae (Fig. 6F); lateral spiracular complex of mesothorax with closed, relict spiracle (Fig. 7A-2) and six verruciform sensilla (Figs. 7A-3) in vertical row posteriorad of spiracle, additional ver- ruciform sensillum posteriorad of fourth- most-vertical sensillum; lateral spiracular complex of metathorax similarly composed of closed, relict spiracle (Figs. 7A-4, B-1) and four verruciform sensilla (Figs. 7A-5, B-2) similarly positioned; lateral spiracular complex of first abdominal segment com- posed of closed, presumably relict spiracle (Figs. 7A-6, C-1) and three verruciform sensilla (Figs. 7A-7, C-2) similarly posi- tioned; posterior spiracular plate (Figs. 7D- 1, 8A) bears three, broadly elliptical rimae (Fig. 8A-1), ca. 0.03 mm long, and four, unbranched, spiniform, interspiracular pro- cesses (Fig. 8A-2), each ca. 0.007 mm long; stelex sensilla dorsolaterad (Figs. 7D-2, 8B), verruciform sensilla laterad (Figs. 7D- 3, 8C), and stelex sensilla ventrolaterad (Fig. 7D-4) of posterior spiracular plate; in- termediate sensory complexes (Figs. 7D-5, 8D) consist of stelex sensillum (Fig. 8D-1) and medusoid sensillum (Fig. 8D-2). Fortunately, the third instars of both spe- cies of Oxyna known from North America have now been described in considerable detail, facilitating comparison between them. For example, the minute acanthae on third instars of O. aterrima are fewer in number, occupy fewer body segments, and 545 different (Goeden 2002b) than the minute acanthae on O. palpalis. Oxyna palpalis has at least nine or 10 oral form patterns ridges (Fig. 6D-1), six of which have ven- trally toothed margins; whereas, O. aterri- ma has only two oral ridges (Goeden 2002b). All five lateral integumental petals of third instar O. aterrima are papillate (Goeden 2002b); whereas, only one of the five lateral integumental petals of the third instar of O. palpalis is papillate, the rest are foliose (Figs. 6B-7, C-8, D-4). The lateral spiracular complex of the mesothorax of O. has four verruciform sensilla (Goeden 2002b); whereas, this same com- plex in O. palpalis has seven verruciform sensilla (Figs. 7A-3). Similarly, the lateral spiracular complex of the first abdominal segment of O. aterrima has four verruci- form sensilla, two pairs in separate vertical rows, (Goeden 2002b); whereas, this same complex in O. palpalis has three, vertical, verruciform sensilla (Figs. 7A-7, C-2). The caudal segments of third instars of these congeners also differ considerably. The dorsolateral, lateral, and ventrolateral sensilla surrounding the posterior spiracular plate of O. aterrima are all verruciform. The intermediate sensory complex of this species also uniquely consists of two ver- ruciform sensilla, and this composition dis- tinguishes O. aterrima from third instars of all other nonfrugivorous tephritids exam- ined by my coworkers and me to date (Goe- den 2002b). In O. palpalis, the dorsolateral and ventral lateral sensilla are stelex in form (Figs. 7D-2, -4E, 8B), and the inter- mediate sensory complex is comprised of a stelex sensillum (Fig. 8D-1) and a medu- soid sensillum (Fig. 8D-2), like all other third instars described by us to date. Differences noted between the first and second instars of O. palpalis include the usual acquisition of an anterior spiracle in the second instar, the increase in the number of lateral integumental petals from one to three, and the drastic change in shape of the interspiracular processes (Figs. 3D-2, 5D- 1). Differences noted between the second aterrima 546 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bice a metathoracic spiracle, 3—verruciform sensillum, 4—mesothoracic spiracle, 5—verruciform sensillum, 6—first abdominal segment spiracle, 7—verruciform sensillum; (B) part of lateral spiracular complex of metathorax, 1— spiracle, 2—verruciform sensillum, (C) part of lateral spiracular complex of metathorax, 1—spiracle, 2—ver- ruciform sensillum; (D) anal segment, 1—posterior spiracular plate, 2—dorsolateral stelex sensillum, 3—lateral verruciform sensillum, 4—ventrolateral stelex sensillum, 5—intermediate sensory complexes. and third instars include a change in the in- cidence of the minute acanthae as described above. Other changes include increases from three (Fig. 4C-7) to five (Figs. 6B-7; B-8; C-9; D-4, -5) in the number of lateral integumental petals, which all are foliose in the second instar, but add a central, papillate petal in the third instar. Two pairs of medial integumental petals are present in the sec- ond instar (Fig. 4C-8); whereas, three pairs occur in the third instar (Fig. 6C-7). The anterior spiracles with five papillae in the second instar (Fig. 5A) compare with three (Figs. 6E, 7A-1) or four papillae (Fig. 6F) in the third instar. The sensilla surrounding the posterior spiracular plate are the same in number as in the second instar; however, Third instar of Oxyna palpalis, continued: (A) lateral spiracular complexes, 1—anterior spiracle, 2— in the second instar the lateral sensilla are stelex (Fig. 5B-2), not verruciform, as in the third instar (Figs. 7D-3, 8C). The inter- spiracular processes on the posterior spirac- ular plate of the second instar are larger, foliose, and upright (Fig. 5D-1) compared to the small, resupinate, spiniform process- es of the third instar (Fig. 8D-2). Puparia: Reniform-ellipsoidal, light-, yellow-, or reddish-brown, rarely white (Fig. 9B), anterior end bears the invagina- tion scar and anterior thoracic spiracles; caudal segment bears posterior spiracular plates (Fig. 9C), each with three broadly el- liptical, raised rimae (Fig. 9C-1) and four, interspiracular processes (Fig. 9C-2). One hundred and five puparia averaged 3.04 + VOLUME 104, NUMBER 3 Fig. 8. interspiracular processes, (B) dorsolateral stelex sensillum, (C) lateral verruciform sensillum, (D) intermediate sensory complex, 1—stelex sensillum, 2—medusoid sensillum, 3—minute acanthae. 0.03 (range, 2.13—3.70) mm in length; 1.60 + 0.016 (range, 1.14—1.99) mm in width. DISTRIBUTION AND HOosTs Oxyna_ palpalis uniquely reproduces as an inquiline in the rosette galls of Rhopa- lomyia florella of terminal buds on branch- es of Artemisia tridentata in southern Cal- ifornia. In this capacity it also functions as a facultative predator on the larvae and pu- pae of R. florella (see below), which also renders this tephritid rare and fascinating in the annals of tephritidology. Novak et al. (1967) reported that the larvae form small, succulent, polythalamous galls on small branches of A. tridentata in Idaho, but as documented in the next section of this pa- per, this interpretation was incorrect, as O. palpalis is not a gall-former. Foote et al. (1993) mapped the distribution of O. pal- Third instar of Oxyna palpalis, continued: (A) posterior spiracular plate, 1—three rimae, 2—four palis to include California, Idaho, Nevada, Oregon, Utah, Washington, and Wyoming. Thus, like that of O. aterrima (Goeden 2001b), the distribution of O. palpalis may coincide wholly with A. tridentata sensu latu, or in part with one or more of its sub- species (Hickman 1993). This is a shrub that inhabits dry soils, valleys, slopes from 300\to: 3000-- mi im the western U:S.)1-e:, north to Washington, the North Central States and south to New Mexico (Hickman 1993). The distribution of A. tridentata also extends into southwestern Canada (Barkley 1986). BIOLOGY Egg.—In each of 31, nearly fully formed rosette galls of Rhopalomyia florella (Gag- ne 1989) already containing larvae of this cecidomyiid (Fig. 1OA), most of 51 eggs of 548 Fig. 9. anterior to left; (B) puparium, habitus, anterior to left; (C) posterior spiracular plate, 1—three rimae, 2—four interspiracular processes. Oxyna palpalis, (A) prepuparium, habitus, O. palpalis were found inserted separately, a few in small clusters, pedicel-last, in leaf bases to depths of half to all of the egg lengths (Fig. 10B). Leaves at the margins or in the centers of the galls received the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON most oviposition, but a few eggs were layed upon, but not penetrating the tissues be- tween adjacent bases of inner or outer leaves. About half of the eggs lay with their long axes parallel to the long axes of the galls (Fig. 10B); the remainder lay at angles of 5° to 45° to the long axes. Oviposition in galls was scattered throughout a stand of A. tridentata, like and sympatric with the galls of R. florella, and not confined to certain individual host-plants galled repeatedly Over successive years (Headrick and Goe- den 1998). This is similar to the pattern of the incidence of galls of O. aterrima (Goe- den 2002b). Some galls bore tephritid eggs on opposite sides that showed different de- grees of embryony, presumably indicative of oviposition by different females at dif- ferent times. An average of 3.4 = 04 (range, 1-10) eggs was found in these 31 galls. Galls containing eggs of O. palpalis (Fig. 1OA) averaged 6.3 + 0.4 (range, 4.3— 11.5) mm in length, and 4.0 + 0.2 (range, 0.6—6.8) mm in width. The linear leaves in- vesting these galls averaged 3.5 + 0.3 (range, 2.0—8.6) mm in length and 0.9 + 0.04 (range, 0.6—1.3) mm in width. Larva.—Upon eclosion, most first instars of O. palpalis tunneled into the center of the gall basad of the young, cecidomyiid larvae and leaf bases (Fig. 10C), where each first instar remained for up to 3 months within an individual, ovoidal, open, smooth-walled cell. An average of 2.3 + 0.2 (range, 1—5) first instars of O. palpalis were found in a total of 53 infested galls sampled monthly during the first 3 months following oviposition, during which time the subspheroidal (smaller) (Fig. 1OA) to hemispheroidal (larger) galls averaged 6.9 + 0.3 (range, 3.1—11.5) mm in length and 4.8 + 0.2 (range, 2.9—7.4) mm in width. The linear leaves laterally surrounding the galls averaged 4.4 + 0.2 (range, 1.4—8.6) mm in length and 0.9 + 0.03 (range, 0.6— 1.4) mm in width. Seventy-three ovoidal to spheroidal cells containing the first instars of O. palpalis averaged 0.78 + 0.09 (range, 0.28—4.6) mm in length and 0.48 + 0.18 VOLUME 104, NUMBER 3 (range, 0.21—0.99) mm in width. The size of most of these cells containing first instars slowly increased during the first stadium. Unfortunately, the number of cecidomyiid larvae was not recorded in most galls; how- ever, 14 galls containing first instars of O. palpalis also were noted to contain an av- erage of 2.2 + 0.5 (range, 1-8) larvae of R. florella. Also, it was subsequently noted that most galls of R. florella without O. pal- palis contained only a single, centrally lo- cated larva (Fig. 11D). Thus, some galls containing first instars of O. palpalis also contained dead, centrally located, cecido- mylid larva(e) presumably killed after ac- cidental contact with a tephritid larva. In- deed, at least for later instars of O. palpalis in those many galls found to be lacking R. florella (see below), this relationship may represent nutritionally advantageous, facul- tative predation. The larva next molted to the second in- star (Fig. 10D), evidenced as the cast ce- phalopharyngeal skeleton remaining in the cell. This instar may also last about 3 months and is one stage found in overwin- tering galls in southern California (see be- low). The external dimensions of 28 galls found to contain second instars measured 6.8 + 0.3 (range, 3.7—11.4) mm in length by 5.7 + 0.3 (range, 2.9-8.6) mm in width. The linear leaves investing these galls av- eraged 4.6 += 0.3 (range, 1.7—9.1) mm in length and 1.1 + 0.04 (range. 0.7—1.7) mm in width. These 28 galls each contained an average of 1.8 + 0.2 (range, 1—6) second instars of O. palpalis (Fig. 10D). Two of these galls (7%) contained two and three larvae clustered together in a central, com- mon, open cavity. The remaining 26 galls contained one or more second instars in separate, open, ovoidal cells (Fig. 10D) that averaged 1.3 + 0.06 (range, 0.6—2.0) mm in length and 0.8 + 0.04 (range, 0.4—1.1) mm in width. Thus, the galls grew little on average during the second stadium, as did the cells containing individual second in- stars of O. palpalis (Fig. 10D). Ten galls were recorded to contain dead larvae of R. 549 florella or none, the latter of which was pre- sumed to reflect complete consumption by O. palpalis, but again, cecidomyid inci- dence was not recorded in another 10 of these galls, so the incidence of cecidomyiid mortality may actually have been higher. Oxyna palpalis also overwinters as third instars in southern California (Figs. 1OE, F). This is the stage during which the greatest amount of larval growth and attendant in- crease in gall cavity size takes place (Figs. 1OE, F). Seventy-eight galls containing third instars of O. palpalis averaged 8.9 + 0.2 (range, 5.1—-13) mm in length by 7.4 + 0.2 (range, 2.1—4.5) mm. The linear leaves laterally surrounding the galls averaged 5.3 + 0.2 (range, 2.2—10.8) mm in length and 1.2 + 0.03 (range, 0.9-1.7) mm in width. Thus, on average, galls increased little, if any, in size during the equally slow growth of the third instars during the winter through early spring, when in response to renewed plant growth following winter rainfall, the cavities that contained the third instars expanded to accomodate the fast- growing, O. palpalis larvae. The 65, frass- lined, ovoidal or irregularly shaped, cen- trally located, open cells that contained sin- gle instars of O. palpalis within these 78 galls averaged, 2:6 = O31 @ange; 1yl=5-1) mm in longest measurement by 1.7 + 0.1 (range, 0.57—2.9) mm in shortest measure- ment, or about twice as large as cells con- taining single second instars. Moreover, in galls containing two or more third instars (Figs. 10E, F) these central cavities aver- aged 3.3 + 0.2 (range, 1.7—5.1) mm in lon- gest length by 2.6 + 0.2 (range, 0.85—4.6) mm in shortest width, or, again, more than twice as large on average than cavities con- taining single second instars, or those with single third instars, sometimes occupying much of the interior of smaller galls. All told, the 78 galls contained an average of 1.7 + 0.1 (range, 1—5) third instars (Figs. 1OE, F). The lateral thickness of the walls of these galls averaged 1.8 + 0.6 (range, 1.1—2.9) mm, which afforded insufficient spatial protection from parasitoids (see be- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 10. Life stages of Oxyna palpalis in galls of Rhopalomyia florella: (A) partly grown gall of R. florella that contained egg of O. palpalis, (B) egg of O. palpalis (arrow) in gall of R. florella, (C) newly eclosed, first instar of O. palpalis (arrow) tunneling into center of gall, (D) two, overwintering second instars of O. palpalis in separate chambers (arrows), (E) four third instars of O. palpalis in common, central cell in gall, (F) 1—newly formed puparium and 2—full-sized larva of O. palpalis, and 3—pupa of R. florella in same gall. Lines = 1 mm. low) that attacked the third instars and pu- paria of O. palpalis. All told, third instars were found in samples over an 8-month pe- riod, 6 months into which, puparia first ap- peared in monthly gall samples. Pupa.—Near the end of the third larval stadium, the third instar transformed into a preparium (Fig. 9A), which in O. palpalis was of relatively short duration. No window to facilitate future adult egress from the galls was formed in the gall wall, unlike the windows made by O. aterrima third instars prior to prepupal formation (Goeden 2002b). The prepuparia of O. palpalis (Figs. 9A, 10F-2) transformed into puparia within the larval cells or among the cottony tomentum (Fig. 1OF-1), which by then also contained pupae (Fig. 1OF-3) or empty pu- pal exuviae of emerged, surviving R. flo- rella, usually found on the periphery of a VOLUME 104, NUMBER 3 51 Fig. 11. gall (Fig. 1OF-3). The anterior end of the puparium of O. palpalis usually faced dis- tad, away from the base of the gall (Figs. 10F-1, 11). Eighty-four mature galls (Figs. 11B, C) sampled over a 2-mo period con- tained an average of 1.5 + 0.1 (range, 1—5) puparia of O. palpalis. These galls exter- nally averaged 9.1 + 0.2 (range, 5.7—15.0) mm in length by 7.9 + 0.2 (range, 5.0- 11.4) mm in width. The linear leaves lat- erally surrounding the galls averaged 7.1 + 0.3 (range, 2.6—-16) mm in length and 1.4 Life stages of Oxyna palpalis in galls of Rhopalomyia florella, continued, (A) two puparia of O. palpalis in gall, (B) full-size gall that contained puparia of O. palpalis, (C) single central puparium of O. palpalis in gall, (D) single, central pupa of R. florella in gall, (E) male adult of O. palpalis, (F) female adult of O. palpalis. Lines = 1 mm. + 0.03 (range, 0.8—2.5) mm in width. Sixty (71%) of the 84 galls each contained a frass-lined, ovoidal or irregularly shaped, central cavity that held a single puparium of O. palpalis (Fig. 11C), which averaged 3.4 + 0.1 (range, 2.3—6.3) mm in longest measurement by 2.0 + 0.1 (range, 1.1—3.4) mm in shortest measurement. Whereas, in galls containing two or more puparia (Fig. 11A) these central cavities averaged 4.1 + 0.2 (range, 2.9—-5.7) mm in longest length by 3.2 + 0.1 (range, 2.3—4.6) mm in short- S52 Table 1. Incidence of Oxyna palpalis in galls of Rhopalomyia florella on Artemisia tridentata on sam- ple dates indicated. R. florella Galls Sampled No. Without No. With Sample Date O. palpalis (%) O. palpalis (%) 10.iv.1996 49 (84) 9 (16) l.v. 1996 51 (84) 10 (16) 7.v. 1996 55 (79) LES (2119) 5.vi. 1996 u ils) (2) 18.vi. 1996 12 (38) 20 (62) 16.vi1. 1996 15 (56) 12 (44) 13.vili. 1996 p25) (7/\\)) 10 (29) 16.1x. 1996 76 (87) MGS) 16.x. 1996 35 (81) 8 (19) 13.x1.1996 60 (79) 16 (21) 18.x11. 1996 40 (70) 17 (30) Lone 997 20 (71) 8 (29) 12997 34 (63) 20 (37) 1S 997 65 (81) Pas) (UL) Sits IL 76 (76) 24 (24) Guan 997 69 (78) 19 (22) 30.iv. 1997 45 (80) 11 (20) est width, again, sometimes occupying much of the interior of smaller galls. The central location of a single pupa of R. flo- rella in a gall without O. palpalis (Fig. 11D), when compared to a similarly located puparium of O. palpalis typical of the 60 galls containing single individuals noted above (Fig. 11C), most of which lacked any sign of R. florella, strongly suggests a high incidence of facultative, or at least acciden- tal predation by the tephritid on the ceci- domyiid. Table | records the incidence of O. palpalis in galls of R. florella on differ- ent sample dates. Adult.—Adults exited galls presumably by pushing aside, between, and outward through the enfolding, apical, immature leaves and tomentum (Figs. 11A, C). Under insectary conditions, 17 males (Fig. 11E) lived an average of 48 + 5 (range, 21-84) days, and 12 females (Fig. 11F) an average of 41 + 6 (range, 12-75) days. These lon- gevities are relatively long in duration for a circumnatal tephritid (Headrick and Goeden 1998). For example, males and females of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON O. aterrima on average respectively lived half as long (Goeden 2002b). Mating behavior.—The premating, mat- ing, and postmating behaviors of O. pal- palis were not studied in the field, nor were these behaviors observed in petri dish are- nas of the type found to be so useful with many other nonfrugivorous, tephritid spe- cies (Headrick and Goeden 1994, Goeden 2002b). In two such arenas, adults exhibited a few behaviors similar to O. aterrima (Goeden 2002b) and typical of other cir- cumnatal, gallicolous species, cf., Proceci- dochares, previously studied in southern California, i.e., a lack of courtship behavior, the exhibition of enantion type wing move- ments by both sexes, and male stalking of females prior to mating (Goeden and Norr- bom 2001; Headrick and Goeden 1994, 2000). Seasonal history.—Oxyna palpalis is a univoltine, circumnatal species (Headrick and Goeden 1994, 1998) reproducing in ro- sette galls of Rhopalomyia florella of ter- minal buds on branches of A. tridentata in southern California (Gagné 1989). Eggs are laid in nearly fully formed galls containing young larvae of R. florella in early summer (June) and O. palpalis passes the summer (June-September) as first instars. These lar- vae molt to the second instar beginning in mid-September in early fall, and to the third instar beginning in mid-October, initially overwintering as both second and third in- stars, but solely as third instars by mid-win- ter (February) of the following year. The third instars complete this instar beginning in early spring (March) at the time of the renewed flush of host-plant growth. Pupar- iation and adult emergence follow in April and the adults mate and probably oviposit in newly formed galls of R. florella on or near the same plants from which both spe- cies of flies emerged. Natural enemies.—Many individual Eur- ytoma sp. (Hymenoptera: Eurytomidae) and six individuals of Eupelmus sp. (Hymenop- tera: Eupelmidae) were reared separately from individual puparia as primary, soli- VOLUME 104, NUMBER 3 tary, probably larval-pupal endoparasitoids. One, three, three, and seven individuals of Lyrcus sp. (Hymenoptera: Pteromalidae) were reared from individual puparia of O. palpalis as gregarious, primary endoparas- itoids. ACKNOWLEDGMENTS My thanks to Andrew C. Sanders, Cu- rator of the Herbarium, Department of Bot- any and Plant Sciences, University of Cal- ifornia, Riverside, for identifications of plants mentioned in this paper. Krassimer Bozhilov in the Central Facility for Ad- vanced Microscopy and Microanalysis, University of California, Riverside, greatly facilitated my scanning electron microsco- py and digital imaging. The parasitoids ini- tially were identified by Harry E. Andersen, Huntington Beach, California, now de- ceased, and later were confirmed or further identified by John Heraty and Jung-Wook Kim, Department of Entomology, Univer- sity of California, Riverside. I also am grateful to Jeff Teerink for technical assis- tance and to David Headrick for his helpful comments on earlier drafts of this paper. LITERATURE CITED Baker, C. EF 1904. Reports on California and Nevadan Diptera, |. Invertebrata Pacifica 1: 17—39. Barkley, T. M., ed. 1986. Flora of the Great Plains. University Press of Kansas, Lawrence. Blanc, F L. and R. H. Foote. 1963. The fruit flies or Tephritidae of California. Bulletin of the Califor- nia Insect Survey 7: 1-117. Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc. Portland, Oregon. Foote, R. H. and F L. Blanc. 1963. The fruit flies or Tephritidae of California. Bulletin of the Califor- nia Insect Survey 7: 1-117. 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. Gagné, R. J. 1989. The Plant-feeding Gall Midges of North America. Comstock Publishing Associates, Cornell University Press, Ithaca and London. Si nn ee) Goeden, R. D. 2001. Life history and description of immature stages of Neaspilota footei Freidberg and Mathis (Diptera: Tephritidae) on Aster occi- dentalis (Nuttall) (Asteraceae) in southern Cali- fornia. Proceedings of the Entomological Society of Washington 103: 191—206. . 2002a. Description of the immature stages of Tephritis stigmatica (Coquillett) (Diptera: Tephri- tidae). Proceedings of the Entomological Society of Washington 104: 335-347. . 2002b. Life history and description of im- mature stages of Oxyna aterrima (Doane) (Dip- tera: Tephritidae) on Artemisia tridentata Nuttall (Asteraceae) in southern California. Proceedings of the Entomological Society of Washington 104: 510-526. Goeden, R. D. and A. L. Norrbom. 2001. Life history and description of adults and immature stages of Procecidochares blanci, n. sp. (Diptera: Tephriti- dae) on /socoma acradenia (E. Greene) E. Greene (Asteraceae) in southern California. Proceedings of the Entomological Society of Washington 103: 517-540. Goeden, R. D. and J. A. Teerink 1999. Life history and description of immature stages of Trupanea vicina (Wulp) (Diptera: Tephritidae) on wild and cultivated Asteraceae in southern California. Pro- ceedings of the Entomological Society of Wash- ington 101: 742-755. 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— 22, ——.. 1998. The biology of nonfrugivous tephritid fruit flies. Annual Review of Entomology 43: 217-241. . 2000. Behavior of flies in the subfamily Te- phritinae. Chapter 2. Jn Aluja, M. and A. L. Norr- bom, eds. Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, London, New York, and Washington, D.C. Hickman, J. C., ed. 1993. The Jepson Manual. Uni- versity of California Press. Berkeley and Los An- geles. Novak, J. A., W. B. Stolzfus, E. J. Allen, and B. A. Foote. 1967. New host records for North Ameri- can fruit flies (Diptera: Tephritidae). Proceedings of Entomological Society of Washington 69: 146— 148. Quisenberry, B. F 1949. The genus Oxyna in the Ne- arctic Region north of Mexico. Pan-Pacific Ento- mologist 25: 71-76. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 554-562 BIOLOGICAL OBSERVATIONS OF CENTISTES GASSENI SHAW (HYMENOPTERA: BRACONIDAE), A PARASITOID OF DIABROTICA SPP. (COLEOPTERA: CHRYSOMELIDAE)! R. EF W. SCHRODER AND M. M. ATHANAS (RFWS) Research Entomologist, retired, (MMA) Entomologist, Insect Biocontrol Lab- oratory, U.S. Department of Agriculture, Agricultural Research Service, Bldg. 011A, Rm. 107, BARC-West, Beltsville, MD 20705, U.S.A. (e-mail: athanasm @ba.ars.usda.gov) Abstract.—In 1992-1993, Centistes gasseni Shaw was imported into the United States, and in the laboratory successfully parasitized: southern corn rootworm, Diabrotica un- decimpunctata howardi Barber; banded cucumber beetle, D. balteata LeConte; western corn rootworm, D. virgifera virgifera LeConte; and striped cucumber beetle, Acalymma vittatum (FE). Males and females of C. gasseni lived an average of 15.4 and 12.9 days (with a maximum of 30 and 29 days), respectively. A single female oviposited in 383 host Diabrotica over her lifetime, from which 158 cocoons were recovered. Additional observations on the biology and rearing of the parasitoid are presented. Key Words: Acalymma vittatum, biological control, Centistes gasseni, Diabrotica bal- teata, Diabrotica barberi, Diabrotica speciosa, Diabrotica undecimpunc- tata howardi, Diabrotica virgifera virgifera, host specificity The northern corn rootworm (NCR), Diabrotica barberi Smith and Lawrence, and the western corn rootworm (WCR), Diabrotica virgifera virgifera LeConte are the most damaging and costly pests of corn in North America (Sutter and Lance 1991). To achieve effective control of these chrys- omelids, soil insecticides routinely have been used on 50—60% of the corn acreage, or 12-16 million ha annually (Metcalf 1986). These insecticides are generally ap- plied prophylactically and are frequently unnecessary (Lance and Sutter 1992). Ac- cording to Metcalf (1986), the cost from crop loss and treatment due to the corn rootworm (CRW) complex is approximate- ly one billion dollars per year. 'This article reports the results of research only. Mention of a proprietary product does not constitute an endorsement or a recommendation by the USDA for its use. In addition, southern corn rootworm (SCR), Diabrotica undecimpunctata ho- wardi Barber; striped cucumber beetle, Acalymma vittatum (FE); western striped cu- cumber beetle, Acalymma trivittatum (Man- nerheim); and banded cucumber beetle, Diabrotica balteata LeConte cause 50—100 million dollars in damage to other crops (Cucurbitaceae, Fabaceae) (Metcalf et al. 1962). Lance and Sutter (1992) cite health risks to growers, livestock and wildlife due to soil insecticides used for CRW _ control. They also refer to reports of these insecti- cides being detected in ground and surface water. Therefore, it is evident that there is a need for alternative control measures that are environmentally safe and cost effective. In 1990, while searching for biocontrol agents of Colorado potato beetle (Coleop- tera: Chrysomelidae) in Rio Grande do Sul, VOLUME 104, NUMBER 3 Brazil, we learned of the existence of an undescribed braconid parasitoid of the adult Neotropical leaf beetle, Diabrotica speciosa Germar (Gassen 1986). It was subsequently described by Shaw (1995) as Centistes gas- seni Shaw, a solitary endoparasitoid of the adult. As there are no known effective par- asitoids of the CRW complex in the United States, we were interested in acquiring and studying C. gasseni as a potential new bi- ological control agent. We traveled to Bra- zil in 1992—93, and, in cooperation with Dirceu Gassen, EMBRAPA-CNPT, collect- ed and imported adults of D. speciosa par- asitized by the braconid. In this article, we present observations on the biology, rear- ing, mating behavior and host specificity of C. gasseni, obtained during initial attempts to colonize the parasitoid in quarantine. MATERIALS AND METHODS Approximately 6,000 D. speciosa adults were collected from April 24—26, 1993, at the University of Passo Fundo, Passo Fun- do, Rio Grande do Sul, Brazil. Collections were made primarily from potato plants, but some adults were also collected from near- by plots of pepper and sweet potato plants. Parasitoids were also recovered from bee- tles collected from flowering weeds and wild cucurbit plants, 98 km west of Passo Fundo. Sub-samples of field collected bee- tles were dissected to determine percent parasitism and the number of parasitoids per host. In subsequent collections, Hei- neck-Leonel and Salles (1997) recovered C. gasseni from D. speciosa collected in corn, beans, melon, cucumber, cabbage, broccoli, spinach and lettuce. Field collected adult beetles were held in 30.5 cm collapsible aluminum screen cages (BioQuip® Products, Gardena, CA). Each cage was inverted and the hinged top served as the bottom. A fitted piece of moistened felt was placed on the bottom as a liner, and covered with a ca 0.5—0.6 cm layer of moistened sand with a 2.5 cm sand- free margin on all sides. The sand and felt served as a substrate in which the parasitoid Si Nn nN larvae spun cocoons. Approximately one cm above the bottom of the cage, we fitted a pre-cut piece of number 12 mesh stainless steel screen, that separated the adult Dia- brotica from emerging C. gasseni larvae, which passed through to pupate. The adult beetles, and 31 parasitoid co- coons recovered prior to departure, were taken to the Maryland Department of Ag- riculture quarantine facility, Annapolis, MD, and held at 21.1°C, 60% RH, and a 16:8 LD photoperiod. The host beetles were kept in the modified cages which were ser- viced three times a week. Parasitoid co- coons were carefully removed from the sand and placed in petri dishes, along with a moistened, 3.8 cm cotton roll to maintain humidity. The containers used for oviposition, and to house the C. gasseni adults, were plex- iglass tubes (7.6 cm in diameter and 10.2 cm long) modeled after those used by Whistlecraft et al. (1984) in their braconid rearing program. To confirm mating, a sin- gle female was confined in an aspirator tube with multiple males until copulation was observed. With Cotesia melanoscela (Ratz- eburg), Weseloh (1977) found that 20 sec- onds was necessary to successfully transfer sperm to the female. We therefore consid- ered a successful mating session as one in which copulation was observed for 20 sec- onds or more. We used several of the meth- ods discussed by Singh (1982) to induce successful mating, including shaking the in- sects to the bottom of the tube, increasing the number of males per female, and peri- odically chilling the females prior to ex- posure to the males. The following procedures were used in quarantine to conduct studies on the biolo- gy, mating behavior, and host preferences of the parasitoid. Host exposure methods.—I/ndividual ex- posure: To expose host Diabrotica adults individually, we placed a parasitoid female into a 35 X 10 mm polystyrene culture dish and introduced a single Diabrotica adult. The host beetle was observed until ovipo- 556 sition occurred, then it was removed. Ten to twenty Diabrotica were exposed to a fe- male per session. Following each session, the female was held for one hour in a tube or petri dish, and given access to water and honey solution. Each female had no more than two exposure sessions per day. A sep- arate cage (#1—3) was set up for the Dia- brotica exposed to each of the three, mated C. gasseni females used. Group exposure with immediate remov- al: Approximately 35 Diabrotica beetles were placed in an oviposition tube, into which a C. gasseni female was then intro- duced. As soon as a host beetle was para- sitized, it was aspirated out of the tube. As above, following each session females were permitted to rest, and limited to two ses- sions per day. A separate cage (#4—6) was set up for the Diabrotica exposed to each of the three, mated C. gasseni females used. Group exposure for one hour: Six C. gasseni females were used to sting a total of 394 BCB over a 7-day span. The females were confirmed mated, and all had emerged in association with other males and females. Beetles were exposed in groups of 15—20 to individual females for one hour, then placed into a single cage (#7). Group exposure for four hours: Six C. gasseni females, selected from a group of females, were used to sting a total of 140 BCB over a 4-day span. All the females had been confined with an equal number of males (17 6 & 17 2) for 72 hours preced- ing exposure to Diabrotica, but mating was not confirmed. As above, all had emerged in association with other parasitoid males and females. BCB were exposed in groups of 20—25 to single C. gasseni females, for four hours duration, then placed into a sin- gle cage (#8). Host species preference.—Adult SCR and BCB were sexed, and three males and three females of each species were placed into an oviposition tube. A C. gasseni fe- male reared from BCB host was introduced and exposed to the beetles for one hour, af- ter which time she was removed and held PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON as previously described. Using nine C. gas- seni females, a total of 792 Diabrotica were exposed. Host sex preference.—To determine if C. gasseni exhibited a preference for either male or female hosts, SCR reared parasit- oids (n = 10) were exposed individually to six male and six female SCR for one hour. One group of hosts (204 2 & 6, cages | & 2, respectively) was exposed to C. gas- seni females 1—3 days old, another group of hosts (372 2 & 6, cages 3 & 4, respec- tively) was exposed to C. gasseni females, the majority of which were I—5 days old. Two cages of unexposed male and female Diabrotica were maintained as controls. Host specificity—Host specificity tests were conducted to determine whether SCR, WCR, BCB, the related SCB and Cerotoma trifurcata (FE) were suitable hosts for C. gasseni. The criteria for these tests were based on observing the parasitoids’ ovipo- sitional behavior for one hour after expo- sure to the hosts, and then holding parasit- oid and hosts together (25—50 per exposure) over night. The female was then removed and the hosts held for parasitoid emergence for 30 days. Additional non-target insects that may occur in the same habitat, including bene- ficial and phytophagous coccinellids were also exposed to C. gasseni to determine their suitability as hosts. In each test, a sin- gle C. gasseni female, less than seven days old, was placed in an oviposition tube. Five Diabrotica (BCB or SCR), which served as controls, and 25—50 adult host beetles (hosts were obtained from laboratory and field sources) were then introduced. Each tube was observed for one hour to confirm any oviposition that occurred. The Diabro- tica were removed after they were stung and set up in separate parasitoid emergence containers. After one hour, observations were discontinued, and females were kept with the test beetles for 24 hours. Host bee- tles were then removed, and held for para- sitoid emergence. Thirty days after expo- VOLUME 104, NUMBER 3 Mable: 41. speciosa. nN nN ~~ Emergence, sex ratios and hyperparasitism of Centistes gasseni from field collected Diabrotica Sex Ratio No %o Centistes Date No. Cocoons No No ; Mesochorous Emergence 4/92 1LO8 1] 19 1:2 12 28 4/93 Bil 5 6 {<1 2 és) 5/93 338 138 64 Zo) 3 60 5/94 LO] 10 13 1h 2) 23 6/944 rele) 8 8 Le] 14 19 “Data from cocoons collected at the ARS, South American Biological Control Laboratory, Buenos Aires, Argentina. sure, host beetles were dissected to deter- mine if they had been parasitized. RESULTS From the 31 cocoons brought back from Brazil, 11 C. gasseni and two hyperparasi- toids, Mesochorous sp. (Hymenoptera: Ich- neumonidae) emerged. From the ~ 6000 D. speciosa collected in Brazil, we recovered 338 C. gasseni cocoons, 66 C. gasseni lar- vae, and 11 puparia of Celatoria bosqui Blanchard (Diptera: Tachinidae) (Table 1). As the number of host beetles collected was approximated, we could not accurately de- termine the percent parasitism. However, in one sub-sample (n = 111) 10.0% were par- asitized. In Pelotas, RS, Brazil, Heineck- Leonel and Salles (1997) reported that par- asitism, of D. speciosa by C. gasseni, ranged from 0-18.9% (* = 8.7, SD = 7.1) in samples obtained at monthly intervals from May 1994—April 1995. Mating data were obtained on a sub-sam- ple of P and F generation parasitoid fe- males. Forty-one percent of the P sample (n 17) successfully mated (Table 2). All C. gasseni pupae obtained from D. speciosa were set up individually, thus all females that emerged were isolated from other males and females. Of those that mated, 29% did so on the first attempt, 43% on the second, and 28% required three or more at- tempts to mate. Unsuccessful mating at- tempts were observed for a minimum of 10 minutes. Seventy-one percent of the F sam- ple (n = 90) successfully mated (Table 2). Of these, 72% occurred on the first attempt, 20% on the second, and 8% required three or more attempts. In the majority of cases, the female would aggressively fight off the advances from any males after she had suc- cessfully mated. Nevertheless, 9% of the fe- males did mate a second time, all within 2.1 minutes of the first. When we exposed pre- viously mated females to new males, 8% mated a second time. Parasitoid female progeny were obtained only from positively mated parental fe- males. When confined with a suitable host, the female mounted the beetle from the side and thrust her ovipositor into the abdomen. The beetle responded to the attack by trying Table 2. Reproductive behavior of Centistes gasseni reared from native and alternative host Diabrotica. %1so2 % % & % 2 with Mean % Mean ? with & with d 2 &o Mean Expos. Mean Length Mean No. Mean Age 6 Host 2 Mated Age (Days) Mated Mated Mated Mated (Min) (Sec) 3 Used (Days) Pgen> 41 Dalia lesry 1) N/A N/A N/A 32852) S20) 93102428 eee 4S Fa °4 BESS Al LES == OLO88 87:8 86 68 57 Spe ee EI BY PS) series "ete 22 I 2.4 + 0.4 “Tsolated from other females and males. °C. gasseni reared from D. speciosa. °C. gasseni reared from North American Diabrotica. 558 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Parasitism by Centistes gasseni using various host exposure methods. Days ? ? Emerged Hosts No. %e No. 3 No. @ % Cage in Use with 3, 9 Exposed Cocoons Parasitism Emerged Emerged Emerged 1 1—5 None 103 SCR? 62 60 q 39 74 2, 1-2 296 12 61 SCR 45 80 Si 0) 82 3p 1=3 296 12 47 SCR 28 60 14 1 54 4 2-13 aXe) 3) 232 SCR 154 73} 102 0) 66 5 5-9 36 32 129 BCBs 61 62 D1 0) 93 6° 2-12 le 383 BCB 158 52 93 0) 59 7 1-9 — 394 BCB 163 52 68 14 50 8 4—6 — 140 BCB 67 64 36 5) 61 “Cages 1-3, hosts exposed individually, cages 4—6, hosts exposed in groups with immediate removal, cage 7 hosts exposed for one hour, cage 8 hosts exposed for four hours. > Female did not mate until 37 of the 47 hosts had been exposed. © Female mated twice. 4 Southern corn rootworm, Diabrotica undecimpunctata howardi Barber. © Banded cucumber beetle, D. balteata LeConte. to brush the C. gasseni from its back, be- coming highly active or agitated, or by dropping to the bottom of the container. Dissections of field collected D. speciosa revealed that parasitized beetles contained a single C. gasseni larva per host. However, laboratory dissections indicated that super- parasitism did occur, with only one larva completing development. At 21.1 + 0.5°C, the parasitoid would complete its life cycle in 33—41 days (x = 34.7 + 1.7 days). At 23.9 + 0.5°C, it took 26-32 days (* = 27.8 10.7, days): At 21.) =O "Ce sasseni females and males lived an average of 12.9 25/0 days and 15/4 =" 2.8 "days; respec- tively. Host exposure methods.—I/ndividual ex- posure: In the hosts exposed individually (cages 1-3), the mean percent parasitism, based upon the number of hosts exposed and the number of parasitoids (cocoons and/or larvae) recovered, was 66.7% (SD = 11.54), and the mean parasitoid emergence from the resultant cocoons was 70% (SD = 14.4) (Table 3). At 32 + 5 days post ex- posure, 89.3% (SD = 5.03) of the host SCR were dead. The female of cage 1, which emerged alone, produced offspring with a sex ratio of 6 2:1 d, the highest ratio of females to males obtained in this study. Group exposure with immediate remov- al: The females of cages 4 and 6 oviposited in 232 SCR and 383 BCB respectively, over a 12 day period (Table 3). These yield- ed 154 and 158 cocoons, respectively, and 102 males and 93 males, respectively. This was the largest number of hosts stung, and number of cocoons and progeny obtained from individual C. gasseni females. Pre-cocoon mortality was higher in the BCB hosts, averaging 22.0% versus 8.5% in the SCR. Eliminating cage 6 from the computations, since this female was ex- posed to an excessive number of hosts to determine oviposition limits, percent para- sitism using this method of exposure aver- aged 67.5% (SD = 7.78). The mean para- sitoid emergence from the cocoons was 72.6% (SD = 18.0), and at 29 + 4 days post exposure, 91.5% (SD = 3.5) of the ex- posed hosts were dead. Parasitoid females 10—12 days old, were the oldest from which viable progeny were recovered. The oldest age attained by a C. gasseni male and female, was 30 and 29 days, respectively. Over the course of the female’s life, she was exposed to 114 SCR, both in groups and individually. As she aged, all hosts were exposed individually. On her 24th day, she stung two SCR, this was the oldest age at which we observed attack behavior. A cocoon was recovered from a SCR parasitized by a female 14—24 days old. There was no emergence from the VOLUME 104, NUMBER 3 Table 4. Host specificity of Centistes gasseni. Para. Emerge from Host Host Species Tested Para. Dissected from Host Para. Emerge from Control Coccinellidae Coleomegilla maculata (DeGeer) Hippodamia variegata (Goeze) Propylea quatuordecimpunctata (L.) Epilachna varivestis Mulsant Cycloneda munda (Say) Chilocorus stigma (Say) Hippodamia convergens Guerin Harmonia axyridis (Pallas) Chrysomelidae Cerotoma trifurcata (F.) Acalymma vittatum (FP.)* Diabrotica balteata LeConte* D. undecimpunctata howardi Barber* D. virgifera virgifera LeConte* | +++4¢4+4+ 44 ++ ++ “The Acalymma and Diabrotica tested were suitable hosts, dissections and controls were therefore not used. cocoon, and a male was subsequently dis- sected out. This proved to be the oldest C. gasseni from which both cocoon, and fully formed offspring have been recovered. Group exposure for one and four hours: The percent parasitism obtained by expo- sure for one (cage 7) and four (cage 8) hours, was 52% and 64%, respectively (Ta- ble 3). From cage 7, 82 parasitoids with a sex ratio of 5 d:1 2, were recovered. From cage 8, we recovered 41 parasitoids with a 7 3:1 2 sex ratio. This was the first in- stance in which parental females that were not confirmed mated, produced female off- spring. Pre-cocoon mortality averaged 23% for cages 7 and 8. Parasitoid emergence from the cocoons was 50% and 61% for cages 7 and 8, respectively. Host species preference.—Although all parental C. gasseni used were BCB reared, they did not exhibit a preference for BCB hosts. The percentage parasitism averaged 32% in both the BCB and SCR. Fifty-seven parasitoids, including seven females, were recovered from SCR hosts, and 55 parasit- oids, including three females, were recov- ered from BCB hosts. All C. gasseni female offspring were recovered from BCB fe- males and SCR males. Pre-cocoon mortal- ity was 14% for both SCR and BCB. Par- asitoid emergence from cocoons was 51% for BCB males and females, 65% for SCR males and 42% for SCR females. Host sex preference.—In cages | and 2, the percent parasitism for female and male hosts was 32% and 23%, respectively. In cages 3 and 4, it was 45% and 38% for the female and male hosts, respectively. Emer- gence averaged 70% from the cocoons ob- tained from female hosts, and 78% from co- coons obtained from the males. For both C. gasseni age groups, all female offspring were recovered from female SCR hosts. Beetle mortality appeared to be excessive with this generation: pre-cocoon mortality averaged 46% for cages | and 2, and 53% and 60% for cages 3 and 4, respectively. Thus it is uncertain whether the recovery of female parasitoids from female hosts was due to parasitoid preference, or selective mortality. Host specificity—The SCR, WCR, BCB and SCB exposed to C. gasseni were suc- cessfully parasitized, resulting in the recov- ery of parasitoids from all four hosts. Nei- ther Cerotoma trifurcata (FE), nor any of the exposed coccinellids were parasitized (Ta- ble 4). In every test, given the choice be- 560 tween the control Diabrotica and the test insect, C. gasseni always searched for, and attacked the Diabrotica first, indicating that the parasitoid was active. The time it took to parasitize all control Diabrotica varied from 10-30 minutes. C. gasseni females approached the test beetles numerous times during the observation period. Usually, they would rapidly tap their antennae on the bee- tle, then move on to another host. Occa- sionally the female would mount the test beetle and probe with the tip of the abdo- men at the same location where she would normally oviposit. DISCUSSION The survival of C. gasseni larvae found during the maintenance of the host cages was extremely low, regardless of whether they were returned to the original cage or set up in petri dishes containing moistened sand. Vance (1932) observed that any Che- lonus annulipes Wesmael larvae that emerge from the host and completes its fi- nal feeding, is unable to construct a cocoon if it is disturbed in any way. Harrison et al. (1993) observed a similar situation with lar- vae of Microplitis croceipes (Cresson). Those larvae died within two hours of ex- iting the host if they had not spun cocoons by then. When inducing C. gasseni to mate, most of the successful copulations occurred when modified, open-ended aspirator tubes were used, and an air current provided. This seemed to aid the males in orienting to the virgin females (Vinson 1978). This suggests the presence of a sex pheromone in C. gas- sent (Matthews 1974). In several cases in which females did not mate, or mating oc- curred after multiple attempts, the males used were =24 hours old. Laing and Cal- tagirone (1969) observed that females of the braconid Habrobracon lineatellae (Fi- scher) will not mate until they are at least 24 hours old, and that insemination is more likely to occur when older males (5—6 days old) are used. As Schlinger and Hall (1960) reported that the sperm supply drops rap- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON idly with each successful mating, whenever possible, we removed each male after it had mated. The host beetle apparently dies soon after parasitoid emergence; when harvesting C. gasseni cocoons we would find almost an equal number of dead beetles. Loan and Holdaway (1961) observed that the curcu- lionid, Sitona sp., would stop laying eggs 1—2 days after being stung, and usually die in 3—4 hours following emergence of the endoparasitic braconid larva, Pygostolus falcatus (Nees). No female parasitoid prog- eny were obtained using group exposure of hosts. The large host to parasitoid ratio (Grinberg and Wallner 1991), and rapid rate of oviposition (Flanders 1956) may have been factors. When exposed to hosts in groups, the females would oviposit at a rate of one host per 0.92 + 0.38 min. When presented with hosts individually, the rate slowed to one host per 2.2 + 0.78 min. Centistes gasseni was successfully reared through nine generations. During this peri- od, the sex ratio varied widely, ranging from all males to 6 2:1 3d. An adequate number of females was obtained to conduct experiments and propagate the colony. By the tenth generation, only males were pro- duced. Although our results showed that fe- males contacting each other did not pre- clude the production of female offspring, it may have affected the sex ratio. With the scelionid Trissolcus, contact with other fe- males of the same species (Viktorov 1968), or trace pheromones of other females (Vik- torov and Kochetova 1973, Buleza 1975), resulted in production of significantly more male offspring. In addition, the fluorescent lighting utilized in quarantine may have hindered successful mating. Nealis and Fra- ser (1988), observed that the braconid Apanteles fumiferanae (Viereck), mated more frequently under natural rather than artificial light conditions. The SCR, WCR, BCB, and SCB were readily attacked and parasitized by C. gas- seni, Whereas other phytophagous and ben- eficial beetles tested proved unsuitable as VOLUME 104, NUMBER 3 hosts. Due to the small number of C. gas- seni females and availability of host beetles, the specificity tests were limited in scope, but generic specificity was evident. Centistes gasseni has potential for use as a biocontrol agent of the North American CRW complex, and the striped cucumber beetle, A. vittatum. The biology, population dynamics and behavior of C. gasseni needs to be studied on its native host D. speciosa in Brazil. ACKNOWLEDGMENTS We gratefully acknowledge the assistance and cooperation of: Ann Sidor, USDA, In- sect Biocontrol Laboratory, Beltsville, Maryland; Guillermo Cabrera Walsh, USDA, Agricultural Research Service, South American Biological Control Labo- ratory, Buenos Aires, Argentina; Dirceu Gassen, EMBRAPA-CNPT, Passo Fundo, RS, Brazil; the Maryland Department of Agriculture, Annapolis, Maryland; and the USDA Vegetable Research Laboratory, Charleston, South Carolina. LITERATURE CITED Buleza, V. V. 1975. 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A. and L. A. B. Salles. 1997. In- cidencia de parasitoides e patoge em adultos de Diabrotica speciosa (Germar) (Coleoptera: Chry- S561 somelidae) na regiao de Pelotas, RS. Anais da So- ciedade Entomologia do Brasil 26: 81-85. Laing, D. R. and L. E. Caltagirone. 1969. Biology of Habrobracon lineatellae (Hymenoptera: Bracont- dae). Canadian Entomologist 101: 135-142. Lance, D. R. and G. R. Sutter. 1992. Field tests of a semiochemical-based toxic bait for suppression of corn rootworm beetles (Coleoptera: Chrysomeli- dae). Journal of 967-973. Loan, C. Economic Entomology 85(3): and E G. Holdaway. 1961. Pygostolus fal- catus (Nees) (Hymenoptera: Braconidae), a para- site of Sitona species (Coleoptera: Curculionidae). Bulletin of Entomological Research 52: 473-488. Matthews, R. W. 1974. Biology of Braconidae. Annual Review of Entomology 19: 15—32. Metcalf, R. L. 1986. Foreword, pp. 7-15. /n Krysan, J. L. and T. A. Miller, eds. Methods for the Study of Pest Diabrotica. Springer-Verlag, New York, 260 pp. Metcalf, C. L., W. P: Plint, and Ri. Metcalf. 1962: Destructive and Useful Insects. 4th edition. Mc- Graw-Hill, New York, 1087 pp. Nealis, V. G. and S. Fraser. 1988. Rate of development, reproduction, and mass-rearing of Apanteles fu- miferanae Vier. (Hymenoptera: Braconidae) under controlled conditions. Canadian Entomologist 120: 197-204. Schlinger, E. I. and J. C. Hall. 1960. The biology, be- havior, and morphology of Praon palitans Mue- sebeck, an internal parasite of the spotted alfalfa aphid, Therioaphis maculata (Buckton) (Hyme- noptera: Braconidae, Aphidiinae). Annals of the Entomological Society of America 53: 144—160. Shaw, S. R. 1995. A new species of Centistes from Brazil (Hymenoptera: parasitizing adults of Diabrotica (Coleoptera: Braconidae: Euphorinae) Chrysomelidae) with a key to new world species. Proceedings of the Entomological Washington 97: 153-160. Singh, P. 1982. The rearing of beneficial insects. New Zealand Entomologist 7: 304—310. Sutter, G. R. and D. R. Lance. 1991. New strategies for reducing insecticide use in the corn belt, pp. 231-249. In Hullar, T. L., chm. Sustainable Ag- riculture Research and Education in the Field: A Proceedings. National Academy Press, Washing- ton, DC, 448 pp. Vance, A. M. 1932. The biology and morphology of Society of the braconid Chelonus annulipes Wesm., a para- site of the European corn borer. United States De- partment of Agriculture, Technical Bulletin No. 294, 48 pp. Viktorov, G. A. 1968. The influence of the population density upon the sex ratio in Trissolcus grandis Thoms. (Hymenoptera: Scelionidae). Zoologi- cheskii Zhurnal 47: 1035—1039. Viktoroy, G. A. and N. I. Kochetova. 1973. The role 562 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of trace pheromones in the regulation of sex ratio moth parasite, Apanteles melanoscelus. Annals of in Trissolcus grandis (Hymenoptera: Scelionidae). the Entomological Society of America 70: 549— Zhurnal Obshchei Biologii 34: 559-562. 554. Vinson, S. B. 1978. Courtship behavior and source of | Whistlecraft, J. W., C. R. Harris, A. D. Tomlin, and J. a sexual pheromone from Cardiochiles nigriceps. H. Tolman. 1984. Mass rearing technique for a Annals of the Entomological Society of America braconid parasite, Aphaereta pallipes (Say) (Hy- 71: 832-837. menoptera: Braconidae). Journal of Economic En- Weseloh, R. M. 1977. Mating behavior of the gypsy tomology 77: 814-816. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 563-570 A NEW SPECIES OF EUTARSOPOLIPUS BERLESE (ACARI: PODAPOLIPIDAE) FROM THE GALAPAGOS ISLANDS, A PARASITE OF AGONUM CHATHAMI VAN DYKE (COLEOPTERA: CARABIDAE) ROBERT W. HUSBAND 1035 Scottdale Drive, Adrian, MI 49221, U.S.A. (e-mail: rhusband @ adrian.edu) Abstract.—Eutarsopolipus brettae, n. sp. (Acari: Podapolipidae), is described from Agonum chathami Van Dyke (Coleoptera: Carabidae) from the Galapagos Islands and compared with related species of Eutarsopolipus. Characters of E. brettae do not fit with existing subgroups of Eutarsopolipus. Key Words: Mites in the family Podapolipidae (Acari: Heterostigmata) are highly specialized ecto- and endoparasites of the insect orders Blat- taria, Orthoptera, Heteroptera, Hymenop- tera, and especially Coleoptera. The genus Eutarsopolipus Berlese is restricted to hosts in Carabidae (Coleoptera) and occurs worldwide. Along with the type species, E. lagenaeformis Berlese 1913, more than 40 others have been discovered, of which 21 were described by Regenfuss (1968, 1974). This is the first record of Eutarsopolipus from the Galapagos Islands. The purpose of this paper is to describe a new species collected from Agonum chathami Van Dyke and compare it with other species of Eutarsopolipus parasitizing the carabid genus Agonum, with Eutarso- polipus from Central and South America, and with related Eutarsopolipus worldwide. MATERIALS AND METHODS Males, larval and adult females, and eggs of Eutarsopolipus brettae were collected from Agonum chathami Van Dyke _ bor- rowed from the California Academy of Sci- ences, San Francisco, California, U.S.A. The technique for removing mites from mu- beetle, Carabidae, parasitic mites, Acari, Podapolipidae, Galapagos, Ecuador seum specimens is described in Husband and Dastych (1998). Measurements, in micrometers (wm), were taken with the aid of a Zeiss micro- scope with a stage micrometer and drawing tube. The terminology used follows Lind- quist (1986). Eutarsopolipus brettae Husband, new species (Figs. 1—3) Diagnosis.—Adult female E. brettae lack stigmata, cheliceral stylets are long, 75-85, and ambulacral II, III claws are well devel- oped (10). The genital capsule of male EF. brettae has concave lateral margins. Plates C and D are separate in larval female E. brettae, cheliceral stylets are long (60—64), setae of the idiosoma and legs are long (Ta- ble 1), and the larval female has 3 femur I setae in combination with | genu I seta (Ta- ble:2): Adult female (Fig. 1).—Gnathosoma: Length 62—69, width 60—67. Palp length 22-25; cheliceral stylet length 75-85 with basal sclerite 12-15, pharynx width 17, walls of pharynx thick, dorsal gnathosomal 564 Fig. 1. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON H 1} ii @ 4d |p iit fit { 0.1mm Eutarsopolipus brettae, adult female: left, ventral aspect; right, dorsal aspect. VOLUME 104, NUMBER 3 setae 32-35, ventral setae 20—25, distance between ventral setae 18—20. No stigmata. Idiosoma: Length 266—450, width 209— 318. Prodorsal plate setae v, 12—15 v, 7-8, sc, 25-30, setae about 10 from posterior margin of prodorsal plate. Distance be- tween setae v, 55—56; v, lateral to line con- necting v, and sc,. Plate C midlength 60— 70, width 237-265, setae c, 7-8, c, 9-10. Plate D length 50—75, width 175-197, setae 0.1mm d 7-8. Plate EF length 40-53, width 130— 145, setae e 7-8, no setae h,, h,. Venter: Apodemes | and 2 weakly de- veloped, meeting sternal apodeme medially; sternal apodeme not extending beyond junction with apodemes 2. Coxal setae la 3—4, 2a 5—6; setae la situated equidistant to apodemes 1, 2. Distance between setae la 37-52. Coxal setae 3a not evident, 3b 6-7. 566 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ran BS oot we ee eK He ‘ > Ae, \ \ { \ \ ‘ ‘ 1B, BE Legs: Leg setation as in Table 2. Am- bulacrum I with a terminal stout claw, am- bulacra IH, II with 2 strong claws. Tarsus I subunguinal seta spinelike, 2 terminal spines on each of tarsi II, III. Tarsus I so- lenidion w 5—6; tarsus II solenidion w 5. = (o aha) eer | Gann == ae Eutarsopolipus brettae, larval female: left, ventral aspect; right, dorsal aspect. Tibia I solenidion 8—9, adjacent seta k 4— 6. Tibial I, II, II setae d 45-50, 30—33 and 20—22 respectively. Male (Fig. 2).—Gnathosoma: length 37—40, width 36-37. Palp length 13-17; cheliceral stylet length 25-31, no_ basal VOLUME 104, NUMBER 3 sclerite; pharynx width 10—11, dorsal gna- thosomal setae thick 2—4, ventral setae 9— 1S: Idiosoma: Length 170-188, width 120— 147. Prodorsal setae v, m, v, m, setae sc, 53-65. Distance between setae v, 20-21, distance between setae v, 52; setae v, lateral to line connecting setae v, and sc,. Plates C and D fused, setae c,, v,, d, and e vestigial or not present. Genital capsule length 28— 30, width 30-35 at base, concave lateral margins. Venter: Apodemes 1, 2 and sternal apo- deme evident; coxae III separated medially. Setae la, 2a, 3b microsetae, 3a not evident. Legs: Leg setation as in Table 2. Am- bulacrum I with | thick claw, length 4, am- bulacra H, If] with small claws 3. Tarsus I spinelike seta s 5—6, tarsi H, III spinelike setae tc’ and u’ 5—7. Tarsus I solenidion w 5—6, tarsus II solendion w 4—5; tibia I so- lendion & 8, adjacent seta k 2—3. Tibiae I, II, IfI setae d 30-33, 15-22, 19 respective- ly. Genu III seta v” evident in only | male. Larval female (Fig. 3).—Gnathosoma: length 44-53, width 42—45. Palp length 15—20, cheliceral stylet length 60—64, stylet basal sclerite 7—8, pharynx width 12-13, dorsal gnathosomal setae 31—36, ventral se- tae 13—14, distance between ventral setae 14-20. Idiosoma: Length 165—192, width 125— 132. Prodorsal plate setae v, 17—20, v, 9- 16, sc, 84-85. Distance between setae v, 34—45; setae v, lateral to line connecting setae v, and setae sc,. Plates C and D sep- arate, setae c, 7-8, c, 8-10, setae d 7-8: distance between setae d 30—33. Setae c, in line with setae c,. Plate EF oval, setae e 7. Plate H broader at base, setae h, 80—90, h, 5—7, distance between setae h, 2. Venter: Apodemes 1, 2 and sternal apo- deme conspicuous but weakly scerotized. Setae la 3—4, 2a 3-5, 3a m and 3b 5. Dis- tance between setae 3a and 3b 18—20, setae 3a visible only in larval exoskeleton asso- ciated with adult female. Legs: Leg setation as in Table 1. Am- bulacrum I with 2 small, parallel claws. 567 Ambulacra II, HI with small, diverging claws. Tarsus I spinelike seta s 5, tarsi I, III spinelike setae tc’ and wu’ 5—7. Tarsus | solenidia o 3—5, tarsus II solenidion w 5. Tibia I solenidion @ 9-10, seta k 3. Setae tc’ 7-9, tc” 9-10. Tibiae I, II, III setae d 35-40, 18-23 and 17-25 respectively. Egg.—Length 210-240, width 120-135. Type, host, and locality data.—Holotype 2? (RWHO10501-1), allotype ¢ and 24 par- atypes: from Chatham Island, Galapagos Is- lands, Ecuador, from under eleytra of Agon- um chathami Van Dyke (Carabidae) col- lected by F S. Williams, 3 October 1905. Type deposition.—Holotype, allotype, 9 adult 2 paratypes, 6 ¢d paratypes, 3 larval 2 paratypes and 2 egg paratypes of Eutar- sopolipus brettae deposited in the Depart- ment of Entomology, California Academy of Sciences, San Francisco, California, U.S.A. One 6, 2 adult 2, 1 larval 2 and 1 vial of paratypes in the collection of the au- thor. Etymology.—The species is named for Roberta Brett, California Academy of Sci- ences, San Francisco, California, in recog- nition of her support of studies of parasitic mites of insects. DISCUSSION Before Regenfuss (1968) described 16 new species of Eutarsopolipus trom Ger- many, only E. lagenaeformis from Italy and E. desani Cooreman 1952 from Central Af- rica were known. Regenfuss (1968) defined 7 subgroups of Eutarsopolipus. As new species were described from Pacific Islands, Australia, India, S. Africa, and the Western Hemisphere, characters of existing sub- groups of Eutarsopolipus no longer worked in placing new species. Husband (1995) created the ochoai subgroup and Husband and Macfarlane (1999) created the catad- romi and secundus subgroups. There are now more than 40 species of Eutarsopoli- pus and many more expected from the more than 25,000 known species of Carabidae. Major characters used to distinguish sub- groups of Eutarsopolipus include: presence 568 Table 1. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Comparisons of characters and maximum measurements of Eutarsopolipus brettae with Eutarso- polipus species in the pterostichi and agonum groups, E. trichognathi from South America and E. ochoai from Central America. Character E. brettae pterostichi agonum E. trichognathi E. ochoai Adult females Stigmata 0) 0) “PF at + Idiosoma 450 510 498 550 580 Amb. II, III claws 10 7 0) O 3, thin Cheliceral stylets 85 48 40 54 67 Dors. gnath. setae 35 19 20 18 29 Vent. gnath. setae 25 6 5 5 25 Setae v, 15 6 9 5 23 Setae v, 8 9 20 5) 18 Femur I v” 19 m 0) O 20 Tibia I sol. 9 10 4 7) 9 Tibia II d 33 10 15 8 31 Genu I v’ 5 2 O O 4 Adult males Idiosoma 188 220) 196 173 222 Amb. HI, HI claws 5) minute 0) minute 3, thin Cheliceral stylets 31 28 22 D3} 31 Dors. gnath. setae 4, thick 13 i 8 13 Vent. gnath. setae 13 4 m m 9 Setae sc, 65 48 S)i/ 32 62 Femur I vy” 0-12 2 0) 0 0 Tibia I d 33 19 13 aT 31 Larval females Idiosoma 192 270 230 332 AT) Amb. I, II claws 4 minute 0 minute 3, thin Cheliceral stylets 64 49 29 32 49 Dors. gnath. setae 36 30 16 16 26 Vent. gnath. setae 14 7 5 m 15 Femur I v” 14 m 0) 0) 10 Tibia I d 40 8 18 10 26 Tarsus IT sol. w 5S) 2 O 5 5 h,-h, distance 2 d 12 4 O, adj. h, 7 22. 30 25 m Genu I v, 0) 0) 0) 0) 5 or absence of stigmata, shape of the male genital capsule, proportions of cheliceral stylets in relation to gnathosoma, presence or absence of idiosomal setae or setae on legs and presence or absence of idiosomal plates. Major characters used to distinguish species include: relative lengths of chelic- eral stylets, lengths and widths of other structures, idiosomal plates of adult females divided or not, plates of larval females fused or not, idiosomal shape, structure of setae (thick, thin, with or without micro- spines), setal position, and relatives lengths and position of setae of the legs and idio- soma. All Eutarsopolipus that are parasites of the carabid genus Agonum fit patterns de- scribed for the biunguis subgroup proposd by Regenfuss (1968). Comparisons of Eu- tarsopolipus brettae with species in the biunguis group yielded more than 40 dif- ferences. All biunguis species have distinct stigmata, no ambulacral I claw, no or mi- nute ambulacra II, HI claws, no femur I se- VOLUME 104, NUMBER 3 Table 2. 569 Leg setation of femur, genu, tibia and tarsus for larval female E. brettae and related species. Solenidia are included. Two numbers in a column indicate variation in setal numbers. Leg I Leg I I G Ni la EF G ri Ta F G ri Ta E. brettae | a 7 O | 4 6 0 | 4 5 E. ochoai 3 2 if 9 0 | 4 7 0 | 4 5 E. stammeri O 2 6 9 O | 4 6 0) | 4 5 E. trichognathi 2 0 7 8/9 0 0) 4 6 0) 0) 4 5 E. biunguis subgroup from Agonum 1/2 0) 7 8 0) O 4 5 0 O 4 5 E. lukoschusi 3 0) 6 9 0) 0 4 6 0) 0 4 5) E. pterostichi subgroup 2/3 0/2 5/7 8/9 0) 0) 4 6/7 0) 0) 4 5/6 tae v" and no genu I v” setae. Eutarsopolipis brettae has no stigmata and has the struc- tures which are missing in biunguis species. Patterns of characters do not fit patterns for the biunguis subgroup, any of the other 6 subgroups proposed by Regentuss or any of the 3 subgroups added since 1968. An attempt was made to place E. brettae with species which have no stigmata: E. stammeri Regenfuss of the stammeri sub- group, E. lukoschusi Husband 1986 and species in the prerostichi subgroup. In con- trast to large claws of female EF. brettae, no claws are present in E. stammeri. In E. lu- koschusi ambulacral claws II, I are minute and ambulacrum I claw is small. Regenfuss (1974) placed EF. inermis Regenfuss from Georgia, U.S.A., within the pterostichi sub- group because it lacked stigmata, the geni- tal capsule of the male was similar to other mites in this group, solenidia were short as was femur I seta d and larval females had slightly separated setae /,. In contrast to other mites in the pterostichi subgroup, adult females of E. inermis have no genu I setae, all ambulacral claws are reduced and coxal seta 3a setae are not present. The ho- lotype of E. pterostichi Regentuss has 3 fe- mur I setae (as does E. brettae), 2 genu I setae in contrast to | genu I setae in E. bret- tae, and short solenidia w in contrast to lon- ger solenida w on tarsi I, Hl in E. brettae. Setae /i, are nearly adjacent in larval female E. brettae and genital capsules of males have concave rather than straight lateral margins. Although adult female E. brettae do not appear to have setae 3a, ambulacral claws are not reduced as in E. inermis. With the exception of EF. inermis in the pterosti- chi subgroup, cheliceral stylets of all instars of E. brettae are distinctly longer (Table 1). Geographically, the nearest Eutarsopoli- pus species to E. brettae are E. ochoai Hus- band 1995 parasite of Pasimachus spp. bee- tles from Costa Rica and E. trichognathi Husband and Eidelberg 1996 from Trichog- nathus marginipennis (Latreille) from Bra- zil. The species FE. trichognathi occurs in Ecuador, Columbia, Brazil, Paraguay and Bolivia (Husband 1999). In contrast to no stigmata in E. brettae, both E. ochoai and E. trichognathi have stigmata. Eutarsopo- lipus trichognathi shares characters with the biunguis subgroup of Eutarsopolipus. All instars of E. ochoai have small ambulacra II, II] claws in contrast to larger claws in E. brettae. Genu I of E. ochoai has 2 setae in contrast to | seta of E. brettae. Males of E. brettae have short, thick dorsal gnatho- somal setae (4) in contrast to longer gna- thosomal setae (13) in males of FE. ochoai. I consider, on the basis of these characters, that E. brettae is not closely related to the 2 species of Eutarsopolipus from Central and South America. Until a reassessment of subgroups of Eu- tarsopolipus is completed, E. brettae cannot be placed in a subgroup. It has combina- tions of unique characters of no other Eu- tarsopolipus. 570 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ACKNOWLEDGMENTS The assistance of Roberta Brett and Da- vid Kavanaugh, Department of Entomolo- gy, California Academy of Sciences, Gold- en Gate Park, San Francisco, California, U.S.A. in acquiring specimens and advice on host beetles is appreciated. LITERATURE CITED Berlese, A. 1913. Acari Nuovi. Redia 9: 27-87. Cooreman, J. 1952. Acariens Podapolipodidae du Con- go Belge. Bulletin Institut de Institute Royal des Sciences Naturelles de Belgique 28(35): 1—10. Husband, R. W. 1986. New taxa of Podapolipidae (Ac- arina) from S. African Coleoptera: Result of the Namaqualand-Namibia expedition of the King Le- opold Foundation for exploration and protection of nature. Bulletin de I’ Institut Royal des Sciences Naturelles de Belgique: Entomologie 56: 5—14. . 1995. A new species of Eutarsopolipus (Ac- ari: Podapolipidae) from Costa Rican Pasimachus spp. (Coleoptera: Carabidae). Entomologischen Mitteilungen aus dem Zoologische Museum Ham- burg 11(151): 157-165. . 1999. American Eutarsopolipus of the biun- guis group and descriptions of previously un- known male and larval female of Eutarsopolipus trichognathi (Acari: Podapolipidae) from South America. International Journal of Acarology 25(1): 13-17. Husband, R. W. and H. Dastych. 1998. A new species of Eutarsopolipus (Acari: Podapolipidae) from Chlaenius sericeus Frost (Coleoptera: Carabidae) from Athens, Georgia, U.S.A. Entomologischen Mitteilungen aus dem Zoologische Museum Ham- burg 12(158): 317-326. Husband, R. W. and M. Eidelberg. 1996. A new spe- cies of Eutarsopolipus (Acari: Podapolipidae) from Trichognathus marginipennis (Coleoptera: Carabidae) from Brazil. International Journal of Acarology 22(3): 193-197. Husband, R. W. and D. Macfarlane. 1999. Two new species of Eutarsopolipus (Acari: Podapolipidae) from Catadromus lacordairei (Coleoptera: Cara- bidae) from Australia. International Journal of Ac- arology 25(4): 297-308. Lindquist, E. E. 1986. The world genera Tarsonemidae (Acari: Heterostigmata): A morphological, phy- logenetic, and systematic revision with reclassifi- cation of family group taxa in Heterostigmata. Memoirs of the Entomological Society of Canada 136: 1-517. Regenfuss, H. 1968. Untersuchungen zur Morpholo- gie, Systematik und Okolgie der Podapolipidae (Acarina, Tarsonemini). Zeitschrift fiir Wissen- schaftliche Zoologie, Leipzig 177(3/4): 183-282. . 1974. Neue ektoparasitische Arten der familie Podapolipidae (Acari: Tarsonemini) von Carabi- den. Mitteilungen aus dem Hamburg Zoolische Museum und Institut 71: 147-163. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 571-575 NEW HOST RECORD, NEW RANGE INFORMATION, AND A NEW PATTERN OF VOLTINISM: POSSIBLE HOST RACES WITHIN THE HOLLY LEAFMINER PHYTOMYZA GLABRICOLA KULP (DIPTERA: AGROMYZIDAE) SONJA J. SCHEFFER Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, Bldg. 005, Rm. 137, BARC-West, 10300 Baltimore Avenue, Belts- ville, MD 20705, U.S.A. (e-mail: sscheffe @sel.barc.usda.gov) Abstract.—The agromyzid leafminer Phytomyza glabricola is reported from the holly Ilex coriacea, a plant not previously reported to host leafminers. The known geographic range of P. glabricola is extended to include Alabama, Florida, Mississippi, North Car- olina, and South Carolina. In these areas, it can be found feeding on sympatric populations of its hosts /. coriacea and I. glabra. Phytomyza glabricola reared from I. coriacea are univoltine, having only a single generation a year, while on /. glabra this species is multivoltine with at least two generations a year. This suggests that either this leafminer exhibits a high degree of host-associated phenotypic plasticity in life history or that host races are present. Key Words: The agromyzid fly Phytomyza glabricola Kulp belongs to a complex of closely re- lated leafminers, all of which feed on var- ious species of holly in the genus //ex (Aquifoliaceae) (Kulp 1968, Scheffer and Wiegmann 2000). Phytomyza_ glabricola was first described from the plant species Ilex glabra (Linnaeus) Gray by Kulp (1968) in his revision of holly-feeding Phytomyza species. On this host, P. glabricola makes a linear-blotch mine and is multivoltine (Kulp 1968, Al-Siyabi and Shetlar 1998). In 1996, I reared P. glabricola from 26 leafmines found on two bushes of /. cori- acea (Pursh) Chapman growing in North Carolina. This plant species has not been previously reported to host agromyzid leaf- miners (Spencer and Steyskal 1986, Spen- cer 1990). Ilex glabra and I. coriacea are shrubby evergreen hollies that grow sympatrically in host race formation, sympatry, Aquifoliaceae, gallberry, inkberry the coastal plain of the southeastern U.S. The range of /. glabra extends from coastal Nova Scotia to mid Florida to eastern Texas and Missouri and is slightly broader than that of /. coriacea which extends only from coastal Virginia to northern Florida to Tex- as [Galle (1997) includes Mexico in the range, but most other sources do not (Stan- dley 1923, Lundell 1961, Gleason and Cronquist 1963, Radford et al. 1968, Cor- rell and Johnston 1970)]. These two plant species are believed to be very closely re- lated and go by the similar common names gallberry (. glabra) and large gallberry (J. coriacea) (Galle 1997). Phytomyza glabricola has been reported from Connecticut, Massachusetts, New Jer- sey, New York, Ohio, and Washington, D.C. (Kulp 1968, Spencer and Steyskal 1986), but not from the southeastern U.S. where both J. glabra and I. coriacea are S72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Collection locations, hosts, and dates for P. glabricola. en eee ee ee eee eee Location Host Date Alabama Covington Co. I. glabra Jan. 1998 Covington Co. I. coriacea Jan. 1998 Florida Calhoun Co. I. glabra Oct. 1999 Highlands Co. I. glabra Jan. 1998, Oct. 1999 Marion Co. I. glabra Jan. 1999 Wakulla Co. I. glabra Jan. 1998, Oct. 1999 Wakulla Co. I. coriacea Jan. 1998 Mississippi Forrest Co. I. glabra Jan. 1998 Forrest Co. I. coriacea Jan. 1998 North Carolina Brunswick Co. I. glabra Feb. 1997 Brunswick Co. I. coriacea Feb. 1997 Craven Co. I. glabra Feb. 2000 Craven Co. I. coriacea Feb. 2000 Moore Co. I. glabra Feb. 1996 Moore Co. I. coriacea Feb. 1996 New Hanover Co. I. glabra Feb. 1997, Jan. 1998 New Hanover Co. I. coriacea Feb. 1996, Feb. 1997, Jan. 1998 New York Suffolk Co. I. glabra Aug. 1996 South Carolina Berkeley Co. I. glabra Berkeley Co. I. coriacea Feb. 1997, Jan. 1998 Feb. 1997, Jan. 1998 common in the pine woods and bays of the coastal plain. The purpose of this study was to investigate the geographic range of P. glabricola and to determine whether this species commonly feeds on /. coriacea. MATERIALS AND METHODS From 1996 to 2000, I periodically trav- eled within the southeastern U.S. looking for populations of /. glabra and I. coriacea. Leaves containing well-developed mines were removed from plants, and adult flies were reared from the pupae found within the mines. Species identity of the leafmi- ners reared from /. glabra and I. coriacea was determined using Kulp (1968) and Spencer and Steyskal (1986). RESULTS Adult flies reared from /. glabra and I. coriacea were readily identified as P. gla- bricola. Flies reared from the two hosts did not noticeably differ from each other in morphological characters. Locations where P. glabricola was reared from leafmines on I. glabra and/or I. coriacea are listed in Ta- ble 1. Collection dates for samples are listed in Table 1; eclosion of adults occurred with- in approximately one month of this date. Leafmines formed by P. glabricola on both I. glabra and I. coriacea are linear- blotch shaped, although in most cases the extensive feeding during the later instars re- moves tissue surrounding the initial linear mine so that only a blotch is apparent. Gen- erally, a leaf will contain only one mine VOLUME 104, NUMBER 3 with a single larva or pupa present. Occa- sionally, a leaf will have more than one mine, indicating the presence of several lar- vae or pupae. This appeared to be more common on the larger-leafed /. coriacea than on /. glabra. Mines on /. glabra have a tendency to be brown with a blistered ap- pearance, while those on /. more often dark green or reddish-green and do not appear blistered. On both hosts, fly pupation occurs in the mine with the ante- rior pupal spiracles emerging through an “epidermal window,” where just prior to pupation, the larva removes mesophyll ad- jacent to the lower epidermis of the leaf. During the course of this study, it became apparent that populations of P. glabricola on /. coriacea exhibit a univoltine life cy- cle. Well-developed mines are only found in January and February, and the adults eclose from late January to early March. Observations made at other times indicate that first-instar leafmines are found in the summer, second-instar mines in late fall, and third-instar mines only in the winter prior to adult eclosion. This univoltine life cycle is very similar to that of the native holly leafminer P. ilicicola on its host /. opaca (Kulp 1968, Potter and Kimmerer 1986), but it is in sharp contrast to the mul- tivoltine life cycle of P. glabricola on I. glabra (Kulp 1968, Al-Siyabi and Shetlar 1998), where the generation time can be as short as one month (Scheffer, personal ob- servation). corlacea are DISCUSSION These findings confirm that /. coriacea is a widely used host for P. glabricola in nat- ural settings and extends the known geo- graphic distribution of P. glabricola to Al- abama, Florida, Mississippi, North Caroli- na, and South Carolina. It is unlikely that further study will un- cover any additional hosts for P. glabricola. Most holly leafminers are strictly monoph- agous (Kulp 1968, Scheffer and Wiegmann 2000), although in artificial situations (e.g., botanical gardens), additional feeding on 573 non-native //ex species or hybrids may be observed (Scheffer, personal observation; host records listed in Spencer and Steyskal 1986, Griffiths and Piercey-Normore 1995). During the course of this study, P. glabri- cola was reared from I. glabra and I. cor- iacea in both natural and ornamental set- tings, but not from other hollies growing in the same locations, including the evergreen species /. cassine Linnaeus, [. myrtifolia Walter, /. opaca Aiton, and I. vomitoria Ai- ton (Scheffer, unpublished data). Although holly leafmining species are known to exhibit different patterns of vol- tinism, no species previously has been re- ported to exhibit more than one pattern. Many holly leafminers have a multivoltine life cycle with two or more generations a year. In most of these species, the larval feeding period may be as short as a week or two, with a total generation time that of slightly over a month (Scheffer, personal observation; the life cycle of P. opacae Kulp is reported to be even shorter (Kulp 1968)). Multivoltine holly leafminers can be found on either evergreen or deciduous hollies and include the species P. ditmani Kulp, P. opacae Kulp, P. verticillatae Kulp, P. vomitoriae Kulp, and several undescri- bed species (Kulp 1968, Spencer and Steys- kal 1986, Scheffer, unpublished data). In contrast, univoltine holly leafminers have only one generation a year and a highly ex- tended larval developmental period lasting up to ten months that appears to be unique among the leafmining Agromyzidae (Cam- eron 1939, Hering 1951, Potter and Kim- merer 1986). Univoltine holly leafminers have been found only on evergreen hollies and include the species P. ilicicola Loew, P. ilicis Curtis, and an undescribed species on /. myrtifolia (Kulp 1968, Spencer and Steyskal 1986, Scheffer, unpublished data). There are several possible explanations for the observation that P. glabricola has a different life history pattern depending on host plant identity. At the two extremes, these hypotheses can be classified as those involving phenotypic plasticity and those 574 involving genetic differences between flies on the two hosts. First, the rate of larval feeding could be a direct physiological re- sult of some unknown quality of host leaf tissue, such that P. glabricola larvae in I. glabra always develop quickly, while lar- vae in J. coriacea develop very slowly. This situation would be a form of host-mediated phenotypic plasticity and would not require genetic differences in the flies. Consider- ation of the life history patterns of related holly leafminers suggests that an hypothesis of host-mediated phenotypic plasticity, even if found to be valid in the case of P. glabricola, cannot generally explain voltin- ism differences in the holly leafmining group; P. ilicicola and P. opacae both ovi- posit into the soft new leaves of /. opaca, but P. ilicicola exhibits a univoltine life cy- cle on this host while P. opacae is multi- voltine (Kulp 1968, Scheffer and Wieg- mann 2000). An alternative hypothesis is that the flies exhibit different patterns of voltinism be- cause they are genetically different and rep- resent host races or incipient or cryptic spe- cies. Under this scenario, the difference in life history on the two hosts could be either a cause of genetic divergence (i.e., the life history differences may be incompatible ad- aptations to each of the two host plants thereby favoring host race formation) or a result of genetic divergence (i.e., host races diverged for reasons unrelated to voltinism patterns and voltinism patterns have sub- sequently diverged). Note that even if ge- netic host races have formed, the difference in voltinism could still be due to phenotypic plasticity, if it exists, such that voltinism would depend on the host, but each host race would exhibit only one pattern because it uses only one host. Whether P. glabricola on I. glabra and I. coriacea represents a single oligophagous species that feeds on two hosts or a com- plex of closely related but genetically di- vergent host races currently is not known. Although the difference in voltinism on the two hosts is suggestive of host races, the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON voltinism difference by itself does not pre- clude flies from the two hosts from inter- breeding. Adult flies from both hosts emerge in late winter (see dates in Table 1) and can be expected to be present in the same general areas together for several weeks. During this one time each year, in- terbreeding between flies from the two hosts is at least possible. Anecdotal evidence from the field indi- cates that host plant preferences suggestive of host races may be present within P. gla- bricola. When the univoltine leafminers on I. coriacea, which are often extremely abundant, emerge en mass in February, adults can be readily observed on /. cori- acea foliage but not on adjacent /. glabra foliage (Scheffer, personal observation), suggesting a possible preference for /. cor- iacea. This is despite the fact that P. gla- bricola trom I. glabra can be taken into the lab and easily reared to the next generation on /. glabra with no apparent hesitancy to oviposit and little or no larval mortality (Scheffer, personal observation). Results to date of phylogenetic analysis of molecular data are inconclusive with re- gard to the question of host races wihtin P. glabricola. Mitochondrial cytochrome oxi- dase sequences of flies from /. glabra and I. coriacea are nearly identical (< 1% pair- wise divergence across more than 2000 bp; Scheffer and Wiegmann 2000), which is typical of sequence variation found within agromyzid species (Scheffer, unpublished data; see also Scheffer 2000). Using this gene region, there is no evidence of host- associated divergence, but given the esti- mated rate of evolution of insect mitochon- drial genes (approx. 2.3% per million years; Brower 1994), mitochondrial sequences would be unlikely to have sufficient varia- tion to track very recent divergence or spe- ciation events. In conclusion, P. glabricola is widely distributed throughout the coastal plain of the eastern United States. It feeds on two holly species, /. glabra and I. coriacea, which are commonly sympatric throughout VOLUME 104, NUMBER 3 much of their ranges. Differences exhibited by P. glabricola in patterns of voltinism on these two hosts suggest either host-associ- ated phenotypic plasticity in life history or the presence of genetically differentiated host races. Additional studies of morpho- logical variation, mating and oviposition behaviors, and/or highly variable molecular markers are needed to determine the status of P. glabricola \eafminers feeding on /. glabra and I. coriacea. ACKNOWLEDGMENTS I thank Leslie Iskenderian and Kevin Omland for excellent assistance in the field. Douglas Miller, Dave Hawthorne, and two anonymous reviewers provided useful com- ments on the manuscript. For permission to collect leafminers, I gratefully acknowledge authorities at Carolina Beach State Park (North Carolina), Francis Marion National Forest (South Carolina), Conecuh National Forest (Alabama), DeSoto National Forest (Mississippi), Croatan National Forest (North Carolina), Apalachicola National Forest (Florida), Ocala National Forest (Florida), and Archbold Biological Station (Lake Placid, Florida). During part of this work, I was supported by a National Sci- ence Foundation/Sloan Foundation Post- doctoral Research Fellowship in Molecular Evolution (Grant no. BIR-9510795). The Holly Society of America provided funds for an extensive holly leafminer collecting trip in 1998. LITERATURE CITED Al-Siyabi, A. A. K. and D. J. Shetlar. 1998. Inkberry leaf miner, Phytomyza glabricola Kulp (Diptera: Agromyzidae): Life cycle in Ohio. Ohio State Ex- tension Research Special Circular 165-99. Brower, A. V. Z. 1994. Rapid morphological radiation and convergence among races of the butterfly Hel- iconius erato inferred from patterns of mitochon- drial DNA evolution. Proceedings of the National Academy of Sciences USA 91: 6491—6495. Sys) Cameron, E. 1939. The holly leafminer (Phytomyza ilicis, Curt.) and its parasites. Bulletin of Ento- mological Research 30: 173-208. Correll, D. S. and M. C. Johnston. 1970. Manual of the Vascular Plants of Texas. Texas Research Foundation, Renner, TX, 1881 pp. Galle, E C. 1997. Hollies: The Genus //lex. Timber Press, Portland, Oregon, 573 pp. Gleason, H. A. and A. Cronquist. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. Willard Grant Press, Boston, MA, 810 pp. Griffiths, G. C. D. and M. D. Piercey-Normore. 1995. A new agromyzid (Diptera) leaf-miner of moun- tain holly (Nemopanthus, Aquifoliaceae) from the Avalon Peninsula, Newfoundland. Field-Naturalist 109: 23-26. Hering, E. M. 1951. Biology of the Leaf Miners. Junk s’Gravenhage, The Hague, The Netherlands, 420 pp. Kulp, L. A. holly leaf miners (Diptera: Agromyzidae). Uni- versity of Maryland Agriculture Experiment Sta- tion Bulletin A-155: 1—42. Lundell, C. L. 1961. Flora of Texas, Vol. 3. Texas Re- search Foundation, Renner, TX, 433 pp. Potter, D. A. and T. W. Kimmerer. 1986. Seasonal al- location of defense investment in //ex opaca Aiton Canadian 1968. The taxonomic status of dipterous and constraints on a specialist leafminer. Oecolo- gia 69: 217-224. Radford, A. E.; H: Ey Ables;and CR. Bells 1968: Manual of the Vascular Flora of the Carolinas. University of North Carolina Press, Chapel Hill, NC, 1183 pp. Scheffer, S. J. 2000. Molecular evidence of cryptic species within the Liriomyza huidobrensis (Dip- tera: Agromyzidae). Journal of Economic Ento- mology 93: 1146-1151. Scheffer, S. J. and B. M. Wiegmann. 2000. Molecular phylogenetics of the holly leafminers (Diptera: Agromyzidae: Phytomyza): Species limits, speci- ation, and dietary specialization. Molecular Phy- logenetics and Evolution 17: 244—255. Spencer, K. A. 1990. Host specialization in the world Agromyzidae (Diptera). Kluwer Academic Pub- lishers, Dordrecht, The Netherlands, 444 pp. Spencer, K. A. and G. C. Steyskal. 1986. Manual of the Agromyzidae (Diptera) of the United States. United States Department of Agriculture Hand- book No. 638, Washington, DC, 478 pp. Standley, P. C. 1923. Trees and Shrubs of Mexico. Contributions from the United States National Herbarium, Vol. 23. Washington, 1721 pp. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 576-588 LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF GOEDENIA RUFIPES (CURRAN) (DIPTERA: TEPHRITIDAE) ON ISOCOMA ACRADENIA (E. GREENE) E. GREENE IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e- mail: richard.goeden @ucr.edu) Abstract.—Goedenia rufipes (Curran) is an oligophagous, nonfrugivorous, fruit fly (Dip- tera: Tephritidae) producing at least two, probably three, annual generations altogether in the flower heads of Chrysothamnus teretifolius (Durand and Hilgard) H. M. Hall and Isocoma acradenia (E. Greene) E. Greene in southern California. Both of these confirmed hosts are Asteraceae belonging to the subtribe Solidagininae of the tribe Astereae. The egg, second- and third-instar larvae, and puparia are described and figured, and selected characteristics of these stages are compared with those of G. timberlakei (Blanc and Foote), the only other well-known species of Goedenia. The egg of G. rufipes is the first pictured for this genus; it bears a prominent pedicel with semicircular to fusiform micro- pyles. The second instar is white, but the third instar has a dark brown to black venter on the meso- and metathorax and a similarly darkened caudal segment. The prothorax and gnathocephalon of the second and third instars are smooth, mostly free of the minute acanthae that circumscribe most other body segments. Minute acanthae cover the posterior end of the truncated caudal segment, which also is perforated by scattered, open pores. The third instar lacks oral ridges. The anterior thoracic spiracle of the second instar bears three papillae, which are reduced to two papillae in the third instar. The life cycle is of the aggregative type and overwintering occurs in dead flower heads as prepuparial third instars, aS puparia in an open, central cells loosely surrounded by floret fragments and intact undamaged achenes, and as unmated, sexually immature adults. Eurytoma sp. (Hy- menoptera: Eurytomidae) and Pteromalus sp. (Hymenoptera: Pteromalidae) are reported as solitary, larval-pupal endoparasitoids, and Eupelmus sp. (Hymenoptera: Eupelmidae) as possible solitary endoparasitoids. Key Words: Insecta, Goedenia, Chrysothamnus, Isocoma, Asteraceae, nonfrugivorous Tephritidae, biology, taxonomy of immature stages, flower-head feeding, aggregative life cycle, seed predation, parasitoids Most indigenous, western North Amer- _ stages of only one of the eight known spe- ican Myopitini (Diptera: Tephritidae: Te- cies of Goedenia have been described in phritinae) formerly assigned to the Pale- detail, i.e., G. timberlakei (Blanc and arctic genus Urophora Robineau-Desvo- Foote), by Goeden et al. (1995). This pa- idy were reclassified in the genus Goe- per describes the life history and selected denia by Freidberg and Norrbom (1999). immature stages of a second species, G. To date, the life history and immature rufipes (Curran). VOLUME 104, NUMBER 3 MATERIALS AND METHODS The present study utilized specimens of adults reared from 1|-liter samples of mature flower heads of Chrysothamnus teretifolius (Durand and Hilgard) H. M. Hall and /so- coma acradenia (E. Greene) E. Greene (As- teraceae) collected throughout southern California since 1980 (Goeden 1987). The life history study and description of the im- mature stages of G. rufipes were based in large part on dissections of samples of ma- ture and immature flower heads of /. acra- denia collected east of Ocotillo at Coyote Wells and at 42-m elevation, southwestern Imperial Co., during 1990-1999, One-liter samples of excised, immature and mature flower heads containing the scarce larvae and purparia were transported in cold- chests in an air-conditioned vehicle to the laboratory and stored under refrigeration for subsequent dissection, photography, de- scription, and measurement. Ten ova dis- sected from a gravid female as well as two second- and 22 third-instar larvae, and nine puparia dissected from flower heads were preserved in 70% EtOH for scanning elec- tron microscopy (SEM). Prepuparia and pu- paria were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temper- ature for adult and parasitoid emergence. Specimens for SEM were hydrated to dis- tilled water in a decreasing series of acid- ulated EtOH. They were osmicated for 24 h, dehydrated through an increasing series of acidulated EtOH and two, I-h immer- sions in hexamethyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, studied and digitally photographed with a Philips XL-30 scan- ning electron microscope in the Institute of Geophysics and Planetary Physics, Univer- sity of California, Riverside. Five arenas each consisting of a clear- plastic, petri dish were provisioned with a flattened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1994). Each arena contained a vir- S77. gin male and female obtained from emer- gence cages that were used for observations of courtship and copulation behavior. 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 (2001la, b, c), Goeden et al. (1993), Goeden and Headrick (1992), Goeden and Norrbom (2001), Goe- den and Teerink (1997), Teerink and Goe- den (1999), and our earlier works cited therein. Means + SE are used throughout this paper. Digitized photographs used to construct text figures were processed with Adobe Photoshop® Version 6. RESULTS AND DISCUSSION TAXONOMY ADULT.—Goedenia rufipes was described from Arizona as Aleomyia rufipes by Cur- ran (1932), who shortly thereafter reclassi- fied it as Euribia rufipes (Curran 1934, as cited by Foote et al. 1993). Foote (1965) assigned it to Urophora, which Steyskal (1979) adopted in his key to Myopitinae genera and species of Urophora. Freidberg and Norrbom (1999) redesignated most of the indigenous, western North American species as Goedenia, with A. rufipes Curran as the type species. The wing was figured by Curran (1934, as. cited! by Foote etealss 1993) sSteyskal (1979), and Foote et al. (1993). Freidberg and Norrbom (1999) provided line draw- ings of the head in lateral and anterolateral view and the hypandrium and phallopode- me in dorsal view. Immature stages.—The egg, second- and third-instar larvae, and puparium of G. ru- fipes are described below. Egg: Sixteen ova dissected from a 17- day-old female were white, opaque, smooth, elongate-ellipsoidal, 0.55 + 0.008 (range, 0.48—0.60) mm long, 0.18 + 0.005 (range, 0.14—0.20) mm wide, smoothly rounded at tapered basal end (Fig. 1A); 578 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. terior to left; (B) pedicel showing pattern and shapes Egg of Goedenia rufipes: (A) habitus, an- of aeropyles (probably distorted). pedicel prominent, 0.03 mm long, circum- scribed apically by different-sized, semicir- cular to fusiform aeropyles arranged singly or in two rows with their long axes parallel to the long axis of the egg (Fig. 1B). The ova of G. rufipes on average equalled in length the ova of G. timberlakei and were slightly wider, but otherwise agreed with the description provided by Goeden et al. (1995). Unfortunately, the scanning electron micrograph of a possibly deformed pedicel of G. rufipes (Fig. 1B) is the only view of this structure obtained for this genus to date. The eggs and first instars of Goedenia spp. are very small, extremely difficult to find and probably are available for only a short period in nature; moreover, their study is further complicated by the tendency of these tephritids to co-occur with other tephritid genera, i.e., Neaspilota, Procecidochares, Trupanea, in flower heads of the same host (“‘symphagy’’, Goe- den 1997), usually in subordinate numbers, as reported by Goeden (1987). Second instar larva: White, cylindrical, tapered anteriorly, bluntly rounded posteri- orly (Fig. 2A); gnathocephalon conical (Fig. 2B), smooth, with few minute acan- thae ventrally (Fig. 2B-1), lacking oral ridg- es (rugose pads); dorsal sensory organ well- defined, round, flattened (Figs. 2B-2, C-1); anterior sensory lobes (Figs. 2B-3, C-2), separated by vertical medial cleft, with ter- minal sensory organ (Figs. 2B-4, C-3), lat- eral sensory organ (Fig. 2C-4), supralateral sensory organ (Fig. 2C-5), and pit sensory organ (Fig. 2C-6); stomal sense organ (Figs. 2B-5, C-7) ventrolaterad of anterior sensory lobe and fused (Fig. D-1) with la- teralmost of five, foliose, protrudent, lateral integumental petals (Figs. 2C-8, D-2) dor- sad of each mouthhook, two vertical pairs of medial integumental petals between an- terior sensory lobes (Fig. 2C-9); mouth- hook (Figs. 2B-6, D-3) bidentate (Fig. 2D- 3); median oral lobe laterally compressed, apically tapered (Fig. 2D-4), separated from labial lobe (Fig. 2D-5); verruciform sensilla circumscribe posterior third of gnathoce- phalon dorsomedially, dorsolaterally, and laterally (Fig. 2B-7); anterior spiracle with three, subglobose papillae (Fig. 2E); minute acanthae (Fig. 2F) posteriorly directed, spatulate, apically rounded on anterior fourth of meso- and metathorax venters and circumscribing all but posterior three- fourths of first abdominal segment (A1), most of A2 to A6, all but posterior three- fourths of A7, and anterior half of A8. Pos- terior surface of caudal segment not viewed. The habitus of the second instar of G. rufipes (Fig. 2A) approximates that of G. timberlakei (Goeden et al. 1995). Differ- ences noted include five lateral integumen- tal petals in G. rufipes (Figs. 2C-8, D-2), not four, as pictured for G. timberlakei (Goeden et al. 1995). Moreover, the latter VOLUME 104, NUMBER 3 579 Fig. 2. Second instar of Goedenia rufipes: (A) habitus, anterior to right; (B) gnathocephalon, frontolateral view, |—minute acanthae, 2—dorsal sensory organ, 3—anterior sensory lobe, 4—terminal sensory organ, 5— stomal sense organ; 6—mouthhook, 7—verruciform sensilla; (C) gnathocephalon, close-up, 1—dorsal sensory organ, 2—anterior sensory lobe, 3—terminal sensory organ, 4—lateral sensory organ, 5—supralateral sensory lateral integumental petals, 9—medial integumental organ, 6—pit sensory organ, 7—stomal sense organ, 8 petals, mouthhook; (D) oral cavity of gnathocephalon, ventrolateral view, 1—stomal sense organ, 2—lateralmost integumental petal, 3—mouthhook, 4—median oral lobe, 5—labial lobe; (E) anterior spiracle; (F) minute acan- thae on dorsolateral aspect of abdominal segments | and 2, anterior to left. 580 species appears to have only a single pair of medial integumental petals (Goeden et al. 1995), not two pairs, like G. rufipes (Fig. 2C-9). The numbers of these integumental petals were not quantified by Goeden et al. (1995), but their presence and general po- sitions were noted and pictured. Another difference is that the anterior spiracle of the second instar of G. rufipes bears three pa- pillae (Fig. 2E), not two papillae, like G. timberlakei (Goeden et al. 1995). Third instar larva: Oblong-ellipsoidal, roundly tapered anteriorly, bluntly truncat- ed posteriorly (Fig. 3A), integument white, but venter of meso- and metathorax with dark brown to black infuscation (Fig. 6B); caudal segment dark brown or black (Figs. 6B, C); outwardly or posteriorly directed, conical, bluntly or sharply pointed (Figs. 3B-1, F-1) or hemispheroidal (Figs. 4C-1, E-2, F-2) minute acanthae circumscribe an- terior fourth of meso- and metathorax and first abdominal segment (A1) and cover all venters thereof, and circumscribe all but an- terior and posterior fifths of A2-A5, all but posterior quarter of A-7, and cover poste- rior plate of caudal segment (Figs. 4D-2, E- 2, F-2); prothorax smooth, lacking minute acanthae (Fig 3B), but circumscribed by verruciform sensilla dorsally, dorsolaterally, laterally, and ventrolaterally (Fig. 3B-2); gnathocephalon conical, anteriorly flat- tened, and medially divided by vertical cleft (Figs. 3B-3, C), pore dorsoposteriorad of each dorsal sensory organ (Fig. 3C-1); dor- sal sensory organ well-defined, hemispher- ical (Figs. 3C-2, D-1); anterior sensory lobe (Figs. 3C-3, D) bears terminal sensory or- gan (Figs. 3C-4, D-2), lateral sensory organ (Fig. 3D-3), supralateral sensory organ (Fig. 3D-4), and pit sensory organ (Fig. 3D- 5); two medial, papilliform integumental petals (Figs. 3C-5, D-6, E-1), four, lateral, spatulate or papilliform, integumental petals (Figs. 3C-6, D-7, E-2) above each mouth- hook (Figs. 3C-7, E-3), lower, lateral petal separate from stomal sense organ (Figs. 3C- 8, D-8, E-4) ventrolaterad of anterior sen- sory lobe; none (Figs. 3C, D), or sometimes PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON one, oral ridge (Fig. 3E-5) laterad of each anterior sensory lobe; mouthhook bidentate, anterior tooth, concave ventrally (Figs. 3E- 6); median oral lobe laterally compressed, apically pointed (Figs. 3C-9, E-7); anterior thoracic spiracle on posterior margin of prothorax bears two subglobose papillae (Figs. 3B-4, F-2); mesothoracic, lateral spi- racular complex with six verruciform sen- silla in vertical series (Fig. 4A-1), mesotho- racic spiracle not seen; metathoracic lateral spiracular complex with nearly closed, lat- eral spiracle (Fig. 4B-1) and four verruci- form sensilla (Figs. 4A-2, B-2) in vertical series posterior to spiracle; lateral spiracular complex of first abdominal segment con- sists of nearly closed spiracle (Figs. 4A-3, C-1) and three verruciform sensilla in ver- tical series posterior to spiracle (Figs. 4A- 4, C-2); caudal segment with pair of pos- terior spiracular plates (Figs. 4D-1, E-1, F) surrounded by hemispherical minute acan- thae (Figs. 4D-2, E-2, F-1) interspersed dor- somedially, medially, and less so, ventro- medially with open pores (Figs. 4D-3, E-3); each posterior spiracular plate bears three, smoothly flattened, ovoid rimae (Fig. 4F-2), ca. 0.01 mm in length, and four spinose in- terspiracular processes, each ca. 0.004 mm long (Fig. 4F-3). The habitus of the third instar of G. ru- fipes resembles that of G. timberlakei (Goe- den et al. 1995). In both species, the venter of the thorax and the caudal segment are darkly pigmented (Figs. 6C, Goeden et al. 1995) and minute acanthae circumscribe the meso- and metathorax and abdomen, and especially noteworthy, the posterior surface of the caudal segment, also 1s dotted prom- inently with scattered pores (Figs. 4D-3, E- 3; Goeden et al. 1995). The prothorax and gnathocephalon of both species are smooth and free of minute acanthae (Fig. 3C, Goe- denver ale 1995): Two medial and four lateral integumental petals are present in G. rufipes (Figs. 3C-5, =6, D=6., -7,, E-1.-2)2 whereas) G. timber- lakei has two medial and six lateral inte- gumental petals (Goeden et al. 1995, un- VOLUME 104, NUMBER 3 58] Fig. 3. Third instar of Goedenia rufipes: (A) habitus, anterior to left; (B) gnathocephalon and prothorax, frontolateral view, |—minute acanthae, 2—verruciform sensilla, 3—gnathocephalon, 4—anterior spiracle; (C) anterior sensory lobe, 4—terminal sensory gnathocephalon, frontal view, 1—pores, 2—dorsal sensory organ, 3 organ, 5—medial integumental petals, 6—lateral integumental petals, 7—mouthhook, 8—stomal sense organ, 9—median oral lobe; (D) gnathocephalon, close-up, 1—dorsal sensory organ, 2—terminal sensory organ, 3— lateral sensory organ, 4—supralateral sensory organ, 5—pit sensory organ, 6—medial integumental petals, 7— lateral integumental petals, 8—stomal sense organ; (E) oral cavity of gnathocephalon, ventral view 1—medial integumental petals, 2—lateral integumental petals, 3—mouthhook, 4—stomal sense organ, 5—posterior con- cavity on anterior tooth of mouthhook, 6—median oral lobe; (F) anterior spiracle, 1—minute acanthae, 2— papillae. 582 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 20 ime, Fig. 4. Third instar of Goedenia rufipes, continued, (A) lateral spiracular complexes, 1—verruciform sensilla on mesothorax, 2—verruciform sensilla on metathorax, 3—spiracle on first abdominal segment, 4—verruciform sensilla on first abdominal segment, (B) 1—metathoracic lateral spiracle, 2—verruciform sensillum; (C) 1— lateral spiracle of first abdominal segment, 2—minute acanthae; (D) caudal segment, 1—posterior spiracular plates, 2—minute acanthae, 3—pores; (E) caudal segment, close-up, 1—posterior spiracular plates, 2—minute acanthae, 3—pores; (F) posterior spiracular plate, |—minute acanthae, 2—rimae, 3—interspiracular processes, 4—ecdysial scar. VOLUME 104, NUMBER 3 published data). The lateralmost integumen- tal petals are separated from the stomal sense organs in both species (Figs. 3C-8, D- 8, E-4; Goeden et al. 1995). Like G. timberlakei (Goeden et al. 1995, unpublished data), the third instar of G. ru- fipes lacks oral ridges on either side of the mouth opening, and ventral or ventrolateral to the stomal sense organ (Fig. 3E). The mouthhooks of the third instars of G. rufipes (Fig. 3E-6), like those of G. tim- berlakei (Goeden 2001c), are bidentate; however, the teeth of the latter species were described as “‘conical’’, but in the former species the anterior tooth is concave ven- trally (Fig. 3E-6). Unfortunately, a vertical view of the oral cavity such as obtained for G. rufipes (Fig. 3E), was not obtained for G. timberlakei (Goeden et al. 1995, unpub- lished data). The anterior spiracle of both species bears only two papillae (Figs. 3B-4, F; Goe- denvet aly995). The lateral spiracular complex of the me- sothorax of G. rufipes includes six verru- ciform sensilla in a vertical series (Fig. 4A- 1); whereas, in G. timberlakei, this same complex includes only two verruciform sensilla (Goeden et al. 1995). Likewise, the metathoracic lateral spiracular complex of G. rufipes includes four verruciform sensil- la (Fig. 4A-2), but again, only two such sensilla in G. timberlakei (Goeden et al. 1995). Finally, three verruciform sensilla in a vertical series comprise the lateral spirac- ular complex of the first abdominal segment of G. rufipes (Figs. 4A-4, C-2), but only one verruciform sensillum is found on this segment in G. timberlakei (Goeden et al. 1995): Differences noted between the second and third instars of G. rufipes include the reduction in the number of papillae on the anterior thoracic spiracle from three (Fig. 2E) to two (Figs. 3B-4, F-2). The number of lateral integumental petals also is re- duced from five in the second instar (Figs. 2C-8, D-2) to four in the third instar (Figs. 3C-6, D-7, E-2). The stability of these num- 583 bers is questionable, and their separation as medial versus lateral integumental petals is problematic because more precise counts and assessments would require consider- ably more replication of similar views of gnathocephala than were available to the author. Puparia: Light (Fig. 6D) to dark (Fig. 6E), reddish brown with dark brown to black, anterior stripe on venter of meso- and metathorax and similarly dark caudal segment, elongate-ellipsoidal, with smooth- ly rounded anterior end and truncated pos- terior end (Fig. 5A). Anterior end bears in- vagination scar and raised, bilobed, anterior thoracic spiracles (not shown). Flattened posterior end of caudal segment studded with smoothly rounded, hemispherical, mi- nute acanthae (Fig. 5B-1) interspersed with open pores (Fig. 5B-2). A pair of raised, oval, posterior spiracular plates (Fig. 5B-3) each bear three oval rimae interspersed with four, peg-like interspiracular processes (not shown, see Fig. 4F and above description of third instar for details). Ten puparia dis- sected from flower heads of /socoma acra- denia averaged 2.73 + 0.07 (range, 2.50-— 3.17) mmin dength:. 1.157 0:04. @ange; 1.05—1.31) mm in width. DISTRIBUTION AND HOostTs To date, Goedenia rufipes only is known from southern California and southwestern Arizona north of Mexico (Foote et al. 1993); however, it probably ranges well into Mexico attacking flower heads at least of Chrysothamnus teretifolius and Isocoma acradenia. The former host is newly re- ported (Goeden 1987, Foote et al. 1993). Both of these confirmed hosts are Astera- ceae belonging to the subtribe Solidagini- nae of the tribe Astereae (Bremer 1994). Accordingly, G. rufipes probably is a nar- rowly oligophagous tephritid, that to date has not been reared by me from flower heads of several other species of these two, common and widespread, plant genera. Chrysothamnus teretifolius occurs on rocky slopes and flats from 600 to 4,000 m in Cal- 584 Figs 5: anterior to left, (B) caudal segment, 1—minute acan- Puparium of Goedenia rufipes: (A) habitus, thae, 2—pores, 3—posterior spiracular plates. ifornia to southern Nevada and northwest- ern Arizona (Hickman 1993), including the higher parts of the Lower Sonoran Zone and arid, lower margins of the Upper Son- oran Zone as delimited by Shreve and Wig- gens (1964). Isocoma acradenia occurs on sandy or clay soils in alkaline or gypsum flats or slopes below 1,300 m in California, Arizona, Nevada, and Baja California, Mexico (Hickman 1993). BIOLOGY Egg.—Egegs of G. rufipes are inserted singly, pedicel-last, parallel to the long axes of and between the outer phyllaries of closed, preblossom flower heads of [. acra- denia. Larva.—Upon eclosion, the first instar tunneled immediately through the inner PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bracts and into an ovule of a preblossom flower head. It fed with its body perpendic- ular to and its mouthparts toward the recep- tacle within an ovule, which it first exca- vated, then exited and entered an adjacent ovule. The receptacle was neither abraded or pitted by such feeding. Second instars (Fig. 6A) mainly contin- ued feeding on ovules in closed, preblos- som flower heads, but a few were found feeding on soft achenes in open, blossom or postblossom flower heads (Fig. 6A). They usually fed within the ovules or achenes with their bodies perpendicular to the re- ceptacles, but always above the receptacles. Receptacles of 10 flower heads containing second instars averaged 1.45 + 0.12 (range, 0.85—1.99) mm in diameter. These 10 flow- er heads each contained a single larva that had damaged an average of 3.2 + 0.6 (range, 1—6) ovules/achenes, or about 28% of the average total of 11.6 = 0.8 (range, 7-14) ovules/achenes per flower head counted within the 10 flower heads. How- ever, more than 600 uninfested flower heads were individually dissected in order to lo- cate these 10 (1.7%) flower heads infested with second instars. Third instars in flower heads fed with their long axes oriented perpendicular to the receptacles, and with their mouthparts di- rected towards the receptacles, that they usually scored or pitted deeply (Fig. 6B, C). Fifty flower heads (three, closed preblos- som; six, open blossom; 41 postblossom) were dissected that contained an average of 1.3 + 0.1 (range, 1—4) third instars. These 55 flower heads averaged 1.8 + 0.05 (range, |.1—2.9) mm in diameter and con- tained an average total of 13.9 + 0.4 (range, 8—20) ovules/achenes, of which on average 7.1 + 0.5 (range, 1-15) ovules/achenes were damaged or ~51%. However, well over 1,000 flower heads were individually dissected to locate these 55 infested flower heads. Receptacles that consistently were pitted suggested that sap constituted at least part of the diet of third instars of G. rufipes. Goeden (1988), Headrick and Goeden VOLUME 104, NUMBER 3 (1990), Goeden and Headrick (1992), Goe- den’ et al. (1993, 1995), Headrick et. al. (1996), Goeden and Teerink (1997) first noted, described, and discussed sap feeding by florivorous species of Tephritidae in the genera Trupanea, Paracantha, Neaspilota, Tephritis, Goedenia (as Urophora), Dioxy- na, and Xenochaeta, respectively. Upon completing feeding, the larvae oriented with their anterior ends away from the re- ceptacles, retracted their mouthparts, and formed puparia (Figs. 6D, E). Pupa.—The receptacles of nine flower heads that contained an average of 1.3 + 0.2 (range, 1—2) puparia (Figs. 6D, E) av- eraged 1.9 + 0.1 (range, 1.4—2.3) mm in diameter. The receptacles were deeply pit- ted in all nine flower heads, further con- firming that sap constituted part of the diet of third instars. The posterior end of the puparium rested in the smooth cup-like de- pression and the middle and anterior part of the puparium was surrounded by excavated floret fragments that formed a close, central cell not glued to the puparium. Adult.—The premating and mating be- haviors of G. rufipes were not studied in the field, but were observed in petri dish arenas of the type found to be useful with many other nonfrugivorous, tephritid species (Headrick and Goeden 1994). The wings of both sexes were held away from the body at about 45° without supination when at rest (Fig. 6F). Both sexes exhibited wing ha- mation (Headrick and Goeden 1994) throughout the day concurrent with other behaviors, 1.e., grooming, resting, and feed- ing; this also was the most common wing movement reported for G. timberlakei (Goeden et al. 1995). Premating behaviors observed with G. rufipes included males and females tracking individuals of the op- posite sex, during which males sometimes swayed and usually exhibited abdominal pleural distension (Fig. 6F). The male ag- gressively mounted a female by jumping upon her, usually from the front, then turn- ing and forcefully grasping her wing bases, thoracic pleura, and aculeus for purchase, 585 while the female usually struggled, resisted his attempts to part her wings, and pushed against the male with her hind legs and tar- si. The males countered by grasping and raising the oviscape with the mid- and hind tarsi, while positioning the apex of the ex- serted aculeus of a receptive female against his epandrium. Nonreceptive females did not exsert their aculeus or pressed the ex- serted aculeus against the substrate so as to hinder or prevent proper positioning by males. But if receptive, the female allowed the male to raise her oviscape and extended her aculeus to its full length in reponse to his copulatory induction behavior (““CIB”’; Headrick and Goeden 1994, 1999). The CIB mainly consisted of the male rapidly rubbing his hind tarsi, back and forth, along her oviscape. The final mating position commonly had the wings of the male parted at about 20° (Fig. 6H), the wings of the fe- male parted at 60° (Figs. 6G, H), with both pairs of wings centered over the midlines of the flies (Figs. 6G, H). The foretarsi of the male grasped the dorsum of the abdo- men of the female laterally at the thoracic juncture, the midtarsi grasped the oviscape at its base, and the hindtarsi crossed under the oviscape or occasionally rested on the substrate (Fig. 61). The body of the female paralled the substrate with the oviscape raised about 30°, while the extended acu- leus pushed the male upward and backward (Fig. 61). Five pairs were observed to mate once or twice per day for a total of nine matings that lasted an average of 96 (range, 20—285) min. Females became restless be- fore termination of mating by pushing against the males with their hind tarsi, by lofting their wings so as to push them against the males, and by fully extending their aculeus. The male in turn countered this behavior with CIB, rocked from side to side to regain purchase or to avoid the fe- male’s pummeling, and sometimes rapidly vibrated his wings, all of which appeared to calm the female and allow coitus to contin- ue. Females sometimes walked about the arenas carrying the males while remaining 586 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Life stages of Goedenia rufipes (A-E in flower heads of [socoma acradenia): (A) second instar (arrow) feeding on floret; (B) third instar (arrow) feeding deeply in receptacle of flower head; (C) third instar showing dark, ventral infuscation (arrow); (D) newly formed, light brown puparium (arrow); (E) dark overwin- tered puparium (arrow); (F) ventral view of adult male with wings in resting position with inflated abdominal pleura; (G) mating pair, dorsal view; (H) mating pair, ventral view; (1) mating pair, lateral view. Lines = | mm. VOLUME 104, NUMBER 3 in copula. One male was observed to at- tempt mating with a newly dead female continuously for 5 h. Separations of three other pairs were observed, during which the male initially moved slightly forward while pulling the aculeus vertically upward, then rapidly turned and laterally walked off the female, and quickly posteriorly away from the female, while pulling free his genitalia. The separation of one pair took 15 s. Seasonal history.—The life cycle of G. rufipes in southern California follows an aggregative pattern (Headrick and Goeden 1994, 1998) in which the third instar, pu- parium, and some adults variously are the overwintering stages. Adults emerge from some of the puparia formed in late-fall, ear- ly winter (October-December) and these unmated, sexually immature adults over- winter. The remaining, nonfeeding third in- Stars, prepuparia, and puparia Overwinter in dead flower heads remaining on dormant Chrysothamnus teretifolius and Isocoma acradenia. These overwintered individuals emerge as adults in late winter (February— March) and either aggregate on one or more, as-yet-unknown, spring blooming, al- ternate hosts to mate and produce an as-yet- undetected spring generation. Or, the over- wintered and newly emerged adults contin- ue to pass the following spring and summer (April—August), possibly still as non-repro- ductive individuals feeding in mountain meadows and riparian habitats bordering the low-elevation, Sonoran Desert. They eventually aggregate on preblossom, fall- blooming, C. teretifolius and I. acradenia, mate, and subsequently oviposit in the small, newly-formed, closed, preblossom flower heads. Natural enemies.—Fourteen Eurytoma sp. (Hymenoptera: Eurytomidae) and 12 Pteromalus sp. (Hymenoptera: Pteromali- dae) were reared from separate puparia of G. rufipes as solitary, larval-pupal endopar- asitoids. Two Eupelmus sp. (Hymenoptera: Eupelmidae) also were reared from insec- tary cagings of mature flower heads as pos- sible solitary endoparasitoids, as reported 587 from G. Goeden et al. (1995). timberlakei by 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 the Institute of Geophysics and Planetary Physics, University of California, River- side, greatly facilitated my scanning elec- tron microscopy. I also am grateful to Jeff Teerink for his technical assistance and to David Headrick for his helpful comments on an earlier draft of this paper. LITERATURE CITED Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc. Portland, Oregon. Curran, C. H. 1932. New North American Diptera, with notes on others. American Museum Novita- tes 526: 1-13. —.. 1934. The families and genera of North Amer- ican Diptera. C. H. Curran (privately printed). Foote, R. H. 1965. Family Tephritidae, pp. 658-678. In Stone et al., eds. A catalog of the Diptera of America North of Mexico. U. S. Department of Agriculture Handbook 276. 1,696 pp. Foote, R. H., KE 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 A. L. Norrbom. 1999. A generic reclassification and phylogeny of the Tribe My- opitini (Tephritinae), pp. 581—627 (Chapter 23). Jn M. Aluja and A. L. Norrbom, eds. Fruit Flies (Te- phritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, Florida. 944 pp. Goeden, R. D. 1987. Host-plant relations of native Urophora spp. (Diptera: Tephritidae) in southern California. Proceedings of the Entomological So- ciety of Washington 89: 269-274. 1988. Life history of Trupanea imperfecta (Coquillett) on Bebbia juncea (Bentham) Greene in the Colorado Desert of southern California (Diptera: Tephritidae). Pan-Pacific Entomologist 64: 345-351. . 1997. Symphagy among florivorous fruit flies (Diptera: Tephritidae) in southern California. Chapter 3. /n K. Dettner, G. Bauer, and W. V6IkI, eds. Vertical Food Web Interactions: Evolutionary Patterns and Driving Forces. Ecological Studies 130: 39-52. Springer-Verlag, Heidelberg, Ger- many. . 2001la. Life history and description on im- mature stages of Neaspilota footei Freidberg and Mathis (Diptera: Tephritidae) on Aster occidental- is (Nuttall) Torrey and A. Gray (Asteraceae) in southern California. Proceedings of the Entomo- logical Society of Washington 103: 191—206. . 2001b. Life history and description on im- mature stages of Tephritis joanae Goeden (Dip- tera: Tephritidae) on Ericameria pinifolia (A. Gray) H. M. Hall (Asteraceae) in southern Cali- fornia. Proceedings of the Entomological Society of Washington 103: 586—600. . 2001c. Life history and description on im- mature stages of Tephritis teerinki Goeden (Dip- tera: Tephritidae) on Hulsea vestita A. Gray (As- teraceae) in southern California. Proceedings of the Entomological Society of Washington 103: 807-825. 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. Goeden R= Ds) DH? Headricks and Wh vA] Meernmnk. 1993. Life history and descriptions of immature stages of Tephritis arizonaensis Quisenberry (Diptera: Tephritidae) on Baccharis sarothroides Gray in southern California. Proceedings of the Entomological Society of Washington 95: 210— 222 . 1995. Life history and description of imma- ture stages of Urophora timberlakei Blanc and Foote (Diptera: Tephritidae) on native Asteraceae in southern California. Proceedings of the Ento- mological Society of Washington 97: 779-790. Goeden, R. D. and A. L. Norrbom. 2001. Life history and description of adults and immature stages of Procecidochares blanci n. sp. (Diptera: Tephriti- dae) on /socoma acradenia (E. Greene) E. Greene (Asteraceae) in southern Calfornia. Proceedings of the Entomological Society of Washington 103: 517-540. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Goeden, R. D. and J. A. Teerink. 1997. Life history and description of immature stages of Xenochaeta albiflorum Hooker in central and southern Cali- fornia. Proceedings of the Entomological Society of Washington 99: 597-607. Headrick, D. H. and R. D. Goeden. 1990. 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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. Shreve, E and I. L. Wiggens. 1964. Vegetation and Flora of the Sonoran Desert. Volume 2. Stanford University Press, Stanford, California. 1740 pp. Steyskal, G. C. 1979. Taxonomic studies on fruit flies of the genus Urophora (Diptera: Tephritidae). Special Publication, Entomological Society of Washington. 61 pp. Teerink, J. A. and R. D. Goeden. 1999. Description of the immature stages of Trupanea imperfecta (Co- quillett). Proceedings of the Entomological Soci- ety of Washington 101: 75-85. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 589-601 DESCRIPTION OF AEGESEUCOELA BUFFINGTON, NEW NAME, WITH NOTES ON THE STATUS OF GRONOTOMA FORSTER (HYMENOPTERA: FIGITIDAE: EUCOILINAE) MATTHEW L. BUFFINGTON Department of Entomology, Texas A&M University, College Station, TX 77843, U.S.A.; current address: Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: mbuff@citrus.ucr.edu) Abstract.—Aegeseucoela Buffington, a replacement name for Moneucoela Dalla Torre and Kieffer 1910, is described. Aegeseucoela contains two previously described species, Aegeseucoela flavotincta (Kieffer), n. comb., and A. grenadensis (Ashmead), n. comb., both of which are redescribed. The status of Gronotoma Forster is discussed, and the synonymy of Eucoilidea Ashmead with Gronotoma is formally documented. Gronotoma nigricornata Buffington, new name, is proposed to replace Eucoilidea nigricornis Kieffer 1908. Thirty-two new combinations in Gronotoma are given, and a checklist of the world species of Gronotoma is provided. Species in both Aegeseucoela and Gronotoma are common parasitoids of Agromyzidae (Diptera). Aegeseucoela is restricted to the New World tropics and subtropics, and Gronotoma is common in the Afrotropics, as well as Asia, Australia, and the Palearctic and Nearctic regions. Key Words: Eucoiline wasps are endoparasitoids of cyclorrhaphous Diptera inhabiting a variety of habitats. These wasps are generally shiny black to dark reddish brown and range in size from 0.5 mm to 5 mm. The Eucoilinae contain 82 genera and nearly 1000 species, and are by far the most diverse of all figitid subfamilies (Ronquist 1999). Only two ma- jor bodies of work have attempted to clas- sify all of the eucoiline genera (Dalla Torre and Kieffer 1910, Weld 1952), and for the most part, eucoiline classification schemes have resulted in a great deal of chaos (dis- cussed in Nordlander 1982b). Presently available identification keys to eucoiline genera (Dalla Torre and Kieffer 1910, Weld 1952) are largely useless due to the reliance on a few key features, none of which are very dependable. Nordlander (1976, 1978, LOSO 19S 12 Aegeseucoela, Gronotoma, Eucoilinae, Figitidae, Agromyzidae, Cynipoidea 1982a, 1982b) was the first to treat eucoi- line classification from a phylogenetic point of view. Through these works, clear generic and species level definitions were provided for the first tme for a number of Palearctic and cosmopolitan taxa. Nordlander (1982b) summarized his findings by proposing in- formal genus groups defined by explicit morphological criteria, a first step towards a more logical and natural classification scheme. An investigation into the phylogenetics and classification of one of these informal genus groups, the Gronotoma group (Buf- fington, unpublished data), resulted in the identification of a clade of eucoiline wasps of questionable taxonomic placement. Two previously described species were found to belong in this clade, described here as Ae- geseucoela, and both species are rede- 590 scribed. Both species are restricted to the New World tropics and subtropics where they have been reared on numerous occa- sions from agromyzid flies (O. Lewis, un- published data). The first indication of the need for a new eucoiline genus was uncovered during an examination of Kieffer’s eucoiline types. The type specimen of Rhabdeucoela flavo- tincta Kieffer did not possess any of the diagnostic features of the genus Rhabdeu- coela Kieffer, and the species was most likely placed in Rhabdeucoela based on the relatively well-developed mesoscutal keel and large scutellar plate (neither of which are universally diagnostic features of Rhab- deucoela). Furthermore, Rhabdeucoela fla- votincta was later moved to two different genera simultaneously by Weld (1952), Mo- neucoela Dalla Torre and Kieffer 1907 and Tropideucoila Ashmead 1903. This species is not readily accommodated in any of the three genera in which it was previously placed, and it was therefore coded separate- ly in a phylogenetic analysis (Buffington, unpublished data), the results of which in- dicate none of the three genera (1.e., Rhab- deucoela, Moneucoela, and Tropideucoila) will remain monophyletic if this species is included within them. Therefore, Aegeseu- coela is proposed to accommodate this spe- cies and a second species (discussed be- low). The second species, Diranchis grenaden- sis Ashmead, also has a confusing taxo- nomic history. It was one of two species originally included in Moneucoela Dalla Torre and Kieffer 1910, which is itself a junior homonym of Moneucoela Dalla Tor- re and Kieffer 1907. Rohwer and Fagan (1917) apparently missed Kieffer’s 1907 publication containing the description of Moneucoela, and determined that the genus was described as new in Dalla Torre and Kieffer (1910). Further, Rohwer and Fagan (1917) designated Diranchis grenadensis Ashmead as the type species of Moneucoe- la Dalla Torre and Kieffer 1910. Weld (1952) considered this a mistake, removed PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON grenadensis (Ashmead) from type status of Moneucoela Dalla Torre and Kieffer, and designated tinctipennis Kieffer (one of two species described in Kieffer 1907) as the type species of Moneucoela Dalla Torre and Kieffer 1907. Weld (1952) did not include grenadensis within the included species list for Moneucoela, resulting in the placement of the species as incertae sedis. In their revision of Moneucoela Dalla Torre and Kieffer 1907, Diaz and Gallardo (1998) did not mention Moneucoela Dalla Torre and Kieffer 1910, nor the two species that were included in this genus when it was proposed (Dalla Torre and Kieffer 1910). Of the two species originally includ- ed in Moneucoela Dalla Torre and Kieffer 1910, one is congeneric with flavotincta and is redescribed below as Aegeseucoela. The second species belongs in Zaeucoila Ashmead (Buffington, unpublished data) and will be treated in a subsequent paper. Furthermore, since grenadensis Ashmead was designated as the type species for Mo- neucoela Dalla Torre and Kieffer 1910 by Rohwer and Fagan (1917), and Moneucoela Dalla Torre and Kieffer 1910 is preoccupied by Moneucoela Dalla Torre and Kieffer 1907, Aegeseucoela is proposed here as a replacement name for Moneucoela Dalla Torre and Kieffer 1910. The second part of this paper is dedicated to a discussion on the status of the eucoiline genus Gronotoma Forster. Gronotoma re- sides within a basal portion of the eucoiline clade (Fontal et al., in preparation), as is the case with Aegeseucoela (though these two genera do not form a monophyletic group) (Buffington, unpublished data). Similar to Aegeseucoela, species of Gronotoma have all been reared from agromyzid flies (Scheffer unpublished data, Davidson 1963, Harding 1965, Viraktamath et al. 1993), mostly found within the genus Melanagro- myza Hendel (Abe and Konishi 1995; Greathead 1969, 1971). The host preference for agromyzid flies appears to be a plesio- morphic feature within the Eucoilinae (Fontal et al. in preparation). VOLUME 104, NUMBER 3 Gronotoma presently contains 47 de- scribed species (including synonymies pro- posed below), making this genus the most diverse of all eucoiline genera that special- ize on agromyzid hosts. Species of Grono- toma have a worldwide distribution and the Afrotropics are particularly speciose (Quin- lan 1986). Species are also very common in the Oriental (Abe and Konishi 1995), the Palearctic, and the Nearctic regions (Dalla Torre and Kieffer 1910, Weld 1952). Mem- bers of this genus are rarely recorded from the Neotropical Region but are frequently collected throughout subtropical Mexico (Buffington, unpublished data). Because of their potential usefulness in the biological control of pest Agromyzidae and the rela- tively high global diversity of species with- in the genus, the status of Gronotoma is reviewed and includes an important syn- onymy. Phylogenetic evidence (Buffington, unpublished data) supports the conclusions of Hedicke (1930) and Beardsley (1988) with respect to the synonymy of Eucoilidea Ashmead with Gronotoma. A world check- list of the species of Gronotoma is present- ed, with nomenclatural notes where appli- cable. BMNH_ The Natural History Museum, London, UK (S. Lewis). Bernice P. Bishop Museum, Hon- olulu, HI, USA. California Academy of Sciences, San Francisco, CA, USA (W. Pu- laski, R. Zuparko). Canadian National Collection of Insects, Ottawa, Canada (J. Hub- ec Je Read “Masner): Cornell University Insect Collec- tion, Ithaca, NY, USA (E.R. Hoe- beke). Institute of Biology at the Acad- emy of the People’s Republic of Romania. International Centre of Insect Physiology and Ecology, Nairobi, Kenya (R. Copeland). Laboratory of Entomology, Kyo- BPBM CAS CNC CUIC IBPR ICIPE KPU 59] to Prefectural University, Kyoto, Japan. Musée Royale de Il’ Afrique Cen- trale, Tervuren, Belgium. National Museum of Natural His- tory, Smithsonian Institution, Washington, DC, USA (D. Smith). Zoological Institute, Russian Academy of Sciences, St. Peters- burg, Russia. Zoologisches Museum, Humboldt Universitat, Berlin, Germany (EF Koch). MRAC USNM ZIN ZMHB Type specimens (holotypes and_ para- types) were generously loaned to me by the CAS (containing a large portion of Kieffer’s eucoiline types originally housed in Po- mona), the USNM (containing many of Ashmead’s and Weld’s eucoiline types), the BMNH (containing Quinlan’s eucoiline types), and the ZMHB (containing many of Forster’s eucoiline types). While sorting unidentified eucoilines in the AEIC in the summer of 2000, I found an extensive series of eucoilines conspecific with the type specimen of Aegeseucoela fla- votincta, all collected from Costa Rica. Ad- ditionally, a long series of Aegeseucoela flavotincta and of A. grenadensis were sent to me for identification by O. Lewis (Sil- wood Park, UK); these specimens were of particular importance since host data ac- companied each specimen. Unidentified specimens of African Gronotoma were gen- erously loaned to me by Dr. Robert Cope- land (ICIPE). Institutions and individuals that donated ethanol preserved specimens for this study were: CNC; TAMU; and Dr. Owen Lewis, Imperial College at Silwood Park, UK. All SEM images utilized in this study were pre- pared digitally on a JOEL JSM-5600 SEM (operated by James Ehrman, Digital Micro- scope Facility, Mt. Allison University, Sackville, NB, Canada). Terminology follows largely that of Weld (1921, 1952), Nordlander (1982b), Ron- 592 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. to lateral ocellus. C, Mesosoma, dorsal view; A = parapsidal hair line; B = well developed lateraldorsal pro- jections of scutellum; C = pronotal triangle. quist and Nordlander (1989), and Ronquist (1995), with the following modifications: parapsidal ridges is preferred over parapsi- dal furrows, as found in Weld (1952); or- bital furrows and pronotal triangle are new terms and are defined as such below. Orbital furrow (Fig. 1B).—A _ distinct Aegeseucoela flavotincta. A, Side view, habitus. B, Head, anterior view; A = orbital furrow complete groove originating at either the lateral ocel- lus or the lateral side of the torulus (de- pending on the taxon) and lining the inner orbit of the eye, terminating at the clypeal margin after fusing with or paralleling the malar sulcus. Pronotal triangle (Figs. 1C, 2C).—An VOLUME 104, NUMBER 3 area on the dorsal surface of the pronotum bordered by the lateral pronotal carina, the pronotal plate and the anterior margin of the mesoscutum; found in genera with well de- veloped pronotal ridges. Aegeseucoela Buffington, new name Moneucoela Dalla Torre and Kieffer 1910: 103, 888. Type species: Diranchis gren- adensis Ashmead, designated by Rohwer and Fagan 1917. Preoccupied by Moneu- coela Dalla Torre and Kieffer 1907. Diagnosis.—Orbital furrows originating either at lateral ocellus or lateral side of to- rulus. Genal carina well developed, often flanged posterior to compound eye. Meso- scutal keel present, at least anteriorly. Par- apsidal ridges absent. Parapsidal hair lines present. Laterodorsal projections of scutel- lum present to absent. RI never tubular or pigmented (radial cell open). Most similar to Zaeucoila and Agrostocynips, but differs by the presence of the extended orbital fur- rows (in some species), the presence of par- apsidal hair lines and R1 incomplete. Description.—Head: Nearly glabrous, with a few scattered setae along lower face, clypeus, inner orbits of compound eyes, malar space and gena; orbital hair patches present. Ventral %4 of lower face with ad- medial clypeal furrows converging towards clypeus. Orbital furrows present, originat- ing at lateral side of torulus or at lateral ocelli (species dependent), terminating at malar sulcus. Malar sulcus simple. Malar space smooth with a single prominent con- ical protuberance. Genal carina present, ex- tending from malar space to lateral ocelli, often undulating posterior to compound eye: Antenna: Female, 13 segments, monili- form, semi-clavate; segments 3—13 sube- qual in length; rhinaria present on segments 3-13. Male, 15 segments, filiform; rhinaria present on segments 3—15; segments 4—15 subequal in length. Segment 3 slightly lon- ger than 4, curved outwardly, excavated lat- erally. 593 Pronotum: Pronotal plate wide, with se- tae along dorsal margin; slightly crested and bifurcate dorsally; pronotal fovea open. Pronotal triangle present (Figs. 1C, 2C). Pronotal impression absent. Lateral aspect of pronotum (below pronotal triangle) smooth and glabrous. Lateral pronotal ca- rina absent. Mesoscutum: Smooth with some setae. Mesoscutal keel present, reaching posterior margin of mesoscutum; tapering posterior- ly. Parapsidal ridges absent. Parapsidal hair lines present (Figs. 1C, 2C). Parascutal im- pression incomplete, narrow. Notauli ab- sent. Mesopectus: Upper part and lower part of mesopleuron glabrous and smooth. Dor- sal margin of mesopleural triangle well de- fined, rounded ventrally. Mesopleural cari- na simple. Lower part of mesopleuron bor- dered by distinct precoxal carina; anterior surcoxal depression present, reticulate. Scutellum: Scutellar plate ranging from medium to large; mid pit placed centrally on plate; plate truncated posteriorly; nearly always bearing tubercles and setae on dor- sal surface. Dorsal surface of scutellum re- ticulate; margined laterally and posteriorly. Laterodorsal projections present, present to nearly absent (Figs. 1C, 2B, 2C); posterior projections absent. Metapectal-propodeal complex: Meta- pectus nearly glabrous with a few scattered setae posteriorly. Spiracular groove with a well defined dorsal margin and a well de- fined to rounded ventral margin. Posterior margin of metapectus ridged. Metapleural ridge absent; submetapleural ridge variable from present to absent. Anterior impres- sions of metepimeron and metepisternum present. Anteroventral cavity semi-circular and setose. Propodeum covered in long se- tae. Lateral propodeal carinae semi-parallel, bowed at junction with auxiliary propodeal carinae; auxiliary propodeal carinae indis- tinct. Nucha glabrous, reticulate. Wings: Hyaline, with base of forewing sometimes darkened; usually setose basally, always setose apically. R1 incomplete, mar- 594 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. lateral projections of scutellum. C, Mesosoma, dorsal view; A projection of scutellum; C = pronotal triangle. ginal cell as long as deep. Apical fringe pre- sent, short. Legs: Fore and mid coxae about the same size, hind coxa about twice size of either fore or mid coxae. Fore coxa variably cov- ered in long setae; mid coxa with anterior and posterior dorsoventral setal bands; hind Aegeseucoela grenadensis. A, Side view, habitus. B, Mesosoma, posterolateral view; A = reduced parapsidal hair line; B = reduced lateraldorsal coxa with a prominent setal band on hind margin. Femora and tibiae sparsely setose; tarsomeres with dense, appressed setae. Length of hind tarsomere | equal to 0.5 combined length of remaining hind tarso- meres. Metasoma: Female: Sub-equal in size to VOLUME 104, NUMBER 3 mesosoma. Base of syntergum with a hairy ring present, ranging from complete to dor- sally bare, comprised of short, semi-ap- pressed setae and longer erect setae; re- mainder of metasoma glabrous. Micropunc- tures present on posterior 4 of syntergum, and on remaining terga. Terga posterior to syntergum directed posteroventrally, result- ing in a 70 degree angle between syntergum and remaining terga. Male: As in female but terga posterior to syntergum abruptly angled ventrally, resulting in a 90 degree angle between syntergum and remaining terga. Biology.—I have examined specimens reared by O. Lewis from species of agro- myzid flies in the genera Haplopeodes Steyskal (on Solanaceae) and Calycomyza Hendel (on Fabaceae). Distribution.—Neotropical Region: Cos- ta Rica, Belize, Mexico (Veracruz, Tamau- lipas), Panama. Nearctic Region: USA: AZ. Previously only known from Guatemala (flavotincta) and Grenada (grenadensis). Etymology.—Aeges; from Greek my- thology, the name of Athena’s dreaded shield, used here to reference the broad, shieldlike pronotal plate present in this ge- nus; eucoela, a suffix frequently used by J.J. Kieffer in his treatments of the Neo- tropical Eucoilinae. Comments.—Though this genus is close- ly related to Zaeucoila and Agrostocynips, phylogenetic evidence (to be found in a forthcoming publication by the author) sug- gests that neither of these genera will re- main monophyletic if the species of Aege- seucoela are placed within them. The par- apsidal hair lines, extended lateral protu- berances on the scutellum and incomplete R1 vein on the forewing are reliable auta- pomorphies for the genus, and reliable syn- apomorphies for the species within the ge- nus. INCLUDED SPECIES Aegeseucoela flavotincta (Kieffer), n. comb. Rhabdeucoela flavotincta Kieffer 595 1908:46, holotype in CAS (#10537). Re- described below. Aegeseucoela grenadensis (Ashmead), n. comb. Diranchis grenadensis Ashmead 1900: 248, holotype in BMNH. Rede- scribed below. Aegeseucoela flavotincta (Kieffer), new combination (Fig. 1) Description.—As in description of genus except as follows: Head: Orbital furrows originating at lateral ocelli (Fig. 1B); genal carina continuing to near lateral ocelli, un- dulating posterior to compound eye (Fig. 1A). Pronotum: Pronotal crest prominent, bifurcate; pronotal ridge well developed. Scutellum: Scutellar plate large, nearly round; laterodorsal projections of scutellum sometimes well developed (Fig. IC). Me- tapectal-propodeal complex: Submeta- pleural ridge usually well developed, con- necting ventral margin of spiracular groove with posterior margin of mesopleuron. Wings: Base of forewing occasionally dark- ened; base of forewing ranging from gla- brous to setose; forewing always setose api- cally. Metasoma: Hairy ring at base of syn- tergum often highly reduced (narrow), but always present. Material examined.—Holotype 2, Champerico, Guatemala. Coll. Baker. CAS #10537; the type specimen is in good con- dition, with Kieffer’s original determination label (a large red label), the collection data labels, depository label and my determina- tion label (slender white label). Additional material: BELIZE: Las Cuevas, Chiquibul Forest, Cayo District, 550 m, (various dates between Oct. 1997 and Sept. 1998), O.T. Lewis (13°23 5"'6)., BOLIVIA:* Yungas; XII.4.84, 2,400 m, Luis Pefia (1 2); COS- TA RICA: S. Rosa Park, Guan., various dates between 27.V.1976 and 10.VIII.1978, D.H. Janzen, Dry Hill (46 6, 90 2). MEX- ICO: Veracruz, 2 km SW Fortin, 8°54'N, 97°00’ W, 2,700’, 23.V1.1997, J.B. Woolley, screen sweep (1 2); Veracruz, El Crucero nr Puente Nacional, 19°20'’N, 96°26'W, 596 13.V1.1997, L.A. Wilson & J.B. Woolley (1 2); Tamaulipas, 97 km Ciudad Victoria, Hwy 70, 3.VII.1986, G. Zolnerowich & R. Trevino (1 2). PANAMA: Colon Prov., 2 km S Sabanitas, 4—15.VII.1999, 120 m, Gillogly & Woolley, MT 99/033 (1 @). U.S.A: Arizona, Portal, 19—23.VIII.1987, H. & M. Townes (4 ¢). Distribution.—Neotropical and southern Nearctic regions (see above list of locali- ties). Biology.—I have examined specimens reared from the agromyzid flies Haplopeo- des sp. on Solanum erianthum D. Don (So- lanaceae) and Calycomyza hyptidis Spencer on Hyptis capitata Jacq. and H. urticoides Kunth (Lamiaceae) (data from O. Lewis). Aegeseucoela grenadensis (Ashmead), new combination (iss) Description.—As in description of genus except as follows: Head: Orbital furrows originating at torulus; genal carina reduced, non-undulating, terminating at dorsal mar- gin of compound eye (Fig. 2A). Pronotum: Pronotal crest reduced, sometimes absent; pronotal ridge sometimes absent. Scutel- lum: Scutellar plate ranging from medium to small; laterodorsal projections usually re- duced/absent (Figs. 2B, 2C). Metapectal- propodeal complex: Submetapleural ridge completely reduced. Wings: Base of fore- wings occasionally darkened; usually entire forewing surface is setose. Metasoma: Hairy ring at base of syntergum thick and densely pubescent. Material examined.—Holotype @; Bal- thazar (windward side), Grenada, West In- dies. H.H. Smith (Coll.), BMNH; the type specimen is in poor condition, with the head plus thorax on one end of a pinned card, and the metasoma on the other end. The diagnostic features discussed below for this genus are all visible. Two ‘type’ labels are present on the specimen, one with a red circle and the second labeled ‘BM Type Hym., y. 50’. Under this lies a label with ‘paratype’ printed on it. Below that is Ash- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mead’s original determination label (in Ashmead’s hand). Below that is the collec- tion data, and finally below that, my des- ignation label. Additional material: BE- LIZE: Las Cuevas, Chiquibul Forest, Cayo District, 550 m, (various dates between Oct. 1997 and Sept. 1998); O:1. Lewist@7 a2; 20 3). MEXICO: Veracruz, 0.7 mi N Jilo- tepecs 19°36 N, 196756 W, 32680" Bscreen Sweep; 14.V1.1997; L.Aq Wilson 5B: Woolley (1 2). PANAMA, Prov. Colon, Quebrada Lopez, MT, 2—4.VII.1999, A. Gillogly (1 2). WENEZUELA: Merida, Merida City, 8°35'54"N, 71°08'42”W, 1860 m, sweep veg. along trib. to Chama R., 3.V.1981, L. Masner (1 @). Distribution.—Neotropical Region (see above list of localities). Biology.—I have examined specimens reared from the agromyzid flies Calycomy- za verbenivora on Verbena sp. (Verbena- ceae); Calycomyza c.f. cassiae (Frost) on Senna cobanensis (Britton) H.S. Irwin & Barneby (Fabaceae); Haplopeodes sp. on Solanum erianthum (Solanaceae) (data from O. Lewis). Comments.—The type specimen of Dir- anchis grenadensis Ashmead, as noted above, is labeled ‘paratype.’ In the original description of D. grenadensis, Ashmead stated “‘described from 1 female speci- men.” Since the single specimen, the lo- cality data, and the description all agree, this must be the holotype. Gronotoma Forster Gronotoma Forster 1869: 342, 346. Type species: Gronotoma sculpturata Forster 1869: 346, by original designation. Eucoilidea Ashmead 1887: 150, 154. Type species: Eucoilidea canadensis Ashmead, by subsequent designation (Ashmead 1903); synonymy by Hedicke (1930) and Beardsley (1988). Eucoelidea Dalla Torre 1893: 1901: 159. Emendation. Afrostilba Benoit 1956: 544. Type species: Afrostilba nitida Benoit, by monotypy. Synonymy by Quinlan (1986). 15; Kieffer VOLUME 104, NUMBER 3 Ashmead (1887) proposed the genus Eu- coilidea to accommodate E. longicornis and E. canadensis, but did not specify a type species. Later, Ashmead (1903) designated Eucoilidea canadensis as type species. Dal- la Torre (1893) emended the original Ash- mead spelling of Eucoilidea to Eucoelidea, and cites the former in brackets. Eucoelidea was recognized in Kieffer (1901), and Kief- fer (1907, 1909) described four new species (and two varieties) in ‘“Eucoelidea Ashm.’ None of Kieffer’s treatments of Euwcoelidea mention the name’s emendation from Ew- coilidea. Based on Article 33.2.1 of the ICZN (1999), Eucoelidea is an emendation of Eucoilidea and is not available. Burks (1979) reported two apparent misspellings of Eucoilidea, namely Eucoilidia (Kieffer 1907) and Eucoelidia (Kieffer 1909); after consulting the original references, neither of these misspellings were found to exist. It is apparent from Ashmead’s (1887) original description of Eucoilidea that he made no comparison with Gronotoma be- fore proposing the new genus, and Hedicke (1930) was the first to propose Eucoilidea as a synonym of Gronotoma. Weld (1952) and Quinlan (1986, 1988), however, sub- sequently maintained Eucoilidea as distinct from Gronotoma. Beardsley (1988) again synonymized these two genera. Though the holotype of E. canadensis is in poor con- dition, with great portions of the body ob- scured by blackish glue, features such as the notauli, the lateral pronotal carinae and the scutellar plate are all clearly visible. Comparing these features with those found in Gronotoma sculpturata clearly indicate that indeed, Eucoilidea is a junior synonym of Gronotoma, supporting the decisions of Beardsley (1988) and Hedicke (1930). Species of Gronotoma can be recognized by the following diagnostic features: lateral pronotal carina present (a distinct ridge, dis- tal to the lateral margins of the pronotal plate, that give species in the this genus the appearance of having a large pronotal plate); notauli present and well-developed in nearly all species; scutellar plate large; 597 hairy ring at base of syntergum absent. Spe- cies of Gronotoma are most easily confused with Diglyphosema Forster, but easily dis- tinguished from that genus by having the scutellum meeting the scutellar plate at an acute angle (meets at a 90 degree angle in Diglyphosema), and the scutellar plate not as elongate (though some species of Gron- otoma tend to have an oval scutellar plate). Previous works treating the synonymy of Eucoilidea with Gronotoma focused pri- marily on the type species; thus, as indicat- ed in the list below, most of the described species have not been formally transferred Hence, 32 new combina- tions are proposed, mainly reflecting the taxonomic work on this genus by Weld (1952) and Quinlan (1986). Species for which type material (holotypes and/or para- types) were examined are indicated by an * The reader should also note that Kieffer (1901) did indeed treat four species in (and we, Gronotoma carinata (Cresson), G. minor (Provancher), G. nigricornis Kietfer and G. ovalis (Thomson) (see in list below). to Gronotoma. Gronotoma not Eucoilidea), Included Species adachiae Beardsley 1988: 39, holotype in BPBM. *advena (Quinlan), n. comb. Eucoilidea advena Quinlan 1986: 262, 263, holotype in MRAC, paratypes in BMNH. allotriaeformis (Giraud). Eucoila allotriae- formis Giraud 1860: 142. Gronotoma al- lotriaeformis: Forster 1869: 346. Type depository presently unknown. *arcuata (Kieffer), n. comb. Eucoelidea arcuata Kieffer 1909: 65. Type in CAS. *bakeri (Kieffer), n. comb. Eucoelidea bakeri Kieffer 1907: 107. Type in CUIC. *bakeri var. cupularis (Kieffer), n. comb. Eucoelidea bakeri var. cupularis Kieffer 1907: 108. Type in CAS. *bakeri var. flavipes (Kieffer), n. comb. Eu- coelidea bakeri var. flavipes Kieffer 1907: 108. Type in CAS. *bucca (Quinlan), n. comb. Eucoilidea 598 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bucca Quinlan 1986: 263, holotype in MRAC, paratypes in BMNH. *canadensis (Ashmead). Eucoilidea cana- densis Ashmead 1887: 154. Gronotoma canadensis: Hedicke 1930; Beardsley 1988. Holotype in USNM. carinata (Cresson). Eucoila ? carinata Cresson 1865: 6. Gronotoma carinata: Kieffer 1901: 159. Type depository pres- ently unknown. *compressa (Quinlan), n. comb. Eucoilidea compressa Quinlan 1986: 263, 264, ho- lotype in MRAC, paratypes in BMNH. *conversa (Quinlan), n. comb. Eucoilidea conversa Quinlan 1986: 264, holotype in MRAC, paratypes in BMNH and MRAC. *crenulata (Kieffer), n. comb. Eucoilidea crenulata Kieffer 1908: 47. Type in CAS. *dilitata (Kieffer), n. comb. Eucoelidea dilitata Kieffer 1907:108. Type in CAS. domestica Girault 1932: 3. Type depository presently unknown. *dubia (Quinlan), n. comb. Eucoilidea du- bia Quinlan, 1986: 264, 265, holotype in BMNH, paratypes in BMNH and MRAC. extraria (Quinlan), n. comb. Eucoilidea ex- traria Quinlan 1986: 265, holotype and paratype in MRAC. *fetura (Quinlan), n. comb. Eucoilidea fe- tura Quinlan 1986: 265, 266, holotype in MRAC, paratypes in BMNH. fulvicornis (Hedicke). Ganaspis fulvicornis Hedicke 1913: 445. Gronotoma fulvicor- nis: Hedicke 1934: 704. Holotype and four paratypes in ZMHB. *furcula (Quinlan), n. comb. Eucoilidea furcula Quinlan 1986: 266, holotype in BMNH, paratypes in BMNH and MRAC. *fuscipes (Kieffer), n. comb. Eucoelidea fuscipes Kieffer 1907: 112. Type in CAS. gracilicornis Cameron 1889: 15. Type de- pository presently unknown. guamensis (Yoshimoto), n. comb. Eucoili- dea guamensis Yoshimoto 1962a: 107, holotype and paratypes in BPBM. hiranoi Abe and Konishi 1995: 309-311, holotype and paratypes in KPU. insularis Ashmead 1895: 743, holotype in BMNH. lacerta (Quinlan), n. comb. Eucoilidea lac- erta Quinlan 1986: 266, 267, holotype and paratype in MRAC. lana (Quinlan), n. comb. Eucoilidea lana Quinlan 1986: 267, holotype and _ para- type in MRAC. *leptis (Quinlan), n. comb. Eucoilidea lep- tis Quinlan 1986: 267, 268, holotype in BMNH. longicornis (Ashmead), n. comb. Eucoili- dea longicornis Ashmead 1887: 154. Type depository presently unknown. maquilingensis (Kieffer), n. comb. Eucoil- idea maquilingensis Kieffer 1914: 184— 185. Type depository presently unknown. *marcellus (Quinlan), n. comb. Eucoilidea marcellus Quinlan 1986: 268, holotype in BMNH, paratypes in BMNH and MRAC. *maurt (Quinlan), n. comb. Eucoilidea mauri Quinlan 1986: 268, 269, holotype and paratype in BMNH. melanagromyzae Beardsley 1988: 40, ho- lotype in BPBM. micromorpha (Perkins). Eucoilidea micro- morpha Perkins 1910: 676. Gronotoma micromorpha: Beardsley 1988: 38. Eu- coilidea rufula Yoshimoto 1962b: 845, synonymy by Beardsley (1988), holotype and paratypes in BPBM. minor (Provancher). Eucoila minor Pro- vancher 1888: 398. Gronotoma minor: Kieffer 1901: 159. Type depository pres- ently unknown. nigra lonescu 1963: 10, holotype in IBPR. *nigricornata Buffington, new name. Eu- coilidea nigricornis Kieffer 1908: 48. Preoccupied by Gronotoma nigricornis Kieffer 1901: 1595 Paratypes miCUIce. nigricornis Kieffer 1901: 159. Type depos- itory presently unknown. *nitida (Benoit), n. comb. Afrostilba nitida Benoit 1956: 544, holotype in MRAC, paratypes in BMNH. Eucoilidea nitida: Quinlan 1986: 269. ovalis (Thomson). Cothonaspis ovalis Thomson 1877: 817. Gronotoma ovalis: Kieffer 1901: 159. Type depository pres- ently unknown. *pallida (Quinlan), n. comb. Eucoilidea VOLUME 104, NUMBER 3 pallida Quinlan 1986: 269—270, holotype in BMNH, paratypes in BMNH MRAC. *parma (Quinlan), n. comb. Eucoilidea parma Quinlan 1986: 270, holotype in BMNH, paratypes in BMNH and MRAC. and presently unknown. *perangusta (Quinlan), n. comb. Eucoili- dea perangusta Quinlan 1986: 270, 271, holotype and paratypes in BMNH. quadrisulcata (Hedicke), n. comb. Eucoil- idea quadrisulcata Hedicke 1922: 229. Two syntypes in ZMHB. rufipes (Gillette), n. comb. Eucoilidea ru- fipes Gillette 1891: 205. Type depository presently unknown. *sculpturata (Forster). Eucoila sculpturata Forster 1855: 257. Gronotoma sculptur- ata: Forster 1869: 342, 346. Holotype in ZMHB. *seychellensis Kieffer 1911: 309. Holotype and paratypes in BMNH. sugonjaevi Belizin 1973: 22, holotype in ZIN; not examined but characters pre- sented in the original description suggest this species probably belongs in Nordlan- deria. *trulla (Quinlan), n. comb. Eucoilidea trul- la Quinlan 1986: 271, holotype in BMNH, paratype in MRAC. *tyrus (Quinlan), n. comb. Eucoilidea tyrus Quinlan 1986: 271, holotype in BMNH. *yurundiensis (Benoit), n. comb. Eucoilidea urundiensis Benoit 1956: 548, holotype in MRAC, paratypes in BMNH and MRAC. Additional material examined.—Sp. 1: Nearctic Region: Canada (AB, BC, MB, ONVOU; PO): USA dE. MD..NY, PA, TX, VA). Neotropical Region: Mexico (Tamau- lipas, Chiapas). Sp. 2: Nearctic Region: Canada (AB, MB); USA. (AZ, CA, FL: IN, MO; NGC, NM, PA, TN, TX, WA). Neotropical Re- gion: Mexico (Veracruz); Venezuela (Mer- ida). 599 Sp. 3: Nearctic Region: Canada (ON); USA (NC). Specimens of other species from: Neo- tropical Region: Mexico (Oaxaca, Guerre- ro); Ecuador; Nicaragua. Afrotropical Re- gion: Uganda; Zaire; Kenya. ACKNOWLEDGMENTS I thank Dr. Bob Wharton and Dr. Jim Woolley for guidance in this project at Tex- as A&M University; I also thank Dr. Géran Nordlander and Dr. Fredrik Ronquist for additional guidance in understanding eucoi- line morphology, taxonomy, and evolution. Special thanks to all those individuals listed under “‘Materials and Methods” who as- sisted with the loan of material for my ex- amination, including James Ehrman _ for preparation of SEM’s and Dr. Owen Lewis for the use of host records. Additional thanks to Dr. Kathy Schick, Dr. David Smith, and one anonymous reviewer for critical comments on an earlier version of this manuscript. Finally, thanks to my wife for her unending support of my research. This research was entirely funded by the NSF PEET Project, #DEB9712543, award- ed to Drs. Wharton and Woolley. All re- search was carried out in the Department of Entomology, Texas A&M University. LITERATURE CITED Abe, Y. and K. 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WASH. 104(3), 2002, pp. 602-613 ENTOMOBALIA, NEW GENUS, THE FIRST MEMBER OF NYCTELIINI (COLEOPTERA: TENEBRIONIDAE) FROM BRAZIL GUSTAVO E. FLORES AND CHARLES A. TRIPLEHORN (GEF) Instituto Argentino de Investigaciones de las Zonas Aridas (ADIZA CRIGYT): Casilla de Correo 507, 5500 Mendoza, Argentina (e-mail: gflores@lab.cricyt.edu.ar); (CAT) Department of Entomology, Museum of Biological Diversity, The Ohio State Uni- versity, 1315 Kinnear Road, Columbus, OH 43212, U.S.A. (e-mail: ctriplhrn@aol.com) Abstract.—Entomobalia, new genus, (Pimeliinae: Nycteliini) is described from north- eastern Brazil and is the first genus of the tribe Nycteliini recorded for that country. This new genus is created on the basis of two previously described species: Asida platynotos Perty and Asida picta Perty (new combinations in Entomobalia), formerly placed in the genus Scotinus Kirby (Pimeliinae: Asidini). Sixteen different character states among the species of Scotinus and the two species assigned to Entomobalia are discussed, 11 of which are shared by Entomobalia with all or some genera of Nycteliini and not with the remaining species of Scotinus. A description for the genus and redescriptions of the two species are provided. Main diagnostic characters for Entomobalia are in sexual dimor- phism and male and female genitalia. Habitus photographs, illustrations of external mor- phology, internal skeletal anatomy, genitalic features, and a distribution map are included. Key Words: ica, Brazil Nycteliini is an endemic Neotropical tribe of Pimeliinae (Doyen 1993), with 285 species distributed in Argentina, Chile, Bo- livia, Peru, Paraguay, and Uruguay (Fig. 1) (Flores 1997). The species of Nycteliini are currently arranged in 11 genera, the nine recognized in previous revisions: Gyrioso- mus Guérin-Méneville, Pilobalia Burmeis- ter, Entomoderes Solier, Nyctelia Latreille, Epipedonota Solier, Psectrascelis Solier, Scelidospecta Kulzer, Auladera Solier, Mi- tragenius Solier (Kulzer 1954, Flores 1997), the recently described genus Pata- gonogenius (Flores 1999), and the restored genus Callyntra Solier (Flores and Vidal 2000). Our examination of a single specimen collected in Brazil by W. Mann from the Field Museum of Natural History (Chicago, Tenebrionidae, Pimeliinae, Asidini, Nyctelini, Entomobalia, South Amer- USA) led us to think we had discovered a new species of Nycteliini. More recently we discovered a long series of 80 specimens of this species and another series of 25 spec- imens of another smaller species, both col- lected in northeastern Brazil by W. Mann, deposited in the National Museum of Nat- ural History, Smithsonian Institution, (Washington, DC). One specimen of the longer series had been determined by Kulz- er as Scotinus platynotos (Perty). After re- questing a loan of all the species of Scotin- us present in The Natural History Museum (London, UK), we confirmed that both spe- cies had already been described by Perty (1830), the larger as Asida platynotos and the smaller as Asida picta. Finally, study of the types of Perty from the Zoologische Staatssammlung, Miinich (Germany) con- VOLUME 104, NUMBER 3 firmed the identity of both series of these species. When Perty (1830) described these two species, he placed them in the genus Asida Latreille (Asidini). Laporte (1840) consid- ered that they should be included in the ge- nus Scotinus Kirby (Asidini). Scotinus picta had not been mentioned in the literature for 160 years, but in 1935, Blair expressed the opinion that S. platynotos “‘is a somewhat aberrant Pilobalia’. Gebien (1910) listed this species under Scotinus, but later he list- ed it under the nycteliine genus Pilobalia (Gebien 1937). This means that both au- thors believed this species belongs to the tribe Nycteliini rather than the Asidini. However, Kulzer (1954) still accepted this species as a member of the genus Scotinus. Through a detailed discussion of the char- acters, we demonstrate that Scotinus platyn- otos and S. picta are not congeneric with the remaining species of Scotinus, that they are not Asidini, that they share most of these characters with all or some genera of Nycteliini, and that they deserve recogni- tion as a separate genus, which we have named Entomobalia. This new genus exhib- its a mosaic of characters present in other genera of Nycteliini, especially Pilobalia and Entomoderes. The inclusion of these two species in any other known genus of Nycteliini would imply a completely differ- ent concept and redefinition of that genus. In addition, the male and female genitalia show unique apomorphies. Species of Nycteliini inhabit arid and semiarid environments, mainly in the bio- geographical provinces Chaco, Monte, Pampa, Central and Arid Puna, Prepuna, Coquimbo, Santiago, Payunia, Western, Central and Fueguinan Patagonia, and San Jorge Gulf (Morrone 2001). The discovery of these two species from northeastern Bra- zil, at least 3,000 km from the Chaco, the closest point of distribution of a genus of Nycteliini (Entomoderes), was unexpected. Thus the range of the tribe is greatly ex- panded to the Caatinga biogeographical province (Morrone 2001). It is possible that 603 other species of Nycteliini may be found in the Cerrado biogeographical province, Which lies between Caatinga and Chaco Gries al): The objectives of this study are to de- scribe this new genus of Nycteliini, rede- scribe the two species assigned to the new genus, and compare the distinctive charac- ter states of the species of Scotinus (Asi- dini) with the two species assigned to En- tomobalia (Nycteliini), which had previous- ly been placed in Scotinus. Specimens were obtained from the fol- lowing institutions: The Natural History Museum, London, UK (BMNH), Field Museum of Natural History, Chicago, IL, USA (FMNH), National Museum of Nat- ural History, Smithsonian Institution, Washington, DC, USA (USNM), and Zool- ogische Staatssammlung, Miinich, Germa- ny (ZSMC). Thanks to the kind generosity of the USNM, we distributed some speci- mens of both species in the following in- stitutions: California Academy of Scienc- es, San ‘Francisco, GA,. USA (CASE), Field Museum of Natural History, Chica- go, IL, USA (FMNH), Instituto Argentino de Investigaciones de las Zonas Aridas, Mendoza, Argentina (IADIZA), Museo Argentino de Ciencias Naturales Bernar- dino Rivadavia, Buenos Aires, Argentina (MACN), Museu de Zoologia da Univer- sidade de Sao Paulo, Sao Paulo, Brazil (MZSP), and The Ohio State University, Columbus, OH, USA (OSUC). Body length was measured dorsally, along the midline, from anterior margin of labrum to elytral apex. For paraproct/coxite length (P/C) we used the ratio proposed by Doyen (1993). For basal lamina of the teg- men/lateral styles length (B/E) and median lobe/tegmen length (L/T) we used the ratios proposed by Flores (1996). Drawings were made with a camera lucida adapted to a ste- reoscopic microscope. TAXONOMIC PLACEMENT OF ENTOMOBALIA The two species that we have assigned to the new genus Entomobalia (Nycteliini) 604 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs 1 Known distribution of the genera of Nycteliini (lined area). Black square: distribution records for Entomobalia platynota; black circle: distribution records for E. picta. | and 2: Caatinga and Cerrado biogeo- graphical provinces (sensu Morrone 2001). had previously been placed in the genus Scotinus, which belongs to the Asidini, a large tribe of Pimeliinae with more than 1,000 species distributed in North and South America, Africa south of Sahara, Madagascar, and the Mediterranean area (Koch 1955). The species of Scotinus are endemic to Brazil (Gebien 1937). We have studied the following characters within the species of Scotinus, some of which have been used to study the relationships be- tween the tribes of Pimeliinae (Doyen 1993) and to define the tribe Nycteliini (Flores 1997), finding that Scotinus platyn- otos and S. picta share all of these charac- ters with all or some genera of Nycteliini and not with the remaining species of Sco- tinus. The species of Scotinus examined from BMNH are the following (T includes type specimens): S. crenicollis Kirby (T) (type species), S. crucifer Eschscholtz, S. propius Wilke, S. quadricollis Eschscholtz, VOLUME 104, NUMBER 3 S. quadricollis incisus Wilke (T), S. tuber- culatus Eschscholtz, S. tuberculatus dispar Wilke (T), S. chapadensis Wilke (T), and S. bajulus Wilke (T). fe i) Size of antennomere I1: Very small, even rudimentary in Scotinus (Figs. 2— 3); in Entomobalia and Nycteliini well developed, sometimes equal to or longer than antennomere 10 (Figs. 4—7). Subgenal process (defined by Doyen 1993: 454): Subadjacent with mentum in Scotinus (Solier 1836: plate XIII, Figs. 1-2, Doyen 1993: Figs. 22—25); in En- tomobalia and Nycteliini it is remote from mentum (Doyen 1993: Fig. 21). Position of ligula: Nearly concealed be- neath mentum in Scotinus (Solier 1836: plate XIII, Figs. 1-2); in Entomobalia it is entirely exposed anterad of mentum (Fig. 8), with the articulating membrane visible. Within Nycteliini, this character state is shared with Gyriosomus, Pilob- alia and Entomoderes. Relative size of ligula: Less than half the size of mentum in Scotinus (Solier 1836: plate XII, Figs. 1-2); in Entomobalia it is larger than half of mentum (Fig. 8). Within Nycteliini, this character state is shared with Gyriosomus, Pilobalia and Entomoderes. Shape of mentum: Cordiform in Scotin- us, With anterior margin twice as long as posterior margin (Solier 1836: plate XIII, Figs. 1-2). Subtrapezoidal in En- tomobalia, with anterior margin 1.5 times the length of posterior margin (Fig. 8). Within Nycteliini, this character state is shared only with Pilobalia (Fig. 10). Mesepisternum: Small, not reaching the mesocoxal cavities in Scotinus (Fig. 11); in Entomobalia and Nycteliini it is large, reaching the mesocoxal cavities and closing laterally (Fig. 12). Mesepimeron: Long, reaching the base of elytral epipleuron in Scotinus (Fig. 11); in Entomobalia and Nycteliini it is short, not reaching the base of elytral epipleuron (Fig. 12). 9. 605 Metepimeron: Transverse in Scotinus, widely separating the metacoxal cavities from the elytral epipleuron (Fig. 11); in Entomobalia and Nycteliini it is not ev- ident, since the metacoxal cavities are very close to the elytral epipleuron (Fig. (2). Lateral Closed laterally by metepimeron and first visible abdominal sternum in Sco- tinus (Fig. 11); closed laterally by me- tepisternum in Entomobalia (Fig. 12). Within Nycteliini, this character state is closure of metacoxal cavities: shared with Gyriosomus, Pilobalia and Entomoderes. 10. Distance between meso- and metacox- ll. ae: Does not exceed half the metacoxal length in Scotinus (Fig. 11); in Ento- mobalia and Nycteliini it exceeds half the metacoxal length (Fig. 12). Abdominal margin of elytral epipleu- ron: Curved where it contacts meso- and metathorax in Scotinus (Fig. 11); in En- tomobalia and Nycteliini it is straight where it contacts meso- and metathorax (Rise 12): Additional character states which differ in Scotinus and Entomobalia: ha i) oS) Antennal length: Short in Scotinus, not reaching the middle of the lateral margin of pronotum; longer in Entomobalia, ex- tending beyond posterior margin of pronotum. Shape and width of antennomere 10: Oval, wider than long in Scotinus, wider than more proximal antennomeres (Figs. 2—3); subrectangular, longer than wide in Entomobalia, no wider than more prox- imal antennomeres (Figs. 4—5). Ventral femoral surface: Covered by sparse setae in Scotinus; densely setose on proximal %4 in Entomobalia. Elytral epipleuron: Not evident in Sco- tinus (Fig. 11); evident throughout in Entomobalia (Fig. 12). In addition, in Entomobalia the elytral epipleuron is equal in width throughout (Fig. 12); within Nycteliini this character state is 606 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-12. External structure. 2-7, Antennae in dorsal view. 2, Scotinus crenicollis. 3, S. crucifer. 4, En- tomobalia platynota. 5, E. picta. 6, Pilobalia decorata (Erichson). 7, P. oblonga (Blanchard). 8—10, Labium and mentum in ventral view. 8, Labium of Entomobalia platynota. 9, Mentum of Entomoderes satanicus Waterhouse. 10, Mentum of Pilobalia decorata. 11-12, Mesothorax, metathorax and abdomen in ventral view. 11, Scotinus quadricollis. 12, Entomobalia picta (female). Abbreviations: 1, ligula, m, mentum, p, labial palpus, mes, mese- pisternum, mep, mesepimeron, met, metepisternum, mtp, metepimeron. Scale bar = | mm. VOLUME 104, NUMBER 3 shared with Gyriosomus, Pilobalia and Entomoderes. Nn Sexual dimorphism: Species of Scotinus do not exhibit sexual dimorphism; spe- cies of Entomobalia exhibit four char- acters different in male and female (see below). In his cladistic analysis of Pimeliinae, Doyen (1993) found that all genera of As- idini (from South African, Madagascan, Mediterranean, North and South American regions) constitute a monophyletic group defined by six synapomorphies, five of which are not present in bridge of tentorium absent or incomplete, Entomobalia: antennae with ten segments plus the re- duced eleventh antennomere, abdominal la- terotergites extremely small, apicodorsal lobe of proctiger ending at coxite base, and baculus of proctiger extending proximad, equal to baculus of paraproct. Furthermore Doyen (1993) pointed out that the first two are synapomorphies unique to Asidini with- in the subfamily Pimeliinae. Within Pimeliinae, Entomobalia must be placed within the Asidine clade (Doyen 1993) by having multiple, long, slender spermathecal tubes which open as a fascicle into the base of the accesory gland duct or into the vagina near the duct (Figs. 17—18). Within the Asidine clade of Doyen (1993), Entomobalia belongs to the subclade of South American tribes Nycteliini, Physo- gasterini and Praocini by having metendos- ternite arms fused with mesocoxal inflex- ions. Entomobalia is placed in the tribe Nycteliini according to the definition of that tribe by Flores (1996, 1997) and the follow- ing change in that tribal concept should be mentioned: in the female genitalia of En- tomobalia the basal lobe of the coxite is separated vertically from apical lobe (Figs. 17-18), while in the remaining genera of Nycteliini it is separated horizontally from apical lobe (Flores 1996). The most recent key provided for the genera of Nycteliini is that by Flores 607 (1997), which is modified at couplet 8 to separate out Pilobalia and Entomobalia: 8. Pronotum with a short central-posterior carina and two longer longitudinal carinae, lateral margin with lobe (Flores 1997: Fig. 25); fem- ora with umbilicate setae; mentum transverse (Fig. 9) Pronotum lacking carinae, lateral margin con- Meg eat ca OO, O NC Entomoderes Solier cave, lacking lobe; femora with simple setae; mentum subtrapezoidal (Figs. 8, 10) ..... 8a 8a. Mentum impunctate or with very small punc- tures; antennomere 10 subspherical or pyri- form (Figs. 6-7); male with protibiae straight and not expanded apically (Flores 1997: Fig. ZG)! + ia coves Beh ee ae — Mentum with large punctures; antennomere Pilobalia Burmeister 10 subrectangular (Figs. 4—5); male with pro- tibiae curved inward and expanded apically (Figs: 19=20)) 2%. See Entomobalia, new genus Entomobalia Flores and Triplehorn, new genus Type species.—Asida platynotos Perty, present designation. Diagnosis.—Distinguished from other Nycteliini by the following combination of characters: ligula sclerotized, ventrally ex- posed, equal to half mentum area; labial palps inserted on posterior half of ligula; mentum subtrapezoidal; pronotum smooth, without punctures, striae or carinae, lateral margin concave, posterior angles acute, overlapping elytral humeri, epipleuron equal in width throughout; metacoxal cav- ities closed laterally by metepisternum; ventral surface of femora densely setose; median lobe of aedeagus subapically ex- panded and strongly sclerotized on distal third; sexual dimorphism: male with proti- bia curved inward and expanded apically, inner surfaces of tibiae densely setose on distal half, and central area of metasternum with setae; female with protibia straight and not expanded apically, inner surfaces of tib- iae with normal setae, and central area of metasternum glabrous and shiny. Description.—Length, 12.1—-19.9 mm; width, 6.4—11.3 mm. Body and legs brown to black, with antenna and maxillary palp brown. Pronotum and elytron with two 608 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 13-18. Internal skeletal anatomy and female genitalia of Entomobalia spp. 13-14, Tentoria. 13, En- tomobalia platynota. 14, E. picta. 15—16, Metendosternites in posterior view. 15, Entomobalia platynota. 16, E. picta. 17-18, Ovipositor (ventral view), spiculum and internal female reproductive tract. 17, Entomobalia pla- tynota. 18, E. picta. Abbreviations: c, coxite, 0, oviduct, p, paraproct, s, spermatheca, sag, spermathecal accessory gland, sp, spiculum, v, vagina. Scale bar = | mm. kinds of short setae, one stout, dark brown, and other finer, golden or light brown. Head: Epipharynx with anterior margin entirely concealed beneath labrum. Labrum, clypeus and frons with abundant short, golden setae. Clypeus with anterior margin concave. Clypeal suture defined by a deep depression with setae at antennal insertion level. Frons without longitudinal or lateral grooves. Eye reniform. Antenna long, ex- tending beyond posterior margin of prono- tum; antennomeres subrectangular, with two different kinds of pubescence: one short and abundant on entire surface, and second consisting of long, scattered setae; unique apical semicircular tomentose sen- VOLUME 104, NUMBER 3 Figs. 19, 20. sory patch on antennomere 9. Ligula scler- otized, articulated with mentum by a nar- row membrane and exposed ventrally, equal to half mentum area (Fig. 8); labial palpi inserted on posterior half of ligula (Fig. 8). Ligula and mentum with setae arising from large punctures. Mentum subtrapezoidal (Fig. 8). Submentum posteriorly continuous with gula. Tentorium with medial straight bridge (Figs. 13-14). Thorax: Pronotum short (length: prono- tum/elytron = 0.33), pubescent and smooth, without punctures, striae or carinae; disc convex; anterior margin slender and central area not broadened; anterior angles round- ed; lateral margin simple, slender, concave, widest behind mid point; posterior angles acute, overlapping elytral humeri; posterior margin biconcave, as wide as base of elytra. Proepisternum with or without grooves and with sparse not umbilicate setae. Proster- num arched, not extended over mesoster- num. Mesosternum inclined forward, sepa- rated from prosternum. Scutellum visible. Meso- and metepisternum without grooves. 609 Habitus in dorsal view. 19, Entomobalia platynota, male. 20, E. picta, male. Elytron: Dorsal surface, pseudopleuron and epipleuron impunctate; with one or two carinae, space between external carinae and lateral margin without grooves; lateral mar- gin straight and sharp, epipleuron equal in width throughout, texture similar to that of elytra. Legs: Procoxal cavity closed posteriorly. Metacoxae separated by one metacoxal width, enclosed laterally by metepisternum. Ventral surfaces of trochanters pubescent, brushlike. Ventral surfaces of femora dense- ly setose on proximal %4. Ventral surfaces of tarsi bearing abundant decumbent setae. Sexual dimorphism: Male with protibia curved inward and expanded apically, inner surfaces of tibiae densely setose on distal half, and central area of metasternum with setae. Female with protibia straight and not expanded apically, inner surfaces of tibiae with normal setae, and central area of me- tasternum glabrous and shiny. Internal skeletal anatomy: Mesendoster- nite with a long and slender dorsal arm, lon- ger than horizontal arm. Metendosternite 610 (Figs. 15—16) with arms long, extending be- yond mesocoxal inflections about half dis- tance to tergum, stem equal to metacoxal width, width of stem exceeding length, and stem narrow at middle. Elytral-abdominal fusion accomplished by a ridge in the ely- tral epipleuron which interlocks in a lon- gitudinal groove of abdominal sterna. Male genitalia: Rods of sternum IX close at basal third, distance between them not exceeding width of aedeagus. Dorsal membrane of proctiger concave, with two sclerotized areas. Basal lamina of tegmen long (B/E > 1.00). Lateral styles of tegmen distally close, with apex straight and long setae on ventral surface; ventral proximal margin convex in ventral view, projecting dorsally over median lobe. Median lobe long (L/T > 1.00), sheath-shaped, one third the width of lateral styles of tegmen, ex- panded subapically and strongly sclerotized on distal third, with apex acute and straight. Female genitalia (Figs. 17-18): Spicu- lum with arms “V-shaped. Paraprocts moderate (1,2 = P/C = 2,0), glabrous. Cox- ites with setae, basal lobe of coxite extend- ed over paraproct and separated vertically from apical lobe, baculi of coxite inclined 45°: midventral sclerite distally broadened. Proctigeral baculus equal to length of par- aproct baculus. Vagina saccate; spermathe- cal tubes shorter than vagina length, all similar in width and branching pattern: spermathecal accesory gland longer than vagina, with duct annulate and thick. Etymology.—The name of the genus re- fers to the similarity to the Nycteliine gen- era Entomoderes and Pilobalia. Gender feminine. Distribution and habitat.—Brazil: States of Ceara, Rio Grande do Norte, Pernam- buco and Bahia, in the Caatinga biogeo- graphical province (Morrone 2001) (Fig. 1). W. Mann, the collector of the large series of both species in Baixa Verde, Rio Grande do Norte, stated that “the country was arid, with much scrub and cacti, but little life in evidence” (Mann 1948: 91). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON KEY TO THE SPECIES OF ENTOMOBALIA 1. Pronotum with lateral margin reflexed; elytron with lateral margin single and protuberances ir- regularly and sparsely distributed on dorsal surface; length usually greater than 15 mm (Fig. 19) E. platynota (Perty) — Pronotum with lateral margin not reflexed; el- ytron with lateral margin double and lacking protuberances; length usually less than 13 mm (Fig. 20) E. picta (Perty) Entomobalia platynota (Perty), new combination (Piessaeaes., 135 ise eZ) Asida platynotos Perty 1830: 56, plate 12, fig. 2. Scotinus platynotos: Laporte 1840: 208; La- cordaire 1859: 165. Scotinus platynotus: Gemminger and Har- old 1870: 1880 (cat.); Gebien 1910: 139 (cat.). Pilobalia platynota: Blair 1935: 104; Ge- bien 1937: 753 (cat.); Blackwelder 1945: 519 (cat.); Kulzer 1954: 254 (rev.); Flo- res 1997: 16 (ist). Scotinus platynotos: Kulzer 1954: 265. Redescription.—Length, 15.0-19.9 mm; width, 8.0—-11.3 mm. Body and femora brown to black, with antenna, tibia and tarsi brown. Stout dark brown setae and golden or light-brown finer setae uniformly and sparsely distributed on pronotum and dorsal surface of elytron, both forming velvet-like patches only on posterior third of elytron. Antenna long, extending three antenno- meres beyond posterior margin of prono- tum. Pronotum with lateral margin reflexed. Male metasternum with tuft of setae on cen- tral area, brush-like. Metepisternum with not umbilicate setae. Elytron with lateral margin single and protuberances irregularly and sparsely distributed on dorsal surface, with one feebly raised complete or incom- plete carina, close to lateral margin (Fig. 19): in some specimens with a secondary carina close to suture, consisting of aligned protuberances. Male with protibia expanded apically, but not excavated. Last abdominal segment truncate in males and females. In addition to three characters of sexual di- VOLUME 104, NUMBER 3 611 Figs. 21-24. Male genitalia of Entomobalia spp. in dorsal and ventral views. 21, 22, Entomobalia platynota. 23, 24, E. picta. Abbreviations: bl, basal lamina of tegmen, Is, lateral styles of tegmen, ml, median lobe. Scale bar = | mm. morphism at generic level, first abdominal sternum with a tuft of brush-like setae on central area in males and glabrous and shiny in females. Male genitalia.—Lateral styles of tegmen with apex wide, widest at mid point, with long setae on distal half of ventral surface (Fig. 21). Median lobe with apical aperture large, strongly expanded subapically (Fig. D2). Specimens examined.—Lectotype: [Type] [Brasilia/Scotinus/platynotos/Perty | (ZSMC). To fix the current interpretation of this name and to ensure stability, we are hereby designating this lectotype: [Lectoty- pus/Asida_platynotos/Perty, 1830/Des. G. Flores-/C. Triplehorn 2001]. Non-type spec- imens. BRAZIL: Rio Grande do Norte. Baixa Verde, W. Mann, 75 [55 (USNM), 3 (CASC), 4 (FMNH), 7 CUADIZA), 2 (MAECN), .2)'\(MZSP):'2 (OSUC)]? Ceara- Mirim, W. Mann, | (USNM). Pernambuco. Ouricuri,' ‘TW=1982,"E.C 9 "Gux (ex Bufo stomach), 3 (OSUC). Bahia. Near Queima- das, 11-VI-1915, PG. Russell, 2 (USNM); S. Salvador, 1918, 2 (USNM); Barra, 1 (BMNH); without more precise data: 2 (BMNH). Entomobalia picta (Perty), new combination (Figs. ie, 127 A, MG 182205235 24) Asida picta Perty 1830: 56, plate 12, fig. 3. Scotinus picta: Laporte 1840: 208; Lacor- daire 1859: 165. Scotinus pictus: Gemminger and Harold [37021880 (cat);) Gebien 1910 139 612 (cat), 19375743) (cat.);, Blackwelder 1945: 517 (cat.). Redescription.—Length, 12.1-15.9 mm; width, 6.4—7.6 mm. Body and femora brown to black, with antenna, tibia and tarsi brown. Stout dark brown setae uniformly and sparsely distributed on pronotum and dorsal surface of elytron, grouped only on outer third of pronotum and elytron. Finer, golden or light brown setae grouped form- ing velvet-like patches on pronotum and dorsal surface of elytron. Antenna long, ex- tending two antennomeres beyond posterior margin of pronotum. Pronotum with lateral margin not reflexed. Male metasternum with setae on central area, not forming a brushlike tuft. Metepisternum with umbili- cate setae. Elytron with lateral margin dou- ble and without protuberances, with two low carinae, equidistant between suture and lateral margin (Fig. 20), visible only for the different pattern of setae. Male with proti- bia expanded apically and excavated in- ward. First abdominal sternum with short setae in male and female. In addition to three characters of sexual dimorphism at generic level, last abdominal segment trun- cate in males and emarginate in females (Figwi2): Male genitalia.—Lateral styles of tegmen with apex narrow, widest at base, with long setae on distal quarter of ventral surface (Fig. 23). Median lobe with apical aperture small, moderately expanded subapically (Fig. 24). Specimens examined.—Lectotype and three paralectotypes without label data (ZSMC). One paralectotype: [Brasilia/Sco- tinus/pictus/Perty] (ZSMC). To fix the cur- rent interpretation of this name and to en- sure stability, we are hereby designating this lectotype and four paralectotypes: [Lec- totypus/Asida picta/Perty, 1830/Des. G. Flores-/C. Triplehorn 2001], the same for the four paralectotypes. Non-type speci- mens. BRAZIL: Rio Grande do Norte. Baixa Verde, W. Mann, 24 [13 (USNM), 2 (CASO) 2 9EMNE) 562, (ADIZA);! 1 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (MACN), 2 (MZSP), 2 (OSUC)]; Natal, W. Mann, | (USNM). Ceara. FE da Rocha coll., 13-IH-1940, 1 (USNM); E da Rocha coll., 2 (USNM). Pernambuco. Pesqueira, V- 1935, L. Castro, 4 (USNM). Bahia. Near Queimadas, 10-VI-1915, PG. Russell, 2 (USNM); without more precise data: | (ZSMC). ACKNOWLEDGMENTS We gratefully acknowledge Maxwell V. L. Barclay (BMNH), Philip P. Parrillo (FMNH), Warren E. Steiner (USNM), and Martin Baehr (ZSMC) for loaning the spec- imens, Peter W. Kovarik for taking the pho- tographs, Claudia Vergara for inking the drawings, Warren E. Steiner for suggestions on improving this paper and for sending copies of the papers of Perty and Solier, and the valuable comments of an anonymous reviewer. This study was supported by the Department of Entomology, The Ohio State University, and by the Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET), Argentina. LITERATURE CITED Blackwelder, R. E. 1945. Checklist of the coleopterous insects of Mexico, Central America, the West In- dies and South America. Part 3. Bulletin of the United States National Museum 185: 343-550. Blair, K. G. 1935. Some synonymic notes in the family Tenebrionidae (Col.). Entomologist’s Monthly Magazine 71: 102—104. Doyen, J. T. 1993. Cladistic relationships among Pi- meliine Tenebrionidae (Coleoptera). Journal of New York Entomological Society 101(4): 443— 514. Flores, G. E. 1996. Estudio comparativo de las estruc- turas genitales en la tribu Nycteliini (Coleoptera: Tenebrionidae). Revista de la Sociedad Entomo- logica Argentina 55: 33-48. . 1997. Revision de la tribu Nycteliini (Cole- optera: Tenebrionidae). Revista de la Sociedad Entomolégica Argentina 56: 1—19. . 1999. Systematic revision and cladistic anal- ysis of the Neotropical genera Mitragenius Solier, Auladera Solier, and Patagonogenius gen. n. (Co- leoptera: Tenebrionidae). Entomologica Scandi- navica 30(4): 361-396. Flores, G. E. and P. Vidal. 2000. Revalidation and sys- tematic revision of the Chilean genus Callyntra Solier (Coleoptera: Tenebrionidae). Annals of the VOLUME 104, NUMBER 3 Entomological Society of America 93(5): 1052 1075. Gebien, H. 1910. Tenebrionidae. I: 1-166; Il: 167-354. Coleopterorum Catalogus 18 pts. 15, 22. Berlin. . 1937. Katalog der Tenebrioniden. Teil I. Pub- blicazioni del Museo Entomologico Pietro Rossi. Duino 2: 505-883. Gemminger, M. and E. von Harold. 1870. Catalogus Coleopterorum hujusque descriptorum synony- miscus et systematicus, Monachii. Vol. 7: 1801— 2180. Koch, C. 1955. Monograph of the Tenebrionidae of Southern Africa, Vol. I. Tentyriinae, Molurini.- Trachynotina: Somaticus Hope. Transvaal Muse- um, Pretoria, xi + 242 pp. Kulzer, H. 1954. Neunter Beitrag zur Kenntnis der Te- nebrioniden (Col.) Eine Studie tiber die Tribus Nycteliini. Entomologische Arbeiten dem Muse- um Georg Frey 5(1): 145-267. 613 Lacordaire, J. T. 1859. Histoire Naturelle des Insectes. Genera des Coléopteres, Vol 5. Paris, 750 pp. Laporte, F L. N. (Comte de Castelnau). 1840. Histoire Naturelle des Insectes, Vol. 2. Paris, 564 pp. Mann, W. M. 1948. Ant Hill Odyssey. Atlantic Month- ly Press Book, Boston. Morrone, J. J. 2001. Biogeografia de América del La- tina y el Caribe. Manuales & Tesis Sociedad En- tomologica Aragonesa, Vol. 3, Zaragoza, Spain, 148 pp. Perty, M. malium Articulatorum, quae in itinere per Brasi- liam annis MDCCCXVII-MDCCCXX jussu et auspiciis Maximiliani Josephi I. Fasc. 1, pp. 1—60, 1830. Insecta Brasiliensia. /n Delectus Ani- illus. Solier, A. J. J. 1836. Essai sur les Collaptérides. 6e Tribu. Asidites. Annales de la Société Entomolo- gique de France 5: 403-512. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 614-623 A NEW SPECIES AND KEY FOR THE GENUS ZONOSEMATA BENJAMIN (DIPTERA: TEPHRITIDAE) ALLEN L. NORRBOM Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: anorrbom @sel.barc.usda.gov) Abstract.—New taxonomic, host, and distribution data for species of Zonosemata are presented. One new species, Z. guybushi, reared from fruit of Solanum lanceifolium in Costa Rica, is described, and a revised key to the eight known species of this genus is provided. A neotype is designated for Z. electa (Say). Key Words: This paper includes the first results from a project on the natural history and system- atics of the fruit flies (Tephritidae) of Costa Rica. In this project, funded by the Re- search and Scientific Exchanges Division, Foreign Agricultural Service, USDA, I have collaborated with parataxonomists from the Instituto de Biodiversidad of Costa Rica, who are collecting and rearing fruit flles. Among the many new species and host records resulting from this project was an undescribed species of Zonosemata Ben- jamin reared by Freddy Quesada in Guan- acaste. In addition to describing this spe- cies, I take this opportunity to provide a revised key to the species of Zonosemata, and to report additional taxonomic, host, and geographic data for other species. Zonosemata includes eight species that are native to the New World (from south- eastern Canada to Colombia and northeast- ern Brazil). The seven previously known species were cataloged by Norrbom et al. (1999: 249), and Smith and Bush (1999: 197) listed their known host plants. All but one of the species have been reared from fruits or at least associated as adults with species of Solanaceae, mainly in the genus Zonosemata, fruit flies, taxonomy, host plant, distribution, Solanum Solanum. One species, Z. electa (Say), has been reared from genera other than Sola- num, and is considered a pest of pepper and eggplant in the eastern United States and Canada. Zonosemata belongs to the subtribe Car- pomyina of the subfamily Trypetinae (Smith and Bush 1999). Jenkins (1996) questioned whether the apical desclerotized area of the female oviscape (syntergoster- nite 7), considered a synapomorphy for the Carpomyina by Norrbom (1989), is present in Zonosemata. The genitalia of the speci- mens from the USNM that he dissected are overcleared and this character can no longer be seen in them, but it is apparent in less cleared dissections. MATERIALS AND METHODS Morphological terminology follows White et al. (1999). Figure 1I shows the names used for wing bands. Acronyms for institutions housing the examined speci- mens follow Thompson (1999). KEY TO SPECIES OF ZONOSEMATA In previous keys for Zonosemata (Bush 1966, Hernandez-Ortiz 1989), the dark VOLUME 104, NUMBER 3 brown (often referred to as “*black”’) mark- ings of the thorax (see Figs. 2 and 3) were important characters in the early couplets. In the following key I have de-emphasized their importance because they appear to vary more than was previously known. For example, the katepisternal spot is consis- tently present in Z. vittigera and Z. mac- gregori, but varies in Z. scutellata, Z. vi- drapennis (present in Central American specimens), and probably Z. minuta (pre- sent but very small in the examined speci- mens). It is even rarely present in Z. electa (in 4 specimens of the more than 100 ex- amined from the USNM collection). The presutural scutal spot is present in Z. vittig- era, Z. macgregori, and Z. minuta (although it is Sometimes just a linear mark in the latter), but it is variable in Z. scutellata. The postsutural brown markings vary consider- ably, for example, in Z. electa from absent to very large (compare Figs. 2A and B). 1. Wing bands extremely narrow; width of discal, subapical and apical bands less than length of crossvein R-M (Fig. 1H). Most of radial cells and cell dm between discal and subapical bands without microtrichia. Scutum without presutural dark brown mark (Fig. 3C). Mexico, Guatemala, Honduras ....... vidrapennis Bush — Wing bands relatively broad; width of discal and subapical bands equal to or greater than length of R-M (Figs. 1A—G, I). Radial cells and cell dm at most with small areas bare of mi- crotrichia between these bands, or if with large bare areas (Z. vittigera, Z. macgregori), scutum with large presutural, sublateral dark brown Marke (BISS SB MD) heels cha eo ae Dw is a ys Apical band extended basad along vein M, to- gether with subapical band forming P-shaped mark (Fig. 1A). Mexico (Morelos, Guerrero, Chiapas) cocoyoc Bush — Apical band not extended along veinM .... 3 3. Distal 3 abdominal tergites (3—5 in male, 4—6 in female) with dark brown lateral spots, those minuta Bush No i) on last 2 tergites large. Jamaica . — Dark brown lateral spots present at most on distal 2 tergites, that on penultimate tergite small if present 4. Apical band, subapical band anterior to vein R,,.5, and accessory costal band faint, much paler than discal band and posterior part of subapical band (Fig. 1G). Accessory costal band often connected to subapical band. 615 Length of first flagellomere of antenna (mea- sured on mesal side) more than 0.7 times height of face (measured to ventral margin of antennal sockets). Colombia, Venezuela, north- €astermnsBraziliees cee war scutellata (Hendel) — Apical and subapical bands evenly developed, similar to discal band in color (Figs. 1B—E, I). Accessory costal band variable, but not con- nected to subapical band. Length of first fla- gellomere less than 0.7 times height of face .. 5 5. Scutum without presutural dark brown marks (Fig. 2). Cells r,,; and dm mostly or entirely microtrichose between discal and = subapical bands. Katepisternum rarely with dark brown ving ey icgpcaaoe von SRM tc cc cs 6 — Scutum with presutural dark brown marks (Figs. 3B, D). Cells r,,; and dm with large bare spot areas without microtrichia between discal and subapical bands. Katepisternum with dark brown spot 6. Discal band (measured along vein M) usually narrower than hyaline area distal to it, at most 1.1 times as broad (Figs. 1B—C). Discal, sub- apical and apical bands orange brown. Subscu- tellum and mediotergite usually orange. South- eastern! Canada-;,eastemiUtS. = see electa (Say) — Discal band (measured along vein M) 1.21— 1.34 times broader than hyaline area distal to it (Fig. 1D). Discal, subapical and apical bands brown. Subscutellum and most of mediotergite dark: browns GostamRicates <2)... ssebueye chee = arg dts Gan BBE ae enatera yg guybushi Norrbom, n. sp. 7. Wing bands relatively narrow, cell r,,, apical to subapical band with large hyaline mark along vein R,,;, at least % width of cell (Fig. 11). Vertex usually entirely yellow except for brown ocellar tubercle. Scutum with submedial stripe usually orange anterior to transverse su- ture (Fig. 3D), occasionally partially to entirely red-brown or dark brown. Accessory costal band distinct. Southwestern U.S. to central Mexico Ssh Boe. 2. Se ae vittigera (Coquillett) — Wing bands relatively broad, cell r,,, apical to subapical band entirely infuscated or with nar- row hyaline mark along vein R,,;, at most % width of cell (Fig. 1E). Orbital plate moderate to dark brown, sometimes connected by a band across vertex. Scutum with submedial stripe usually dark brown anterior to transverse su- ture (Fig. 3B). Accessory costal band some- times faint. Mexico (Baja California, Sonora) PL: Cb CS oo See macgregori Hernandez-Ortiz Zonosemata cocoyoc Bush (Fig. 1A) References.—Bush 1966: 319 [descrip- tion]; Aluja et al. 1987: 324 [Chiapas]; Norr- bom 1990: 55 [host]. 616 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON subapical Fig. 1. Wings. A, Zonosemata cocoyoc. B-C, Z. electa. D, Z. guybushi. E, Z. macgregori. F, Z. minuta. G, Z. scutellata. H, Z. vidrapennis. 1, Z. vittigera. VOLUME 104, NUMBER 3 Recognition.—The P-shaped apical and subapical bands distinguish Z. cocoyoc from all other species of Zonosemata. Biology.—An undetermined species of Solanum was listed by Norrbom (1990) as a host plant. Distribution.—Known only from Mexico (Chiapas, Guerrero, and Morelos). The fol- lowing specimen is the first reported from Guerrero. Specimen data. —MEXICO: Guerrero: Cacahuamilpa, 2.1 mi. NW of, 23 Jul 1981, Bogar, Schaffner & Friedlander, 19 (TAMU, USNM52777). Zonosemata electa (Say) (Figs. 1B—C, 2A—B) References.—Say 1830: 185 [descrip- tion]: Bush 1966: 314 [revision]. See Foote et al. (1993: 494) for additional information and references. Type data.—Say (1830) described this species as 7rypeta electa based on an un- stated number of specimens from Indiana. The syntypes, like the rest of Say’s Diptera types, have been lost (Stone 1980: 35). To clarify the concept of this species and pre- vent its confusion with the similar species Z. guybushi (see above key for diagnostic characters, and Benjamin 1934, Bush 1966 and Foote et al. 1993 for descriptions of Z. electa), a male in the National Museum of Natural History (USNM) is here designated as neotype. It bears the following labels: “USA: MARYLAND: Montgomery Co., Plummer’s Island, 11.VIII.1988, A. L. Norr- bom, on Solanum carolinense L.”, ““USNM ENT 00052480”’, [red] ““NEOTYPE ¢ Try- peta electa Say, 1830 desig. Norrbom 2001°’, and “‘Zonosemata electa (Say) det. Norrbom 1988”’. Zonosemata guybushi Norrbom, new species (Figs. 1D; ZC) Recognition.—This species runs to Z. electa in the key of Bush (1966). It differs in having broader and darker wing bands (the discal band is broader than the hyaline 617 area between it and the subapical band mea- sured along vein M). Cell cu, lacks or has only a very small nonmicrotrichose area, and the subscutellum and at least a broad medial stripe on the mediotergite are dark brown; in Z. electa these characters are var- lable, but the subscutellum and medioter- gite are usually entirely orange and there is usually a large nonmicrotrichose area in cell CU: Description.—Head: Frons with 3—4 frontal and 2 orbital setae; ocellar seta well developed. First flagellomere 0.53-—0.57 mm long, 0.61—0.63 times facial height, with distinct dorsoapical point. Arista with minute pubescense (sparse in holotype). Thorax: Mesonotum 3.14—3.20 mm long. Dorsocentral seta aligned with postalar seta. Wing (Fig. 1D): 6.17—6.58 mm long. Pat- tern, as typical for genus, with following bands: subbasal, discal (covering crossvein R-M), subapical, and anterior apical bands. Bands brown, relatively broad. Discal band (measured on vein M) 1.21—1.34 times as broad as hyaline area distal to it. Discal and subapical bands broadly connected in cell cu, although both are fainter in that cell than more anteriorly. Wing mostly micro- trichose, with parts of basal and costal cells and most of hyaline areas of cells br and dm between subbasal and discal bands bare; cell cu, at most with narrow bare area aligned with that in cu,. Abdomen: Orange. Tergite 6 with large lateral dark brown spot. Oviscape orange, 1.66 mm long. Aculeus 1.96—2.04 mm long, tip similar to other spe- cies of Zonosemata, slender with minute apical notches. Type data.—Holotype—? (INBio, IN- BI0002153791), COSTA RICA: Guana- caste: Area Conservacion Guanacaste, San Cristobal, LN 317400 383400, 620 m, emerged 8 Apr 1998 from fruits of Solanum lanceifolium Jacq. coll. 10 Sep 1997, 97-F A. Quesada-134. Paratype, same data ex- cept emerged 7 May 1998, 12 (USNM, IN- BIO002 153688). Biology.—The type specimens were reared from Solanum lanceifolium Jacq., PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 618 oe << \/ y f a - > Te if ‘1YSNGAns “Z “+ ‘a[eulay pure seul ‘Vj2aJa YIpUIasoUOZ ‘g—y ‘sMIqeY [esiog ‘T “BI 619 VOLUME 104, NUMBER 3 ‘paasuna "7 ‘q ‘Stuuuadvapia "7 ‘DQ ‘Mosaisovul ‘7 * “DIDIJAINIS DIDUASOUOZ ‘yo csmiqey [esi0od ‘ \\l {! yh) ic ‘Sly 620 which occurs in the Greater and Lesser An- tilles and from Oaxaca, Mexico to Peru. Distribution.—Known only from Costa Rica. Etymology.—This species is named in honor of Guy L. Bush, who produced the latest comprehensive revision of Zonose- mata. Zonosemata macgregori Hernandez-Ortiz (Figs. 1E; 3B) References.—Hernandez-Ortiz 206 [description]. Recognition.—In general, Z. macgregori has the most extensive dark brown mark- ings of any species of Zonosemata (Fig. 3B), although it sometimes cannot be sep- arated from Z. vittigera by body color pat- tern alone. Both of these species have a large dark brown spot on the katepisternum and a large presutural sublateral brown mark on the scutum. In Z. macgregori, the submedial stripe is usually dark brown an- terior to the suture, whereas it is usually orange in Z. vittigera, although there is overlap in this character. The width of the apical band (cell r,,, apical to subapical band entirely infuscated or nearly so) dis- tinguishes Z. macgregori from Z. vittigera. The orbital plates are moderate to dark brown, sometimes connected by a band across the vertex, whereas the top of the head is usually entirely yellow except for the ocellar tubercle in Z. vittigera. Biology.—No host plant data are known. Distribution.—Known only from Mexico (Baja California Norte, northern Baja Cal- ifornia Sur, and Sonora). The first record from Sonora is reported below, based on a female that was compared with the type se- ries by Vicente Hernandez-Ortiz at my re- quest. Comments.—There is some intergrada- tion in the scutal color pattern with Z. vit- tigera. The submedial band is partly to mostly orange anterior to the transverse su- ture in 2 of 5 specimens of Z. macgregori examined, and in some specimens of Z. vit- tigera trom Mexico it is partially or entirely 1989: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON red brown or dark brown. Whether the sub- medial stripe and presutural lateral spot are connected (considered a diagnostic charac- ter for Z. macgregori by Hernandez-Ortiz) varies in both species, although the connec- tion is orange if present in Z. vittigera, whereas it may be brown in Z. macgregori. The difference in the width of the wing bands, especially the apical band, seems consistent, although this is based on a rather small sample of Z. macgregori. Study of larger samples of specimens or analysis us- ing other methods would be useful to con- firm that these populations are not conspe- cific with those of Z. vittigera. Discovery of their host plant would also be helpful in this regard. Solanum elaeagnifolium Cav., the only known host plant of Z. vittigera, occurs in Sonora and at least the extreme north of Baja California Norte (Wiggins 1980), but I do not know if it occurs in the other areas from which Z. macgregori 1s known. Specimen datas—MEXICO: Baja Califor- nia Sur: near Candelaria, 17 Mar 1992, R. Garces & R. Wharton, 1d (USNMS52781); San Jose Viejo, 20 Mar 1992, R. Wharton & R. Garces, 12 (TAMU USNMS52783); same, 18 Oct 1991, 12 (TAMU USNMS2782); Santiago, 16 Mar 1992, R. Garces & R. Wharton, 12 (USNM52780). Sonora: San Jose Guaymas, 3 Apr 1989, FRB-13-89, 12 (USNM53368). Zonosemata scutellata (Hendel) (Figs. 1G, 3A) References.—Hendel 1936: 73 [descrip- tion, Brazil], Steyskal 1974: 234 [descrip- tion as Z. ica, Colombia]; Norrbom 1990: 53 [synonymy, Venezuela, hosts]. Comments.—This species is known from Colombia, Venezuela, and northeastern Brazil. Its biology is poorly known. Norr- bom (1990) listed data for several speci- mens from Venezuela that were at least as- sociated with a species of Solanum; wheth- er on not they were reared is uncertain. The following specimens were clearly reared from a Solanum species and provide anoth- VOLUME 104, NUMBER 3 er data point for the distribution of Z. scu- tellata. Specimen dataa—VENEZUELA: Zulia: Mision del Tokuko (9°42'51"N 72°46'33”"W), 300 m., [reared] ex fruto de cojon de gato (Solanum sp.), 15 Aug 1995, K. Katiyar & J. Orono, 2612 (USNM48730-32). Zonosemata vidrapennis Bush (Figs. 1H, 3C) References.—Bush 1966: 321 [descrip- tion]; Norrbom 1990: 55 [host]. Recognition.—The extremely narrow wing bands distinguish Z. vidrapennis from all other species of Zonosemata (Fig. 1H). Only it, Z. vittigera, and Z. macgregori have large nonmicrotrichose areas in the ra- dial cells and cell dm between the discal and subapical bands. Both of the latter spe- cies have a dark brown, presutural, subla- teral mark on the scutum that is absent in Z. vidrapennis (Fig. 3C). Distribution.—Mexico (Michoacan, Mex- ico, Puebla, Veracruz, Oaxaca, Chiapas), Guatemala, Honduras. Biology.—The Honduran specimens were collected on fruits of Solanum lanceo- latum Cay., which is native from southern Mexico to western Panama and is an intro- duced weed in California. As this is a likely host plant, Z. vidrapennis may be a poten- tial biological control agent for this inva- sive plant. An undetermined species of So- lanum was listed by Norrbom (1990) as a host plant in Mexico. Comments.—The specimens from Gua- temala and Honduras differ from Mexican specimens in having a large, U-shaped dark brown mark on the scutum and scutellum, a small dark brown area on the notopleuron, a small brown area on the katepisternum, and a mostly to entirely dark brown sub- scutellum and mediotergite. These areas are entirely orange in the Mexican specimens, although all of the specimens have large spots on the last abdominal tergite, on the anepimeron, and in males, on the epan- drium, so this does not appear to be age- related variation. More detailed study of ad- 621 ditional specimens, particularly reared se- ries from both Mexico and Central Ameri- ca, is needed to confirm if this is geographic variation as it is here interpreted. Specimen data.—GUATEMALA: Es- cuintla: Escuintla, Grutas de San Pedro Martir, 10 Jul 1965, PJ. Spangler, 1d (USNM52409). HONDURAS: Intibuca: Yamaranguila, Las Hortencias, 14°21'11.9"N 88°14'57.5"W, 2150°m, on fruits of Sola- num lanceolatum Cav., 8 Jun 1998, L. Can- as, 26 (USNM52410-1). MEXICO: Chia- pas: San Cristobal de las Casas, 35 mi. E, 26 Jul 1957, J.A. Chemsak & B.J. Rennells, 12 (UCB). Michoacan: Tzintzuntzan, 7000 ft., 8 Aug 1954, R.E. Ryckman, D. Spencer & C.P. Christianson, 12 (USNMS52408). Oaxaca: El Punto, 10.8 mi S of, 6100 ft., 19 Jul 1987, R. Wharton, 12 (TAMU USNM52779); La Cumbre, 3.2 mi. SW of, 18 Jul 1985, Jones & Schaffner, 16 (TAMU USNM52778); Mitla, 5500 ft., 25 Jul 1962; BeEs=Moillironk464 (CNE USNMS52398—401) 1612 (USNMS52403— 4); same except 27 Jul 1962; 19 (CNC USNM52402); Oaxaca, Monte Alban, 22 Jul 1963, A. Gillogly, 12 (UCR). Puebla: Calcaloapan, 19 mi. NW, 30 Jul 1963, J. Doyen, 12 (UCB USNMS52407). Zonosemata vittigera (Coquillett) (Figs. 11.31) References.—Coquillett 1899: 261 [de- scription]; Cazier 1962: 181 [biology]; Bush 1966: 315 [revision]; Goeden and Ricker 1971: 417 [biology]. See Foote et al. (1993: 496) for additional references. Recognition.—This species differs from other species of Zonosemata, except Z. vi- drapennis and Z. macgregori, by having most of cells r,,; and dm without microtri- chia between the discal and subapical bands. It differs from Z. vidrapennis in hav- ing broader wing bands (particularly the apical band) (Fig. 11) and a presutural sub- lateral dark brown spot on the scutum (Fig. 3D) and a dark brown spot on the katepis- ternum. It differs from Z. macgregori in having a narrower apical band (cell r,,; 622 with a large hyaline spot between the apical band and vein R,,;). The submedial band on the scutum is usually orange anterior to the transverse suture, whereas in Z. mac- gregori it is usually dark brown, but there is overlap in this character in these two spe- CIES. Biology.—Solanum elaeagnifolium Cav. is the only known host (Cazier 1962, Goe- den and Ricker 1971). Distribution.—Mexico (Chihuahua, Coa- huila, Durango, Guanajuato, Michoacan, San Luis Potosi, Sonora, Tamaulipas, Za- catecas; Distrito Federal’) and southwestern United States (southeastern California to Oklahoma and Texas). In Mexico, Z. vittig- era occurs at least as far south as Guana- juato (Bush 1966). The record from Distrito Federal (Foote 1967) may be an error based on the statement by Bush (1966: 318) that Z. vittigera “‘ranges from just north of Mex- ico, D.R” The Specimen Data section in- cludes previously unreported records from Mexico. Comments.—Of 53 specimens from Mexico that I examined, 16 (2 of 5 from Chihuahua, 2 of 7 from Coahuila, and 3 of 8 from Durango, 11 of 31 from Zacatecas) have the submedial stripes at least partially red-brown to dark brown anterior to the transverse suture. One Durango female in the CNC (USNM52414) is unusual in hav- ing a pair of submedial and a pair of sub- lateral dark brown spots on tergite 5 in ad- dition to the normal pair on tergite 6. A female from Coahuila (USNM52435) has the discal and subapical bands broadly con- necteduni cell-cu;: Specimen data——MEXICO: Chihuahua: Chihuahua, 17 mi. W, 11 Jul 1964, J. Powell or J.A. Chemsak, 22 (UCB USNM53369— 70); Ciudad Jiménez, 16 km N, 26 Aug 1991, T. Griswold, 1 2 (USU USNM52434); Mapini dunes, N of, 22 Aug 1991, T. Gris- wold, 12 (USU); Samalayuca, 6 Aug 1950, R.E Smith, 12 (AMNH USNM52433). Coahuila: 20 km. from Saltillo, 26 Jun L9SO55 )Wapshete s1ce) {Sesunall 26 ue (USNM52435, 53371—2); Nueva Rosita, 29 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Aug 1974, G. Bohart & W. Hanson, 1d (USNM52436); Saltillo, 12.4 mi. S of, 4 Jul 1985, J. Wooley & G. Zolnerowich, 36 (TAMU). Durango: Durango, 5 mi. W, 6500 ft., 14 Jul 1964, J.E McAlpine, 145,12 (CNC USNM52412-13); Durango, Hwy. 45, 25 mi. S, 24 Jul 1964, L.A. Kelton, 1° (CNC USNMS52414); La Zarca, 2 mi. S, 16 Jul 1964, J.A. “Chemsak,” 1G (UGB USNM52438); Pedricena, 5 mi. S of, 29 Jul 1966, P. M. & P. K. Wagner, 1¢ (TAMU USNM52798); Tlahualilo, 4 Sep 1904, A.W. Morrill, 1 2 (USNM52437); Yerbanis, 1 mi. SW of, 12 Aug 1965, H. Burke & J. Meyer, 26 (TAMU USNM52796-7). Mi- choacan: Italia, 28 Apr 1965, A.H. Lewis, 12 (USNMS52439). San Luis Potosi: San Lorenzo} lami. S: of, 25) Jul 1976) Reisler Gruetzmacher, R. & M. Murray, Schaffner, 12 (USNMS52784). Tamaulipas: Matamo- ros) 2) Jul 1:955,« TE -ReiStephens. sae? (USNM52443); Reynosa, 10 Jul 1935, EO. Swan, 1d (USNM52444). Zacatecas: Con- cepcion del Oro, 13 mi. SW of, 9 Jul 1983, Kovarik, Harrison, Schaffner, 1¢,1@ (TAMU); Concepcion del Oro, 30 mi. SW of, 9 Jul 1983, Kovarik, Harrison, Schaff- ner, 42 (TAMU); Concepcion del Oro, 4 mi. NE of, 4 Jul 1984, Schaffner, Woolley, Carroll, Friedlander, 11¢,22 (TAMU) 43,12 (USNM52790-4); same, J.B. Woolley, 36,52 (TAMU); Concepcion del Oro, 6 sm Sof; 4) Jul \1984)«@arrolle Schaffner, Friedlander, Woolley, 146 (USNM52795); Luis Moya, 14 mi. N of, 26 Jul 1959, R.B. Selander & J.C. Schaff- ner, 12 (USNM52442). ACKNOWLEDGMENTS I thank Freddy Quesada (formerly IN- Bio) and Manuel Zumbado (INBio), who through diligent effort discovered the new species from Costa Rica; Luis Canas (Pur- due Univ.), for permission to include his data for Z. vidrapennis from Honduras; and Kamta Katiyar (Univ. del Zulia) for the specimens of Z. scutellata. 1 am also grate- ful to the curators of the other institutions who loaned study material. Vicente Hernan- VOLUME 104, NUMBER 3 dez-Ortiz kindly compared several speci- mens with the type series of Z. macgregori, and M. A. Condon, A. Freidberg, M. Po- gue, G. J. Steck and N. E. Woodley re- viewed the manuscript. Lucrecia Rodriguez produced the wing images, and Taina Lit- wak the habitus images. This work was par- tially supported by USDA, FAS, RSED (Project No. CS-ARS-6, Grant No. FG-CR- HOT): LITERATURE CITED Aluja S., M., M. Cabrera, E. Rios, J. C. Guillen A., H. Celedonio H., J. P. Hendrichs, and P. Liedo F 1987. A survey of the economically important fruit flies (Diptera: Tephritidae) present in Chiapas and a few other fruit growing regions in Mexico. Florida Entomologist 70: 321-329. Benjamin, FE H. 1934. Descriptions of some native try- petid flies with notes on their habits. United States Department of Agriculture Technical Bulletin 401, 95 pp. Bush, G. L. 1966. The genus Zonosemata, with notes on the cytology of two species. Psyche (Cam- bridge) (1965) 72: 307-323. Cazier, M. A. 1962. Notes on the bionomics of Zo- nosemata vittigera (Coquillett), a fruit fly on So- lanum (Diptera: Tephritidae). Pan-Pacific Ento- mologist 38: 181-186. Coquillett, D. W. 1899. Notes and descriptions of Try- petidae. Journal of the New York Entomological Society 7: 259-268. Foote, R. H. 1967. Family Tephritidae (Trypetidae, Trupaneidae), 91 pp. Jn Papavero, N., ed. A Cat- alogue of the Diptera of the Americas South of the United States. 57. Departamento de Zoologia, Secretaria da Agricultura, Sao Paulo. Foote, R. H., E L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Comstock Publish- ing Associates, Ithaca, x11 + 571 pp. Goeden, R. D. and D. W. Ricker. 1971. Biology of Zonosemata vittigera relative to silver-leaf night- shade. Journal of Economic Entomology 64: 417— 421. Hendel, EF G. 1936. Ergebnisse einer zoologischen Sammelreise nach Brasilien, insbesondere in das Amazonasgebiet, ausgefiihrt von Dr. H. Zerny. X Teil. Diptera: Muscidae Acalyptratae (excl. Chlo- 623 ropidae). Annalen des Naturhistorischen Museums in Wien 47: 61—106. Hernandez-Ortiz, V. 1989. Una especie nueva de Zo- nosemata (Diptera: Tephritidae) y clave de iden- tificacion de las especies del género. Anales del Instituto de Biologia, Universidad Nacional Au- tonoma de Mexico, Serie Zoologia 60: 205-210. Jenkins, J. 1996. Systematic studies of Rhagoletis and related genera (Diptera: Tephritidae). Dissertation, Michigan State University, East Lansing, 184 pp. Norrbom, A. L. 1989. The status of Urophora acuti- cornis and U. sabroskyi (Diptera: Tephritidae). Entomological News 100: 59-66. . 1990. Notes on Zonosemata Benjamin (Dip- tera: Tephritidae) and the status of Cryptodacus scutellatus Hendel (= Z. ica Steyskal syn. n.). An- nalen des Naturhistorischen Museums in Wien 91: 53-55. Norrbom, A. L., L. E. Carroll, E C. Thompson, I. M. White, and A. Freidberg. 1999. Systematic data- base of names, pp. 65—251. Jn Thompson, FE C., ed. Fruit Fly Expert Identification System and Systematic Information Database. Myia (1998) 9, vil + 524 pp. & Diptera Data Dissemination Disk (CD-ROM) (1998) 1. Say, T. 1830. Descriptions of North American dipter- ous insects [concl.]. Journal of the Academy of Natural Sciences of Philadelphia (1829) 6: 183-— 188. Smith, J. J. and G. L. Bush. 1999. Phylogeny of the subtribe Carpomyina (Trypetinae), emphasizing relationships of the genus Rhagoletis, pp. 187— 217. In Aluja, M. and A. L. Norrbom, eds. Fruit flies (Tephritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, [16] + 944 pp. Steyskal, G. C. 1974. A new species of Zonosemata Benjamin from Colombia (Diptera: Tephritidae). Journal of the Washington Academy of Science 64: 234-235. Stone, A. 1980. History of Nearctic Dipterology. Flies of the Nearctic Region 1(1): 1-62. Thompson, FE C. 1999. Data dictionary and standards, pp. 49-63. In Thompson, F. C., ed. Fruit Fly Ex- pert Identification System and Systematic Infor- mation Database. Myia (1998) 9, vii + 524 pp. & Diptera Data Dissemination Disk (CD-ROM) C1998)" 1 White, I. M., D. H. Headrick, A. L. Norrbom, and L. E. Carroll. 1999. Glossary, pp. 881—924. Jn Aluja, M. and A. L. Norrbom, eds. Fruit flies (Tephriti- dae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, [16] + 944 pp. Wiggins, I. L. 1980. Flora of Baja California. Stanford University Press, Stanford, 1025 pp. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 624—628 A NEW GENUS AND NEW SPECIES OF CEPHIDAE (HYMENOPTERA) FROM SULAWESI UTARA, INDONESIA DAviD R. SMITH AND AKIHIKO SHINOHARA (DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Insti- tution, Washington, DC 20560-0168, U.S.A. (e-mail: dsmith@sel.barc.usda.gov); (AS) Department of Zoology, National Science Museum (Natural History), 3-23-1 Hyakunin- cho, Shinjuku-ku, Tokyo, 169-0073, Japan (e-mail: shinohar@kahaku.go.jp) Abstract.—Sulawesius grandoculus, n. gen., n. sp., is described from Sulawesi Utara, Indonesia. The genus does not fit into the current classification of the Cephidae, but it is provisionally placed in the Athetocephinae, a subfamily previously known only from Madagascar. Key Words: Most cephids are found in the temperate regions of the Northern Hemisphere. Only a few occur south into subtropical or trop- ical regions, and very few occur south of the equator. In the New World, the south- ernmost record is a species of Hartigia from the State of Chiapas, Mexico (Smith 1988). In the Old World, however, the trop- ical fauna is more diverse. The subfamily Athetocephinae, with one genus and two species, occurs in Madagascar (Benson 1935, 1946), Janus and Urosyrista occur in the tropics in Myanmar and Viet Nam (Nai- to et al. 1998), and one species of Janus is known from western Kalimantan, Indonesia (Smith 1994, 1997). Goulet (1992) reported seeing a species from Australia. It is appar- ent that more cephids are to be discovered in the Old World tropics, but, unfortunately, only single or a few specimens of several species form the basis of our knowledge. Such is the case with the unusual species we report on here. It is a single specimen from Sulawesi Utara in Indonesia, so un- usual that it does not conform to, and, in fact, confounds the existing classification, Sulawesius, Indonesia, Athetocephinae even at the subfamily level, and unusual in that it is one of the few known cephids that occur in the Southern Hemisphere. It war- rants attention and adds another segment of information to our understanding of the family. PLACEMENT IN CEPHIDAE The only world classification available is that by Benson (1946) who recognized two subfamilies, Athetocephinae (misspelled Achetocephinae in some literature) and Ce- phinae. Athetocephinae occur only in Mad- agascar and Cephinae are Holarctic and Oriental. The subfamily Cephinae includes three tribes, Pachycephini (Mediterranean and Near East), Hartigiini (Holarctic and Oriental), and Cephini (Holarctic). The genus we describe does not conform to the definition of either of Benson’s (1946) subfamilies but shares certain char- acteristics with each. It shares the following with Athetocephinae: Labial palpus 3-seg- mented and without a sensory pit, much longer and stouter than the 4-segmented maxillary palpus; lack of a genal carina (al- VOLUME 104, NUMBER 3 though also absent in a few Pachycephus and Janus); antennae constricted only at the base and thickest in the middle (Fig. 2); left mandible bidentate (though with a different shaped inner tooth, Fig. 3); inner tooth of the tarsal claw longer and stouter than the outer tooth (but a small, rounded basal lobe present); radial crossvein of the forewing meeting the stigma at about its center: and front ocellus separated from each of the hind ocelli by a distance not more than the diameter of a single ocellus. It does not agree with Athetocephinae (characters in parentheses) by the following, most of which would place the new genus in Ce- phinae, tribe Hartigiini, in Benson’s key (1946): Forewing with intercostal area less transparent than the rest of the wing mem- brane (same transparency); hindwing with 5—7 hamuli (11-15 hamuli); female saw- sheath with the main axis almost straight, as in Fig. 4 (sheath strongly directed up- ward); head in dorsal view strongly narrow- ing behind eyes, as in Fig. 2 (large and ex- panded behind eyes); distance between the antennal sockets about equal to the distance between an antennal socket and the nearest anterior tentorial pit, as in Fig. 3 (distance between antennal sockets twice distance be- tween antennal socket and anterior tentorial pit); hindtibia with one preapical spine (no preapical spine); and tarsal claw with indis- tinct, rounded basal lobe (basal lobe ab- sent). The new genus has the following unique characters that separate it from Athetoce- phinae and Cephinae and all the genera therein: Eyes extremely large, covering most of side of head, in lateral view eye close to and parallel with hind margin of head, in dorsal view, head strongly narrow- ing behind eyes (Figs. 1, 2); left mandible bidentate with inner tooth broad, rounded to truncate (Fig. 3); and tarsal claw with a small, rounded basal lobe. The labial and maxillary palpi, antennae, position of the radial vein of the forewing, lack of a genal carina, and closeness of the ocelli are similar to Athetocephus; the man- 625 dibles are similar to Pachycephus; the tarsal claws are similar to Janus, Hartigia, and Athetocephus; and the less transparent in- tercostal area of the forewing, antennal-an- tennal to antennal tentorial distance, and sawsheath are similar to the Cephinae. The current subfamilies could be redefined, or a new subfamily or tribe could be justified: however, we prefer to provisionally assign this genus to Athetocephinae because we believe more significant characters, as cited above, are shared with that subfamily than with the Cephinae. Within the Athetocephinae, the new ge- nus is separated from Athetocephus by the large eyes; head strongly narrowing behind the eyes; large, blunt inner tooth of the left mandible; presence of a rounded basal lobe on the tarsal claw; the intercostal area of the forewing less transparent than the rest of the wing membrane; presence of only 5— 7 hamuli on the hind wing; and presence of a preapical spur on the hind tibia (though this can be variable in Cephidae). Benson (1935) described Athetocephus and again (1946) separated it from other Cephidae, Paulian (1961) gave an illustration of the dorsal habitus of one species, and Chevin (1984) gave some additional notes and de- scribed the female of one of the species. Sulawesius Smith and Shinohara, new genus Type species.—Sulawesius grandoculus Smith and Shinohara, n. sp. Description.—Antenna constricted to middle of 4th segment, then broadened and slightly tapering at apex, thus somewhat thickened in the middle (Fig. 2); scape 2 longer than broad; pedicel as long as broad; 3rd segment slightly longer that 4th seg- ment; apical 5—6 segments about as long as broad. Labial palpus 3-segmented, without sensory pit. Maxillary palpus apparently 4- segmented (difficult to see in specimen), shorter and more slender than labial palpus. Left mandible bidentate with outer tooth acute and inner tooth broad and rounded to truncate; right mandible with two acute Fig i: teeth, inner tooth shorter than outer, and with rounded basal region (Fig. 3). Genal carina absent. Eyes extremely large, cov- ering most of side of head, slightly diverg- ing below with lower interocular distance equal to or slightly greater than eye length and upper interocular distance slightly less than eye length; in lateral view, eye close to and parallel with hind margin of head (Figs. 1-3); in dorsal view, head strongly narrowing behind eyes (Fig. 2). Malar space slightly less than diameter of front ocellus (Fig. 3). Front ocellus separated from each hind ocellus by a distance not more than diameter of a single ocellus. Forewing with intercostal area less trans- parent than rest of wing membrane; radial crossvein (2r) meets stigma at about its cen- Sulawesius grandoculus, habitus, lateral view. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ter. Hindwing with 5—7 hamuli on front margin; with closed cells M and RS; rest of venation typical of Cephidae. Hindtibia with one preapical spine. Tarsal claws with subapical tooth longer and broader than api- cal tooth, and with indistinct, rounded basal lobe. Etymology.—The genus name is based on the Indonesian province in which it was collected. Gender: masculine. Sulawesius grandoculus Smith and Shinohara, new species (Figs. 1—5) Female.—Length, 8.1 mm; forewing length, 9.0 mm. Yellow; apex of mandible reddish brown; antenna with Ist flagellar segment and basal 4% of 2nd flagellar seg- VOLUME 104, NUMBER 3 Figs. 2-3. Sulawesius grandoculus. 2, Head, dor- sal view. 3, Head, front view. ment reddish brown; apical %4 of 2nd fla- gellar segment to apex black; mesopleuron whitish. Wings hyaline, with intercostal area of forewing darkened; veins and stig- ma black with basal half of costa and sub- costa yellow. Head and body shining, im- punctate; only mandible with somewhat elongated pits and center of clypeus rough- ened. Antenna 22-segmented, length 2.4 head width (Figs. 1, 2). Clypeus with slight cen- tral circular emargination (Fig. 3). Distance between antennal sockets only slightly lon- ger than distance from antennal socket to anterior tentorial pit (as 1.2:1.0) (Fig. 3). Distances between eye and lateral ocellus, between hind ocelli, and from hind ocelli to posterior margin of head as 1.1:1.0:1.1; postocellar area about 2 broader than long (Fig. 2). Hind basitarsus slightly shorter than remaining tarsal segments combined (as 1.0:1.1). Cercus length subequal to length of sheath (Fig. 4). Sheath and basal plate in a straight line. Lancet well sclero- tized, with truncate serrulae (Fig. 4) (not dissected, partly exerted in holotype). Male.—Unknown. 627 Figs. 4-5. Sulawesius grandoculus, apex of ab- domen and sheath, two aspects. Holotype.—¢@, labeled as_ follows: “9604021 coll. BMNH,” “Indonesia: Su- lawesi Utara, Dumoga-Bone N.P., May 1985,” “Gunung Mogogonipa, 1000 m, Malaise, J.S. Noyes, J.H. Martin.” Depos- ited in The Natural History Museum, Lon- don. Etymology.—The specific epithet refers to the unusually large eyes of this species. ACKNOWLEDGMENTS We thank Christine Taylor, The Natural History Museum, London, U.K., for allow- ing examination of this specimen. Cathy Anderson, Systematic Entomology Labo- ratory (SEL), USDA, Washington, DC, took the photographs and arranged the plates. Our thanks to Nathan Schiff, U.S. Forest Service, Stoneville, MS, and A. S. Konstantinov and T. J. Henry, SEL, Wash- ington, DC, for review of the manuscript. 628 LITERATURE CITED Benson, R. B. 1935. On the genera of the Cephidae, and the erection of a new family Syntexidae (Hy- menoptera, Symphyta). Annals and Magazine of Natural History (10)16: 535-553. . 1946. Classification of the Cephidae (Hyme- noptera Symphyta). Transactions of the Royal En- tomological Society of London 96: 89-108. Chevin, H. 1984. Note sur les Hyménopteéres Tenthre- doides (XI). Bulletin Mensuel de la Société Lin- néenne de Lyon 53: 303-309. Goulet, H. 1992. The genera and subgenera of the saw- flies of Canada and Alaska, Hymenoptera: Sym- phyta, Part. 20. Jn The Insect and Arachnids of Canada, Research Branch, Agriculture Canada, Publication 1876, 235 pp. Naito, T., D. R. Smith, and F-S. Huang. 1998. Records of Cephidae (Hymenoptera) from China and southeastern Asia, with two new species of Janus PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Stephens. Japanese Journal of Systematic Ento- mology 4: 237-242. Paulian, R. 1961. XIII. La Zoogéographie de Mada- gascar et des iles voisines. /n Faune de Madagas- car, |’ Institute de Recherche Scientifique, Tanan- arive—Tsimbazaza, 484 pp. Smith, D. R. 1988. A synopsis of the sawflies (Hy- menoptera: Symphyta) of America south of the United States: Introduction, Xyelidae, Pamphili- idae, Cimbicidae, Diprionidae, Xiphydriidae, Sir- icidae, Orussidae, Cephidae. Systematic Entomol- ogy 13: 205-261. . 1994. A new species of Janus Stephens (Hy- menoptera: Cephidae) from Indonesia. Treubia 31: 59-63. . 1997. A new species of Janus (Hymenoptera: Cephidae) from Indonesia. Entomological News 108: 24—28. [Description of same species inad- vertently published in 1994; 1997 publication a more complete description with figures. ] PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 629-639 LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF GOEDENIA SETOSA (FOOTE) (DIPTERA: TEPHRITIDAE) ON ERICAMERIA BRACHYLEPIS (A. GRAY) H. M. HALL IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: richard.goeden @ ucr.edu) Abstract.—Goedenia setosa (Foote) is a rare, monophagous or nearly monophagous, univoltine fruit fly (Diptera: Tephritidae) that feeds in the flower heads of Ericameria brachylepis (A. Gray) H. M. Hall in southern California. This sole, newly reported host belongs to the family Asteraceae tribe, Astereae, and subtribe Solidagininae, like all other known hosts of Goedenia spp. The third-instar larvae and puparia are described and figured, and selected characteristics of these stages are compared with the same stages of two other species of Goedenia. The prothorax and gnathocephalon of the second and third instars are smooth and mostly free of the minute acanthae that circumscribe most other body segments. The third instar of G. setosa lacks oral ridges, as do the third instars of two other congeners studied to date. The anterior thoracic spiracle bears two papillae in the third instar. Minute acanthae cover the center of the posterior end of the truncated, posteriorly sclerotized, caudal segment, that also is perforated by scattered, open pores, and this central area is ringed by concentric series of shallow, elliptical depressions. The life cycle is of the aggregative type. Overwintering limitedly occurs as sexually immature adults, but more frequently in dead flower heads as prepuparial third instars and puparia in apically open, vasiform cells consisting of floret and achene fragments impregnated and hardened with dried liquid feces and sap. The flattened, sclerotized posterior of the caudal segment of the overwintering prepuparium serves to plug the apical opening of the cell and thus possibly helps to shield it at least from predators. However, Colotrechnus sp. (Eulophidae), Eurytoma sp. (Eurytomidae), Eupelmus sp. (Eupelmidae), Halticoptera (Pteromalidae), and Mesopolobus sp. (Pteromalidae) are chalcidoid, primary parasitic Hy- menoptera commonly reared from the overwintered puparia. Key Words: Insecta, Goedenia, Ericameria, Asteraceae, nonfrugivorous Tephritidae, bi- ology, taxonomy of immature stages, flower-head feeding, aggregative life cycle, seed predation, parasitoids Indigenous, western North American Myopitini (Diptera: Tephritidae: Tephriti- nae) formerly assigned to the Palearctic ge- nus Urophora Robineau-Desvoidy were re- designated as Goedenia by Freidberg and Norrbom (1999). To date, the life history and immature stages of two of the eight known species of Goedenia have been de- scribed in detail, 1.e., G. timberlakei (Blanc and Foote) and G. rufipes (Curran), by Goe- den et al. (1995) and Goeden (2002), re- spectively. This paper describes the life his- tory and selected immature stages of a third species, G. setosa (Foote). MATERIALS AND METHODS The present study utilized specimens of adults reared from 1|-liter samples of mature flower heads of Ericameria brachylepis (A. Gray) H. M. Hall collected at a single lo- cation in southern California: South of Ban- ner along Chariot Canyon Road at 980-m elevation in northeastern San Diego Coun- ty. The life history study and description of the immature stages of G. sefosa were based in large part on dissections of sam- ples of live, immature and mature, flower heads and dead, overwintered flower heads collected during 1995-1999. One-liter sam- ples of these flower heads containing the scarce larvae and puparia were transported in cold-chests in an air-conditioned vehicle to the laboratory and stored under refrig- eration for subsequent dissection, photog- raphy, description, and measurement. Six- teen third-instar larvae and nine puparia dissected from flower heads were preserved in 70% EtOH for scanning electron micros- copy (SEM). Prepuparia and puparia were placed in separate, glass shell vials stop- pered with absorbant cotton and held in hu- midity chambers at room temperature for adult and parasitoid emergence. Specimens for SEM were hydrated to distilled water in a decreasing series of acidulated EtOH. They were osmicated for 24 h, dehydrated through an increasing series of acidulated EtOH and two, 1-h immersions in hexa- methyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, studied and digitally photographed with a Philips XL-30 scanning electron mi- croscope in the Institute of Geophysics and Planetary Physics, University of California, Riverside. A few adults reared from isolated pre- puparia and puparia were individually caged in 850-ml, clear-plastic, screened-top cages with a cotton wick and basal water reservoir and provisioned with a strip of pa- per toweling impregnated with yeast hydro- lyzate and sucrose. These cages were used for studies of longevity in the insectary of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the Department of Entomology, University of California, Riverside, at 25 + 1°C and 14/10 (L/D) photoperiod. A virgin male and female obtained from emergence cages were held in a clear-plastic, petri dish pro- visioned with a flattened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1994) for observa- tions of courtship and copulation behavior. Plant names used in this paper follow Hickman (1993) and Bremer (1994); tephri- tid names follow Foote et al. (1993). Ter- minology and telegraphic format used to de- scribe the immature stages follow Goeden (2001a, b, c; 2002), Goeden et al. (1993), Goeden and Headrick (1992), Goeden and Norrbom (2001), Goeden and Teerink (1997), Headrick et al. (1996), Teerink and Goeden (1999), and our earlier works cited therein. Means + SE are used throughout this paper. Digitized photographs used to construct text figures were processed with Adobe Photoshop® Version 6. RESULTS AND DISCUSSION TAXONOMY Adult.—Goedenia setosa was described by R. H. Foote (in Blanc and Foote 1987) in the genus Urophora based on a single male collected from Tar Canyon, Kings County, California. Freidberg and Norrbom (1999) reclassified most of the indigenous, western North American species of Uro- phora as Goedenia. A wing photograph of G. setosa was provided by Blanc and Foote (1987) and Foote et al. (1993), and the head was figured in dorsal view in Foote et al. (1993): Foote et al. (1993) noted that ““most” G. setosa and “most” G. timberlakei, along with Neomyopites (as Urophora) claripen- nis (Foote) are the only U.S. and Canadian Myopitini with an entirely black scutellum. Two males (9%) of 22 adults of G. setosa reared from flower heads at the study site had the middle third of the scutellum yel- low; the scutella of the remaining flies were entirely black. Similarly, 60 (20%) of 302 VOLUME 104, NUMBER 3 adults of G. timberlakei reared from 11 dif- ferent hosts (Goeden 1987, unpublished data; Goeden et al. 1995) had the apex of the scutellum ocherous to bright yellow in- stead of all black. Immature stages.—Third instar larva: Elongate-ellipsoidal, roundly tapered ante- riorly, bluntly truncated posteriorly (Fig. 1A), integument white, but venters of meso-, metathorax, and abdominal seg- ments Al—A4 with dark brown to black in- fuscation; caudal segment, including spira- cles, dark brown or black; minute acanthae conical, bluntly or sharply pointed (Figs. 1B-1, E-1), or hemispheroidal (Figs. 2A-1, C-1), and outwardly or posteriorly directed, circumscribe anterior fifth and all of venters of meso- and metathorax, and all but mid- dle half of abdominal segment A1, includ- ing all of venters, circumscribe all of A2— A7, and cover dorsal half and posterior part of caudal segment; prothorax smooth, but venter with flattened, pointed, posteriorly directed minute acanthae (Figs. 1B-1, C-1), and circumscribed by verruciform sensilla medially (Fig. 1B-2); gnathocephalon con- ical and medially divided by a vertical cleft (Fig. 1C-2), pore (Fig. I1D-1) above each dorsal sensory organ; dorsal sensory organ well-defined, hemispherical (Figs. 1D-2); anterior sensory lobe (Figs. 1C-3, D-3) bears terminal sensory organ (Fig. 1D-4), lateral sensory organ (Fig. 1D-5), suprala- teral sensory organ (Fig. 1D-6), and pit sen- sory organ (Fig. 1D-7); two medial, papil- liform integumental petals (Fig. 1D-8), five, lateral, spatulate or papilliform, integumen- tal petals (Fig. 1D-9) above each mouth- hook (Fig. 1C-4), lateralmost integumental petal (Fig. 1D-10) separate from stomal sense organ (Figs. 1C-5, D-11) ventrolater- ad of anterior sensory lobe; mouthhook bi- dentate (Fig. 1C-4); median oral lobe lat- erally compressed, apically pointed (Fig. 1C-6); verruciform sensilla circumscribe gnathocephalon dorsolaterally, laterally, and ventrolaterally (Figs. 1C-7, D-12); an- terior thoracic spiracle on posterior margin of prothorax bears two doliform papillae 631 (Figs. 1B-3, E-2, F-1); mesothoracic, lateral spiracular complex with five verruciform sensilla in vertical series (Fig. 1F-2), me- sothoracic spiracle not seen; metathoracic lateral spiracular complex with nearly closed, lateral spiracle (Figs. 1F-3, 2A-2) on posterior margin of anterior, circumnatal band of minute acanthae (Fig. 2A-1); five verruciform sensilla (Figs. 1F-4, 2A-3) in a vertical series posterior to lateral metatho- racic spiracle; lateral spiracular complex of abdominal segment Al consists of a nearly closed spiracle (Figs. 1F-5) and three ver- ruciform sensilla in vertical series posterior to spiracle (Fig. 1F-6); caudal segment with pair of posterior spiracular plates (Fig. 2B- 1) surrounded by hemispherical minute acanthae (Figs. 2B-2, C-1) interspersed dor- sally, medially, and less so, ventromedially with open pores (Figs. 2B-3, C-2), these structures in turn ringed by two to three, concentric series of shallow, elliptical de- pressions (Fig. 2B-4), with verruciform sen- silla dorsolaterally, laterally, and ventrolat- erally outside these depressions (Fig. 2B-5);: two, tapered stelex sensilla (Figs. 2B-6, C- 3, D) and another verruciform sensillum (Figs. 2B-7, C-4, E) also dorsolateral to each posterior spiracular plate; posterior spiracular plate bears three, smoothly flat- tened, elliptical rimae (Fig. 2F-1), ca. 0.02 mm in length, and four, spinose, interspi- racular processes, each ca. 0.005 mm long (Fig. 2F-2). The habitus of the third instar of G. se- tosa resembles that of G. timberlakei (Goe- den et al. 1995) and G. rufipes (Goeden 2002). In all three species, the venter of the thorax and the caudal segment are darkly pigmented (Fig. 4B, Goeden et al. 1995, Goeden 2002) and minute acanthae circum- scribe the meso- and metathorax and ab- domen, and especially noteworthy, the cen- tral, posterior surface of the caudal seg- ment, which also is prominently dotted with scattered pores (Figs. 4D-3, E-3; Goeden et al. 1995). However, in G. setosa the minute acanthae and pores appear less numerous and more centralized (Figs. 4D-3, E-3) than 632 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Third instar of Goedenia setosa: (A) habitus, anterior to left; (B) gnathocephalon and prothorax, frontolateral view, 1—minute acanthae, 2—verruciform sensilla, 3 tolateral view, 1—minute acanthae, 2—vertical, medial cleft, 3 anterior spiracle; (C) gnathocephalon, fron- anterior sensory lobe, 4—mouthhook, 5— stomal sense organ, 6—median oral lobe, 7—verruciform sensilla; (D) gnathocephalon, close-up, 1—pore, 2— dorsal sensory organ, 3—anterior sensory lobe, 4—terminal sensory organ, 5—lateral sensory organ, 6—supra- lateral sensory organ, 7—pit sensory organ, 8—medial integumental petals, 9—lateral integumental petals, 1O— lateralmost integumental petal, 11—stomal sense organ, 12—verruciform sensillum; (E) anterior spiracle, 1— minute acanthae, 2—papillae; (F) lateral spiracular complexes, |—anterior spiracle, 2—verruciform sensilla on mesothorax, 3—lateral spiracle on mesothorax, 4—verruciform sensilla on metathorax, 5—spiracle on first ab- dominal segment, 6—verruciform sensilla on first abdominal segment. VOLUME 104, NUMBER 3 Third instar of Goedenia setosa, continued, (A) close-up of lateral spiracular complex of metathorax, 1—minute acanthae, 2—spiracle, 3—verruciform sensillum; (B) caudal segment, 1—posterior spiracular plates, 2—minute acanthae, 3—pores, 4—shallow, elliptical depressions, 5—verruciform sensilla, 6—stelex sensilla, 7—-verruciform sensillum; (C) close-up of caudal segment, 1—minute acanthae, 2—pore, 3—stelex sensillum, 4—-verruciform sensillum; (D) stelex sensillum, (E) verruciform sensillum, (F) posterior spiracular plate, 1— Big 2: rimae, 2—interspiracular processes. 634 in G. timberlakei (Goeden et al. 1995, un- published data) and G. rufipes (Goeden 2002), and in G. setosa also are replaced peripherally by concentric series of shallow, elliptical depressions (Fig. 2B-4). Because G. setosa also exhibits shallow depressions (Fig. 2B-4) surrounding the center of the caudal segment, what I have termed pores may instead be deep depressions. The prothorax and gnathocephalon of G. timberlakei and G. rufipes are smooth and free of minute acanthae (Goeden et al. 1995, Goeden 2002); whereas, the protho- racic venter of G. setosa anteriorly bears minute acanthae (Figs. 1B-1, C-1). Two medial and five lateral integumental petals are present in G. setosa (Figs 1D-8, -9); whereas, G. timberlakei has two medial and six lateral integumental petals (Goeden 2002). The lateralmost of these lateral in- tegumental petals is separated from the sto- mal sense organ in all three species (Figs. 1C-4, D-10; Goeden et al. 1995, Goeden 2002). The third instars of the three species of Goedenia studied to date lack oral ridges on either side of the mouth opening, and ventral or ventrolateral to the stomal sense organs (Figs .1€) D; Goeden etal, 1995; 2002a, unpublished data). The mouthhooks of the third instars of G. setosa (Fig. 1C-6), like those of G. tim- berlakei (Goeden et al. 1995) and G. rufipes (Goeden 2002), are bidentate; however, a vertical view of the oral cavity like that fig- ured for G. rufipes (Goeden 2002) was not obtained for either G. timberlakei (Goeden et al. 1995, unpublished data) nor G. setosa. Thus, the concave, posterior surface of the anterior tooth described for G. rufipes could not be compared with the other two species. The lateral spiracular complex of the me- sothorax of G. setosa includes five verru- ciform sensilla in a vertical series (Fig. 1F- 2); whereas, in G. timberlakei, this same complex includes only two verruciform sensilla (Goeden et al. 1995), but six such sensilla in G. rufipes (Goeden 2002). The anterior spiracle of all three species PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bears only two papillae (Figs. 1E-2, F-1; Goeden et al. 1995, Goeden 2002). The metathoracic lateral spiracular com- plex of G. setosa includes five verruciform sensilla (Fig. 1F-5, 2A-3), but again, only two such sensilla in G. timberlakei (Goeden et al. 1995) and four such sensilla in G. rufipes (Goeden 2002). Finally, three ver- ruciform sensilla comprise the lateral spi- racular complex of the first abdominal seg- ment of G. setosa (Fig. 1F-7) and G. rufipes (Goeden 2002), but only one verruciform sensillum is found on this segment in G. timberlakei (Goeden et al. 1995). Puparia: Light to dark, reddish brown with dark brown to black, anterior stripe on venter of meso- and metathorax and ab- dominal segments A1l—A4, and_ similarly dark caudal segment, elongate-ellipsoidal, with smoothly rounded anterior end and truncated posterior end (Fig. 3A). Anterior end bears invagination scar (Fig. 3B-1) and raised, bilobed, anterior thoracic spiracles (Fig. 3B-2). Flattened posterior end of cau- dal segment studded with smoothly round- ed, hemispherical, minute acanthae (Fig 3C-1) interspersed with open pores (Fig. 3C-2). A pair of raised, hemispheroidal, posterior spiracular plates (Fig. 3C-3) each bear three elliptical rimae interspersed with four, peg-like, interspiracular processes. These structures are ringed by shallow, el- liptical depressions (Fig. 3C-4). Thirty-two puparia dissected from flower heads of Er- icameria brachylepis averaged 2.68 + 0.02 (range, 2.42—2.85) mm in length; 1.10 + 0.02 (range, 0.94—1.28) mm in width. DISTRIBUTION AND HOSTS To date, Goedenia setosa only is reported from two locations in southern California (Blanc and Foote 1987), 1.e., the type lo- cality and the Chariot Canyon study site, and from its newly reported, sole known host plant, Ericameria brachylepis. Like all other known hosts of Goedenia spp. (Goe- den 1987, Foote et al. 1993), E. brachylepis belongs to the family Asteraceae, tribe As- tereae, and subtribe Solidagininae (Bremer VOLUME 104, NUMBER 3 Bigs anterior to right, (B) anterior end, Puparium of Goedenia setosa: (A) habitus, |—invagination scar, 2—anterior spiracles; (C) caudal segment, 1— 2—pores, minute acanthae, plates, 4—shallow, elliptical depressions. 1994). Accordingly, G. setosa apparently is a monophagous, or nearly monophagous, tephritid. The distribution of E. brachylepis includes the southern Peninsular Ranges and South Coast of California [regions de- 3—posterior spiracular liniated in Hickman (1993)] and northern Baja California, Mexico, below 1500 m (Hickman 1993). As the type locality lies outside this distribution, it is possible that another, as-yet-to-be-determined, alternate host plant exists for G. setosa in southern California, which if so, probably also be- longs to the genus Ericameria. BIOLOGY Because G. setosa co-occurred with Tru- panea wheeleri Curran and Neospilota vir- idescens Quisenberry in separate flower heads of E. brachylepis at the Chariot Can- yon study site (‘“‘symphagy’’, Goeden 1997), where G. setosa was much less com- mon than the former tephritids, flower heads containing eggs and first instars of G. setosa could not be distinguished or de- scribed. Larva.—Upon eclosion, the first instar presumably tunneled immediately through the inner bracts and into an ovule of a pre- blossom flower head like G. rufipes (Goe- den 2001a). There, it fed with its body per- pendicular to and its mouthparts toward the receptacle within an ovule, which it first ex- cavated, then exited and entered an adjacent ovule. The receptacle was neither abraded or pitted by such feeding. An average of 1.2 + 0.2 (range, 1—2) sec- ond instars was found feeding on ovules in Six, separate, closed, preblossom flower heads (Fig. 4A). They usually fed within an ovule with their bodies perpendicular to, but always above, the receptacles (Fig. 4A). Receptacles of the six flower heads con- taining second instars averaged 0.95 + 0.05 (range, 0.85—1.14) mm in diameter. These larvae had damaged an average of 3.7 + 1.0 (range, 1—7) ovules, or about 46% of the average total of 8.0 + 0.6 (range, 6—10) ovules counted within each of the six flower However, more than 1,200 flower heads were individually dissected in order to locate these six flower heads infested with second instars. Third instars fed with their long axes ori- heads. ented perpendicular to the receptacles, and 636 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Life stages of Goedenia setosa in flower heads of Ericameia bachylepis: (A) second instar (arrow) feeding on floret; (B) third instar (arrow) feeding on soft achenes and receptacle in flower head; (C) puparium in flower head (arrow); (D) mating pair, dorsal view; (E) mating pair, ventral view; (F) mating pair, lateral view. Lines = | mm. with their mouthparts directed towards the receptacles, which they scored or pitted deeply in 11 (44%) of 25 flower heads ex- amined (Fig. 4B). The 25 flower heads (eight, closed preblossom; five, open blos- som; 12 postblossom) were dissected and contained an average of 1.0 + 0.04 (range, 1-2) third instars. These 25 flower heads av- eraged 1.01 + 0.03 (range, 0.85—1.42) mm in diameter and contained an average total of 8.6 + 0.4 (range, 6-10) ovules/achenes, of which on average 6.9 + 0.8 (range, 2— 10) ovules/achenes were damaged or about 80% (Fig. 4B). Again, well over 1,000 flower heads were individually dissected to locate these 25 infested flower heads. The receptacles that were pitted (Fig. 4B) suggested that sap constituted at least part of the diet of third instars of G. setosa. Goe- den (1988), Headrick and Goeden (1990), Goeden and Headrick (1992), Goeden et al. (1993, 1995), Headrick et al. (1996), Goe- VOLUME 104, NUMBER 3 den and Teerink (1997) first noted, scribed, and discussed sap feeding by flo- rivorous species of Tephritidae in the gen- era Trupanea, Paracantha, Neaspilota, Te- phritis, Goedenia (as Urophora), Dioxyna, and Xenochaeta, respectively. Sap could also be transported to the feeding larvae through the corniculae attaching the basal fragments of the ovules/soft achenes to the receptacle. Upon completing feeding, the larva constructed a vasiform cell consisting of ovule/achene/floret fragments impregnat- ed with and hardened by dried, liquid feces and sap. The inside wall of this cell was smooth and ringed by buried pappus frag- ments, reflecting the turning motions of the larva during the construction of the apically open-mouthed cell. The flattened, sclero- tized caudal segment apparently serves as a plug that tightly closes the mouth of the cell, yet allows respiration, and thus also may help to shield the third instar from nat- ural enemies, especially predators. Prior to pupariation the prepupa turns 180° and ori- ents with its anterior end away from the re- ceptacle, retracts its mouthparts, and forms a puparium (Figs. 4C). Pupa.—The receptacles of 29 flower heads each containing a single puparium (Figs. 4C) averaged 1.12 + 0.02 (range, 1.10—1.14) mm in diameter. The posterior of the puparium faced the receptacle, rested in the cuplike base of the cell, but free of the inner wall of the cell (Fig. 4C). Adult.—Under insectary conditions, eight males of G. setosa lived an average of 49 + 3 (range, 36-61) days, and two females lived for 41 and 75 days. These longevities were slightly longer than the mean longevities reported for G. timberlak- ei (Goeden et al. 1995), but fully commen- surate with the aggregative type of life his- tories of both tephritids (Headrick and Goe- den 1994, 1998; Goeden et al. 1995). The premating and mating behaviors of G. setosa were not studied in the field, but were observed for a male and female paired when |-day old in a petri dish arena of the type found to be useful with many other de- 637 nonfrugivorous, tephritid species (Headrick and Goeden 1994). Both sexes exhibited wing hamation (Headrick and Goeden 1994) throughout the day concurrent with other behaviors, i.e., grooming, resting, and feeding; this also was the most common wing movement reported for G. timberlakei (Goeden et al. 1995) and G. rufipes (Goe- den 2002). Premating behaviors observed with G. setosa included males and females tracking individuals of the opposite sex, during which males sometimes swayed and usually exhibited abdominal pleural disten- sion. During mating, the wings of the male were parted at about 20°, the wings of the female were parted at 60—70°, with both pairs of wings centered over the midlines of the files (Figs. 4D, E). The foretarsi of the male grasped the female abdomen dor- solaterally at the thoracic juncture, the mid- tarsi grasped the oviscape at its base, and the hindtarsi crossed under the oviscape or rested on the substrate (Fig. 4F). During copulation, the body of the female was par- allel to the substrate with the oviscape raised about 20°, while the extended acu- leus pushed the male upward and backward at an angle of about 20° anteriorly (Fig. 4F). This pair was observed to mate once or twice per day, on several successive days during a 41-day period, for a total of 21 matings that lasted an average of 158 (range, 30—320) min. The female evidenced restlessness before termination of mating by pushing against the male with its hind tarsi, by lofting its wings so as to push them against the male, and by fully extending its aculeus. The male in turn countered this be- havior with copulatory induction behavior (CIB), rocked from side to side to regain purchase or to avoid the female’s pummel- ing, and sometimes rapidly vibrated his wings, all of which appeared to calm the female and allow coitus to continue. During separation the male turned and _ rapidly walked off and away from the female while pulling his genitalia free, a process lasting about 5 s (n = 2). Seasonal history.—The life cycle of G. 638 setosa in southern California follows an ag- gregative pattern (Headrick and Goeden 1994, 1998) in which the prepuparia and puparia, and some adults variously are the overwintering stages. Adults emerge from a few puparia formed in late-fall (October— November) and these unmated, sexually immature adults overwinter. The remaining prepuparia, and puparia overwinter in dead flower heads on dormant Ericameria bra- chylepis. These overwintered individuals emerge as adults in late winter (February— March) and pass the following spring and summer (April—August), probably as non- reproductive individuals in riparian habi- tats. They eventually aggregate on preblos- som, fall-blooming, E. brachylepis, mate, and subsequently oviposit in the small, newly-formed, closed, preblossom flower heads. Natural enemies.—Overwintered puparia of G. setosa were heavily parasitized and yielded a variety of chalcidoid, primary parasitic Hymenoptera, 1.e., Colotrechnus sp. (Eulophidae), Eurytoma sp. (Eurytomi- dae), Eupelmus sp. (Eupelmidae), Halticop- tera sp. (Pteromalidae), and Mesopolobus sp. (Pteromalidae). 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 the Institute of Geophysics and Planetary Physics, University of California, River- side, greatly facilitated my scanning elec- tron microscopy. I also am grateful to Jeff Teerink for his technical assistance and to David Headrick for his helpful comments on an earlier draft of this paper. LITERATURE CITED Blanc, L. and R. H. Foote. 1987. Taxonomic obser- vations on United States Tephritidae (Diptera), with descriptions of new species. Proceedings of the Entomological Society of Washington 89: 425-439. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc., Portland, Oregon. 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 A. L. Norrbom. 1999. A generic reclassification and phylogeny of the Tribe My- opitini (Tephritinae), pp. 581—627 (Chapter 23). In Aluja, M. and A. L. Norrbom, eds. Fruit Flies (Te- phritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, Florida. 944 pp. Goeden, R. D. 1987. Host-plant relations of native Urophora spp. (Diptera: Tephritidae) in southern California. Proceedings of the Entomological So- ciety of Washington 89: 269-274. 1988. Life history of Trupanea imperfecta (Coquillett) on Bebbia juncea (Bentham) Greene in the Colorado Desert of southern California (Diptera: Tephritidae). Pan-Pacific Entomologist 64: 345-351. . 1997. Symphagy among florivorous fruit flies (Diptera: Tephritidae) in southern California, Chapter 3. /n Dettner, K., G. Bauer, and W. VOIk], eds. Vertical Food Web Interactions: Evolutionary Patterns and Driving Forces. Ecological Studies 130: 39-52. Springer-Verlag, Heidelberg, Ger- many. . 2001a. Life history and description on im- mature stages of Neaspilota footei Freidberg and Mathis (Diptera: Tephritidae) on Aster occidental- is (Nuttall) Torrey and A. Gray (Asteraceae) in southern California. Proceedings of the Entomo- logical Society of Washington 103: 191-206. . 2001b. Life history and description on im- mature stages of Tephritis joanae Goeden (Dip- tera: Tephritidae) on Ericameria pinifolia (A. Gray) H. M. Hall (Asteraceae) in southern Cali- fornia. Proceedings of the Entomological Society of Washington 103: 586—600. . 2001c. Life history and description of im- mature stages of Tephritis teerinki Goeden (Dip- tera: Tephritidae) on Hulsea vestita A. Gray (As- teraceae) in southern California. Proceedings of the Entomological Society of Washington 103: 807-825. . 2002. Life history and description of imma- ture stages of Goedenia rufipes (Curran) (Diptera: Tephritidae) on Jsocoma acradenia (E. Greene) E. Greene in southern California. Proceedings of the Entomological Society of Washington 104: 576— 588. 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. VOLUME 104, NUMBER 3 Goeden, R. D., D. H. Headrick and J. A. Teerink. 1993. 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Description of the immature stages of Trupanea imperfecta (Co- quillett). Proceedings of the Entomological Soci- ety of Washington 101: 75-85. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 640-645 A NEW SPECIES OF MICROPTEROUS DAMSEL BUG (HETEROPTERA: NABIDAE) FROM NUKU HIVA, MARQUESAS ISLANDS DAN A. POLHEMUS Department of Systematic Biology, Entomology, National Museum of Natural History, Smithsonian Institution, bishopmuseum.org) Washington, DC 20560-0105 (e-mail: bugman@mail. Abstract.—A new species of micropterous damsel bug, Nabis nukuhiva, is described from the summit area of Nuku Hiva in the Marquesas Islands, and compared to other endemic Marquesan Nabidae. Illustrations of the male paramere, male endosoma, and female dorsal habitus are provided. Key Words: As discussed previously (Polhemus 2000), the isolated islands of the Eastern Pacific are notable for their paucity of Red- uviidae, and apparently compensatory local radiations of Nabidae. Although the nabid fauna of the Hawaiian Islands has received considerable attention over the last century, with 27 endemic species now described (Polhemus 1999), the similarly interesting nabid biota of the Marquesas Islands has been far less thoroughly investigated. The only previous Heteroptera collections of any significance from these islands were made by the Pacific Entomological Survey in the 1930s, with the Nabidae sent to E. P. Van Duzee in California, who described the four endemic Nabis species recorded up to the present time (Van Duzee 1932, 1934). Limited visits were also made by individual researchers from the Bishop Museum in 1977, 1984, and 1988, but these resulted in only limited collections of Heteroptera. More recently, a pair of joint Smithsonian/ Bishop Museum expeditions to the Mar- quesas in 1999 and 2001 reached the is- lands of Nuku Hiva, Ua Huka, Ua Pou, Hiva Oa, Tahuata, Mohotane, and Fatu Hiva, obtaining extensive Heteroptera col- Heteroptera, Nabidae, Marquesas Islands lections including a new endemic microp- terous Nabis species from the cloud forests of upland Nuku Hiva, described herein. Ex- amples of still other new nabid species from both the Marquesas and Society islands are at hand, but many of them are represented by old specimens in such poor condition that it seems best to defer their description in the hope that additional fresh material may accrue from future survey work. All measurements are given in millime- ters; CL numbers following locality cita- tions refer to collection locality numbers used to cross reference specimens, photo- graphs, and field notes. Material depository codons are explained in the Acknowledg- ments section. Nabis nukuhiva Polhemus, new species (Figs, 15.254) Description.—Micropterous male: Gen- eral coloration of living specimens pale yellowish brown to orange brown (often fading to tan in dried specimens), with scat- tered darker brown markings dorsally on head, thorax and abdomen; legs maculate with darker brown; limited markings along VOLUME 104, NUMBER 3 641 Fig. 1. Nabis nukuhiva, female, dorsal habitus. central section of abdominal dorsum bright red. Length 9.95 mm, maximum (across abdomen) 2.25 mm. Head elongate, well produced ahead of width eyes, length/width = pale, semi-recumbent setae; width of vertex 1.8 the dorsal width of an eye (0.45/0.25); length of preocular portion of head approx- imately 1.9 the dorsal length of an eye 1.55/0.95, bearing 642 (0.70/0.37); length of postocular portion of head 1.1 x the dorsal length of an eye (0.42/ 0.37); ocelli absent; length of antennal seg- ments I-IV = 2.35/3.30/2.70/1.50; rostrum length 3.05, reaching to mesocoxae, lengths of segments I-IV = 0.25/1.25/1.10/0.45; coloration of head pale brown, with dark brown patches laterally behind eyes, tylus orange, posteriorly directed V-shaped brown marks present centrally at base of tylus and on frons; antenna dark yellowish brown to orange brown, segment II with a single dark brown annulation near apex, re- maining segments unicolorous. Pronotum narrow, anterior collar elon- gate, anterior lobe moderately swollen and tumescent, posterior lobe shorter than an- terior lobe, with sides weakly convergent posteriorly; length/width = 1.60/1.20, bear- ing numerous very short, pale, recumbent setae; coloration pale yellowish to reddish brown, central tumescent section sparingly marked with dark brown maculations, an- terior collar shallowly punctate, posterior lobe weakly rugulose, both bearing dark brown longitudinal midline stripe, this stripe becoming indistinct on central tumes- cent section. Scutellum elongate, roughly triangular, length/width = 1.00/0.70, basal half weakly tumescent, set off from angulate posterior half by incipient ridges extending inward from lateral margins; surfaces of both sec- tions bearing a few very short, pale, recum- bent setae; coloration pale yellowish to red- dish brown, a pair (1+1) of elongate dark brown patches present on basal half to ei- ther side of midline. Hemelytra short, micropterous; hemely- tron length 1.80, reaching to basal section of abdominal tergite I, bearing short, pale, recumbent setae; clavus evident throughout its length, venation of corium obscure, weakly suggested by remnants of cubital and radial veins, membrane present as tiny rounded flap folded slightly downward to- ward underlying tergites; coloration pale brown, veins yellowish to pale orange. Legs elongate, with fore coxa approxi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mately 1.80 as long as thick (0.90/0.50); fore femur fusiform, over 6.60 longer than wide (4.00/0.60), covered with numer- ous short, pale setae; fore tibia slender, wid- ening at apex, inner margin bearing two parallel rows of about 50 tiny black teeth bordered by pale, erect setae, outer margins bearing semi-erect pale setae, these setae becoming longer distally; middle and hind femora, tibiae and tarsi slender, covered with short, pale, semi-erect setae, middle tibia with inner margin bearing two parallel rows of about 30 tiny black teeth on basal half; coloration of legs yellowish brown to pale orange with complex darker brown maculations on all femora; all tibiae uni- colorous, lacking annulations or macula- tions, becoming slightly infuscated near tips; lengths of leg segments as follows: fore femur/tibia/tarsal 1I/tarsal 2/tarsal 3 = 4.00/3.35/0.10/0.15/0.20; middle femur/tib- ia/tarsal 1/tarsal 2/tarsal 3 = 3.70/4.00/0.10/ Q.20/0.37; hind femur/tibia/tarsal 1/tarsal 2/ tarsal 3 = 5.60/6.50/0.10/0.60/0.55. Abdomen with segments I-IX exposed, lateral margins broadly arcuate, postero- medial portions of tergites lacking raised tumescences, tergites I-V dorsally concave, with lateral margins curved upward, tergite VI broadly raised centrally, tergites VII-IX narrowed, forming a box-like genital cap- sule; all tergites covered with scattered short, pale, recumbent setae; ground color of tergites yellowish brown to orange brown, broadly overlain with dark brown maculations and scattered small bright red markings, these markings darker and more dense along midline, anterolateral angles dark brown; lengths of exposed abdominal tergites I-VIT = 0.90/0.85/0.75/0.70/0.65/ 0.75/0.20. Ventral surface of head and thorax me- dium brown, paratergites along lateral ab- domen dark blackish brown, abdominal ventrites pale yellowish, with irregular dark brown markings present along midline on abdominal ventrites I-VI; entire venter clothed with pale, recumbent setae. Genitalia with distal section of paramere VOLUME 104, NUMBER 3 643 Figs. 2-4. Micropterous Nabis species endemic to the Marquesas, male genitalic structures. 2, N. nukuhiva, paramere. 3, N. longipes, paramere. 4, N. nukuhiva, endosoma. sickle-shaped, tip sharp and acute (Fig. 2); Micropterous female: Similar to microp- endosoma with two small basal sclerites, terous male in general structure and color- plus four clusters of long sclerotized spi- ation with following exceptions: length nules in distal section (Fig. 4). 11.20 mm, maximum width (across abdo- 644 men) 3.00 mm; overall form slighly more robust than male, with abdomen more broadly expanded (Fig. 1); combined ab- dominal tergites VIII and IX roughly tri- angular, longitudinally raised medially to form a tube-like structure over upraised tip of retracted ovipositor; dark markings on dorsum of body and legs similar to those in male; venter with ventrolateral sections of abdominal segment VII not strongly swol- len on either side of ovipositor sheath. Etymology.—The name “‘nukuhiva” is a noun in apposition, and refers to the island of Nuku Hiva, to which this species is en- demic. Material examined.—Holotype, microp- terous 6: MARQUESAS ISLANDS, Nuku Hiva, cloud forest along crest of Tekau Ridge, E. of new road crossing, 1160—1220 m. (3800-4000 ft.), 8°51'39"S, 140°10’ 24"W, 21 October 1999, 10:00—15:00 hrs., Gi.60075, D> Aveand. Jz 15) Rolhemus (USNM). Paratypes (all micropterous): MARQUESAS ISLANDS, Nuku Hiva: 10 3d, 6 &, same data as holotype (USNM, BPBM, MHNP); 3 6, 1 2, same data as holotype except 23—24 August 2001, D. A. and J. T. Polhemus (USNM); 1 6, N. side of Tekao Ridge, along old road, 1020 m. 6350) f)28°51'357SP 14010 44 W, 24 Au- gust 2001, beaten from Weinmannia par- viflora, D. A. & J. T. Polhemus (USNM); 1 ?, Tapuaooa, 3000 ft., 1 June 1931, under dead leaves, Le Bronnec and H. Tauraa, Pa- cific Entomological Survey (BPBM). Discussion.—Although four endemic species of Nabis have been previously de- scribed from the Marquesas (Van Duzee 1932, 1934), Nabis nukuhiva is only the second micropterous species discovered in the archipelago. The other, N. /ongipes Van Duzee, is also restricted to the cloud forest zone, occurring along the crest of Temetiu Ridge on the island of Hiva Oa; a similar pattern of habitat preference is also exhib- ited among micropterous Nabis species col- lected by the author in Hawaii and the So- ciety Islands. Nabis nukuhiva may be separated from PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON N. longipes by the following characters: in N. nukuhiva the margins of the posterior lobe of the pronotum are slightly conver- gent posteriorly (Fig. 1), while in N. lon- gipes they are divergent posteriorly; the scutellum of N. nukuhiva is conspicuously elongate and shows an incipient division into anterior and posterior lobes (Fig. 1), while that of N. longipes is more evenly triangular and shows no such incipient transverse division; the ground color of N. nukuhiva ranges from straw yellow to pale orange brown, while the general coloration of N. longipes is distinctly dark reddish brown, with the reddish color predominat- ing in living specimens; and finally the dis- tal section of the male paramere of N. nu- kuhiva is relatively slender, curving, and sickle-shaped, while that of N. longipes is more massive and half-moon shaped (com- pare Figs. 2 and 3). To date, three Marquesan Nabis species, N. mumfordi Van Duzee, N. ancora Van Duzee, and N. nukuhiva, have been dis- sected to evaluate the internal characters of the endosoma. All possess comb-like en- dosomal sclerites, a relatively unusual char- acter state among Nabidae. Such endosomal combs are seen only in a few other groups, including the speciose endemic Hawaiian radiation in the subgenus Milu Kirkaldy, the eastern Palearctic subgenus Reuteronabis Kerzhner, and a few taxa in Australia and North America. This strongly suggests that such comblike sclerites may be a synapo- morphy linking the insular Marquesan and Hawaiian nabid radiations as monophyletic sister clades, although this hypothesis has not yet been tested in the context of a broader-scale cladistic analysis. In N. nu- kuhiva, such endosomal sclerites appear to be highly modified into clusters of long spi- nules, notably different from the short- spined combs seen in the Hawaiian taxa and certain other Marquesan species; this char- acter state, combined with the narrow pron- otum and elongate scutellum, suggests that N. nukuhiva may represent a more diver- gent lineage within this putative clade. VOLUME 104, NUMBER 3 The type series of N. nukuhiva was taken amid dense, low-stature cloud forest along the summit of Tekau Ridge, a high, narrow, elongate ridgeline that forms the crest of Nuku Hiva, representing a remnant section of the island’s old outer caldera rim. Spec- imens were collected primarily by light py- rethrin fogging of moss mats growing on the trunks and branches of intertwined trees and shrubs along the extreme crest of the ridge; a few were also taken by beating vegetation. If N. nukuhiva is indeed con- fined to cloud forest environments, then its total range comprises a very limited strip of such vegetation remaining along the Tekau ridgeline. Adamson (1936) noted of this area “The high mountains above Aakapa are one of the richest regions for collecting in the Marquesas. Unfortunately it is one of the least accessible, being a day’s journey on horseback from Taiohae, and nearly as much from Hakaui. So far as I know, these mountains cannot be ascended from the north.” This situation has now changed greatly with the construction of several new roads subsequent to the 1930s, but ironi- cally these same roads that have provided access to this rich area for biologists have also brought increasing impact from human use, weeds, and feral ungulates, all of which threaten the integrity of this fragile ecosystem. ACKNOWLEDGMENTS I thank Ron Englund of the Bishop Mu- seum, Steve Jordan of the University of 645 Connecticut, and John T. Polhemus of En- glewood, Colorado, for companionship and assistance during field work in the Marque- sas. Special thanks are also due to Jean- Yves Meyer, who provided essential help in securing permits to conduct research in French Polynesia. The holotype of Nabis nukuhiva is deposited in the National Mu- seum of Natural History, Smithsonian In- stitution, Washington, DC (USNM); para- types are held in that collection, at the Bish- op Museum, Honolulu, HI (BPBM), and in the National Museum of Natural History, Paris (MHNP). Field work in the Marque- sas was supported by the Bishop Museum and the Smithsonian Institution’s Drake Fund. This paper represents Pacific Biolog- ical Survey Contribution 2002-001. LITERATURE CITED Adamson, A. M. 1936. Marquesan Insects: Environ- ment. Bernice P. Bishop Museum Bulletin 139: 1— 7A Polhemus, D. A. 1999. A new species of riparian Na- bidae (Heteroptera) from the Hawatian Islands. Proceedings of the Entomological Washington 101: 868-874. . 2000. A revision of the endemic Hawaiian Society of reduviid genus Saicella Usinger, with descriptions of four new species (Heteroptera: Reduviidae: Emesinae). Proceedings of the Entomological So- ciety of Washington 102: 1—20. Van Duzee, E. P. 1932. New Hemiptera-Heteroptera from the Marquesas. Bernice P. Bishop Museum Bulletin 98: 177-190. 1934. A second report on Hemiptera-Heter- optera from the Marquesas. Bernice P. Bishop Mu- seum Bulletin 114: 313-326. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 646-654 PAUESIA COLUMBIANA, N. SP. (HYMENOPTERA: BRACONIDAE: APHIDITNAE) ON JUNIPER APHIDS, AND A KEY TO RELATED SPECIES K. S. PIKE, P STARY, G. GRAF, AND D. ALLISON (KSP) Entomologist and (DA, GG) Research Technicians, Washington State University, Irrigated Agriculture Research and Extension Center, 24106 N Bunn Road, Prosser, WA 99350, U.S.A. (e-mail: kpike @tricity.wsu.edu); (PS) Entomologist, Institute of Entomol- ogy, Academy of Sciences of the Czech Republic, Ceské Budéjovice, Czech Republic Abstract.—Pauesia columbiana Pike and Stary, n. sp., reared from Cinara burrilli (Wilson) on Juniperus occidentalis in Oregon is described. The North American species of Pauesia are reviewed in relation to a newly introduced grouping of the genus, and a key is provided for species of one of the two groups. Key Words: Research on aphidiine parasitoids in the Pacific Northwest has led to the discovery of numerous new species and aphid host re- lationships (Pike et al. 2000). The genus Pauesia Quilis is one of more than a dozen different aphidiine genera in western North America. Its members are restricted to uti- lizing conifer-feeding aphids in the subfam- ily Lachninae, the most common being in the genus Cinara Curtis. In the present work, a new species of Pauesia reared from Cinara burrilli (Wilson) feeding on Juni- perus occidentalis Hook in Oregon is de- scribed, a finding which has prompted a closer study of the genus. The North Amer- ican Pauesia are reviewed, a new diagnos- tic character involving apical setae on the Ovipositor sheath is presented to distinguish new groupings, and a key is provided for species of one of two groups. Group | is keyed; Group 2 is not keyed because of a lack of available quality material to study. MATERIAL AND METHODS Material evaluated.—Museum specimens afforded fundamental material for compar- aphid, parasitoid, Cinara, Pauesia ative morphological evaluation and review. Included was an examination of separate body parts preserved in glycerine-contain- ing glass capsules obtained from the late C. F Smith, North Carolina State University, Raleigh. A large part of the study included Pauesia reared from authors’ collections of determined aphids taken from a range of Pacific Northwest habitats (Pike and Stary 1996, Pike et al. 1996). In some cases, it was necessary to dissect and slide mount specimens for more detailed view. In such cases, specimens were boiled in 10% KOH for 20—60 seconds, washed in distilled wa- ter, and then mounted in DeSwann medium. References.—References used to under- stand original descriptions, and for compar- ison with type material, taxonomic lists, and description of new species included Gahan (1911), Smith (1944), Mackauer (1968), Mackauer and Stary (1967), Marsh (1979, 1991), Stary and Remaudiére (1982), Pike and Stary (1996), and Pike et al. (1996, 2000). Diagnostic characters.—Morphological nomenclature followed prevalently Huber VOLUME 104, NUMBER 3 647 mee (ees. pl 7 13 Figs. 1-13. Various features of Pauesia (drawn from females, in different scale). 1—4, P. ahtanumensis. 1, Mesonotum. 2, Propodeum. 3—4, Ovipositor sheath and apex (close-up). 5—8, P. bicolor. 5, Mesonotum. 6, Propodeum. 7—8, Ovipositor sheath and apex (close-up). 9-12, P.juniperaphidis. 9, Mesonotum. 10, Propodeum. 11, Ovipositor sheath and apex (close-up). 13, P. pinaphidis. Ovipositor sheath. 648 and Sharkey (1993). Sculpturing of the me- sonotum and propodeum are characters generally used to define the species (Gahan 1911, Smith 1944). We have found it ad- visable to make observations of both dry material and slide-mounted material. In slide form, setae, circular pits, and rugosi- ties, are often more readily distinguished. The circular pits on the mesonotum referred to by Gahan (1911) and Smith (1944) may occur more or less on all lobes (Fig. 22), merely at the base (Fig. 16), or in concert with conspicuous rugosities (Fig. 5). Other characters of the mesonotum include gran- ulation or feeble rugosities. Similarly, the propodeum also possesses useful charac- ters. The central areola may be well-de- fined, almost smooth, with few cross-cari- nae (Figs. 2, 10), or coarsely rugose with numerous cross-carinae (Figs. 6, 17). In some species, the longitudinal carinae may be significantly reduced to absent. Here we introduce the shape of the setae at the apex of the ovipositor sheaths as a new diagnostic character. These setae are of two types: (1) their base is either simple (Bigs 7458. 12, 15, 19) on @)etubitorm (Figs. 21, 23). The character is best viewed when specimens are dissected, slide-mount- ed, and observed at a minimum of 200 magnification. Sedlag (1971) emphasised the apparent importance of different types of ovipositor sheaths and accessory prongs in Pauesia and related genera. He distinguished four morphological types. The differences were presumed to be due to such factors as strong ant attendance, high mobility of host aphids and their occurrence between conifer needles. In a broader sense, Stary (1976) supported this approach, by distinguishing two basic types of ovipositor sheaths in Pauesia, and raising two groups to subge- neric level (Pauesia Quilis s. str., type spe- cies: Pauesia albuferensis Quilis, 1931, = unilachni Gahan 1927; and Paraphidius Stary 1958, type species: Aphidius califor- nicus Ashmead 1889). Later, Sedlag and Stary (1980) described another subgenus, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pauesiella from Central Europe, which manifests a spatulate-shaped ovipositor sheath with a strongly narrowed apical por- tion and bearing a group of strong, long se- tae in its dorso-apical fifth. Kiriac (1993) reviewed and keyed the Palaearctic species of Pauesia which manifest acutely nar- rowed ovipositor sheaths, compared to the prevailing broad oval shape. However, he did not take into consideration the differ- ences in the shape of the apical setae. Regarding setae on the apical portion of the sheaths, the tubiform type appeared ini- tially to be associated with the broadly oval-shaped sheaths (Fig. 20), and the sim- ple type to be associated with the narrowed arcuate shape (Figs. 7, 14), but some spe- cies such as P. juniperaphidis (Fig. 11), P. xanthothera (Fig. 18), P. salignae (Fig. 23), and others, varied from this association. Abbreviations.—The following are used in the text for collections: USNM = Na- tional Museum of Natural History, Smith- sonian Institution, Washington, DC; CFS = C. E Smith collection, Raleigh, NC; WSU = Washington State University collection, Prosser, WA. REVIEW OF NORTH AMERICAN PAUESIA SPECIES, INCLUDING SPECIES FROM MEXICO The list below includes all new, and pre- vious classified species (Marsh 1979, 1991; Pike and Stary 1996; Pike et al. 2000; Stary and Remaudiére 1982). The generic classi- fication by Marsh (1979) is followed, ex- cept that the species are arranged in two groups based on the apical setae of the ovi- positor sheath. Group |. Setae at ovipositor sheath apex with simple base: P. ahtanumensis Pike and Stary (Figs. . bicolor (Ashmead) (Figs. 5—8) . columbiana sp.n. (Figs. 24—32) . juniperaphidis (Gahan) (Figs. 9-12) . pinaphidis (Ashmead) (Fig. 13) . scorpinica (Smith) (Figs. 14—15) . xanthothera (Smith) (Figs. 16—19) 1 Se ec vcs VOLUME 104, NUMBER 3 Group 2. Setae at ovipositor sheath apex with tubiform base: . californica (Ashmead) . cinaravora Marsh . gillettei (Gahan) . macrogaster (Ashmead) . nigrovaria (Provancher) . pahtonis Pike and Stary . ponderosae Pike and Stary (Figs. 20— i) WS FOS) aa re ao) 0 as . ponderosaecola Pike and Stary ? P. procephali (Ashmead) P. pseudotsugae Pike and Stary P. rufithorax Stary and Remaudiere (Mexico) P. rugosa Stary and Remaudiere (Mexi- co) (Fig. 22) P. salignae (Watanabe) (Fig. 23) P. takomaensis (Smith) P. varigata (Smith) Group 1—ANNOTATED LISTING OF MATERIAL EXAMINED P. ahtanumensis Pike and Stary. WSU—ex Cinara ponderosae (Wilson) on Pinus ponderosa and Pinus sp. [material ex- amined from Idaho, Montana, and Wash- ington (see: Pike and Stary 1996; Pike et al 1996,72000)]: P. bicolor (Ashmead). USNM—1/Mary- land, Montgomery Co., Takoma _ Park [parasitoids labelled as Aphidius bicolor Ashm, det. Gahan; Aphidius bicolor Ashm, det. C. E Smith, C. N. Ainslie col- lection]. 2/Wisconsin, Oneida Co., Amer. Legion St. For, 16 & 24-VH-1957, ex aphids on Pinus banksiana |parasitoids labelled as Aphidius (P.) bicolor Ashm, det. Mues, coll. P A. Jones] [Figs. 5—8 drawn from this material]. CFS—Ohio, Kocking Co., 26-VII-1938, ex Cinara strobi (Fitch)? [parasitoids labelled as 468 and 471, Aphidius bicolor Ashmead, det. C. E Smith]. WSU—Idaho, Bound- ary Co., Bonners Ferry, 29-VII-1996, ex Cinara sp. on Pinus contorta. P. columbiana, n.sp. See description below. 649 P. juniperaphidis (Gahan). USNM—Colo- rado, Boulder Co., Boulder [parasitoids labelled as 61600, 944, antenna 22 seg- ments, type, USNM 14361, Aphidius jun- iperaphidis, allo-lectotype]. CFS—Colo- Boulder Co., Boulder, 6-VI-1908 [parasitoids labelled as Type 944, para- type no. 14361 USNM, A. juniperaphi- dis]. WSU—Washington, Yakima Co., Yakima Arboretum, 3-VI-94, ex Cinara pilcornis (Hartig) on Picea pungens. P. pinaphidis (Ashmead). USNM—1/Flor- ida, Duval Co., Jacksonville [parasitoid labelled as type, Aphidius, Aphidius (pi- naphidis) = bicolor det. C. F Smith, coll. Ashmead]. 2/Louisiana, Orleans Parish, New Orleans, 10 & 25-IV, Pinus palus- tris |parasitoids labelled as Aphidius pi- naphidis Ashm., det. Gahan, No. 24504, coll. H. K. Plank]. 3/Florida, Alachua Co., Gainesville, ex Lachnus pini [para- sitoid labelled as Aphidius pinaphidis Ashm. det. Gahan. coll. A. C. Mason]. P. scorpinica (Smith). USNM—1/Virginia, Fairfax Co., Vienna, 18-IV-1915, on pine [parasitoids labelled as Aphidius scorpin- icus det. C.F Smith; allotype, Aphidius scorpinicus. coll. R. A. Cushman]. 2/Vir- ginia, Fairfax Co., Vienna, 19-IV-1912, on pine, coll. J. C. Bridgwell. CFS—Vir- ginia, Fairfax Co., Vienna, 27-III-1914, on pine, coll. R. A. Cushman [parasitoid labelled as female paratype, Aphidius scorpinicus sp. n. det. C.E Smith]. P. xanthothera (Smith). USNM—1/North Carolina, Wake Co., Raleigh, 20-VII- 1940, ex black pine aphid [parasitoid la- belled as Allotype Aphidius xanthotherus Sp. mn. det; G-RSmith,;coll.. S.C..Schnell]. 2/Virginia, Norfolk City Co., Norfolk, 14-V-1931, ex Dilachnus strobi Fitch [parasitoid labelled as 6805, paratype Aphidius xanthotherus Smith, coll. G.E. Gould] [Figs. 16-19 drawn from this ma- terial]. CFS—North Carolina, Raleigh, 20-VII-1940, ex black pine aphid [para- sitoid labelled as paratype, Aphidius xan- thotherus Smith, coll. S. C. Schnell]. rado, 650 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON \ Sk \ Je (ESO ZO Figs. 14-23. Various features of Pauesia (drawn from females in different scale). 14-15, P. scorpinica. Ovipositor sheath and apex (close-up). 16-19, P. xanthothera. 16, Mesonotum. 17, Propodeum. 18—19, Ovipos- itor sheath and apex (close-up). 20-21, P. ponderosae. 20, Genitalia. 21, Ovipositor sheath apex with tubiform base setae. 22, P. rugosa. Mesonotum. 23, P. salignae. Ovipositor sheath and apex (close-up). VOLUME 104, NUMBER 3 DESCRIPTION Pauesia columbiana Pike and Stary, new species (Figs. 24-32) Diagnosis.—The new species is similar to P. juniperaphidis (Gahan), which is also associated with Cinara aphids on Junipe- rus. It is easily distinguished from the latter species by the prevalent yellow coloration of the body, by the sculpture of the pro- podeum and, less distinctly, by the mean number of antennal segments. Etymology.—Named after the Columbia River. Female description.—Eye medium sized, with sparse setae. Malar space 0.4 of eye length. Tentorio-ocular line 0.8 to equal to intertentorial line. Antenna 18—19 (17, 20) segmented, not thickened to apex, and length about as long as head, mesosoma, and half of metasoma. Flagellomere | (F1) (Fig. 27) twice as long as broad, with 2—3 longitudinal placodes; F2 (Fig. 27) equal to Fl, with 3 placodes. Middle flagellomeres (Fig. 28) as broad as basal segments. Mesosoma: Mesonotum (Fig. 24) with notaulices distinct anteriorly, effaced on disc, with sparse setae, feebly rugose (Fig. 25). Propodeum (Fig. 29) areolated, areola inside with irregular carinae to rugosities, lateral longitudinal carinae irregular to ru- gose. Forewing (Fig. 30): Stigma length about 2.5 to 3.0 times width; distal abscissa of RI (= metacarpus) somewhat shorter than length of stigma; rs vein somewhat shorter than width of stigma, subequal to 3/Rs vein; 2/Rs vein shorter than rs. Metasoma: Petiole (Fig. 26) length 2.5 to 3.0 width at spiracles, with sparse long setae in the apical portion, rugose, about 0.6 wider at apex than at spiracles. Genitalia (Fig. 31): Apical setae of ovi- positor sheath with simple bases (Fig. 32). Coloration: Head brownish, frontal part yellowish, palpi yellow, mandible with brownish apices, antenna brown, scape lighter. Mesosoma yellow, lower part of 651 mesopleuron with brown spot, metanotum with darker patterns. Wing venation brown, stigma brown. Legs yellow, pretarsi brown. Metasoma yellow, ovipositor sheaths dark brown. Length of body: About 2.2 to 2.3 mm. Male description.—Antenna 20-21 (+19) segmented. Coloration: Head brown with yellowish face and inner orbits. An- tenna light brown, palpi yellow. Mesosoma yellow brown, base of the central lobe of mesonotum, lower part of mesopleura, me- tanotum, and propodeum brown or merely with brownish patterns. Scutellum yellow brown. Wing venation brown. Legs yellow, apices of tarsi infuscated. Metasoma yellow brown, darkened to apex, petiole yellow. Material.—Holotype (2) ex Cinara bur- rilli (Wilson)—USA, Oregon, Grant Coun- ty, Hwy 7, 6 miles West of Whitney, 27- VII-2000, on Juniperus occidentalis, WSU Sample AOGS582, coll. George and Della Graft. Deposited in USNM. Paratypes: 11 2°, 10 3 (sample AOGS5S82), same data as holotype. Deposited in part in WSU and collection of P. Stary, Ceské Bu- déjovice, Czech Republic. 1 2 (sample 99K068) ex Cinara burrilli (Wilson)—USA, Oregon, Wasco County, Cow Canyon, 2 miles N of Antelope Junction, 24-[X-99, on Juniperus occidentalis, coll. K. Pike and G. Graf. Deposited in WSU. STATUS AND SEPARATION OF PAUESIA SCORPINICA (SMITH) AND RELATED SPECIES Smith (1944) redescribed P. bicolor (Ashmead), and distinguished another spe- cies, P. scorpinica as a new species, while suppressing P. pinaphidis (Gahan) as a new synonym of bicolor. Mackauer (Mackauer and Stary 1967, Mackauer 1968) reclassi- fied pinaphidis as a valid species, but placed scorpinica as a new synonym of bi- color. This classification was followed later by Marsh (1979). In the present study, we recognize all of the aforementioned taxa (bicolor, pinaphi- dis, and scorpinica) as distinct, valid spe- cies. They are comparable in overall col- 652 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 3] eae Figs. 24-32. Various features of Pauesia columbiana, female paratypes (illustrations not to equal scale). 24, Mesonotum. 25, Surface detail of central lobe of mesonotum. 26, Petiole. 27, Flagellomere 1 and 2. 28, Fla- gellomere 8. 29, Propodeum. 30, Forewing, in part (terminology after Huber and Sharkey 1993). 31, Genitalia. 32, Ovipositor sheath apex (close-up). 30 VOLUME 104, NUMBER 3 oration and in the more or less large circular pits on the mesonotum. Pauesia bicolor and P. scorpinica agree in the number of anten- nal segments (female, 21—22), but the shape and the distribution of the setae on the ovi- positor sheaths are different, as correctly described and figured by Smith (1944). Pauesia pinaphidis manifests ovipositor sheaths similar to P. bicolor, but the dense circular pits on the mesonotum are distinct from P. bicolor. Also, the 18—19 segmented antenna (female) of P. scorpinica is an in- dication that the species is different from P. bicolor and P. pinaphidis (Gahan 1911). The variation of “I9—22”’ antennal seg- ments in the redescription of bicolor by Smith (1944) is due to an apparent confu- sion of the two species. In the USNM ma- terial, even the series of “bicolor Ash- mead” and (pinaphidis) = bicolor, det. C. F Smith, with an earlier label pinaphidis, det. Gahan are separated and correspond to distinct species. In general, the material available on pinaphidis is relatively poor, and may need to be reexamined when new material becomes available. KEY TO NORTH AMERICAN SPECIES OF PAUESIA, GROUP 1 (FEMALES) [Group character: Setae with simple base at apex of ovipositor sheaths] Ne Ovipositor sheath slender and rather nar- rowed to apex (Figs. 7, 13) to claw-shaped (Fig. 14) - Ovipositor sheaths broader, suboval, not to conspicuously narrowed to the apex (Figs. 35 11 18) Ovipositor sheath slender and rather nar- rowed to apex, with | (to 0?) long setae in middle (Figs. 7, 13). Antenna 18—19 or 21— 22 segmented. Mesonotum with dispersed or dense large circular pits (Figs. 5, 16) .. 3 ~ Ovipositor sheaths slender, claw-shaped, with several long setae in middle (Fig. 14). Antenna 21—22 segmented. Mesonotum with dispersed large circular pits .. scorpinica . Antenna 18-19 segmented. Mesonotum coarsely rugose with dense large circular pits, the distance between them approxi- mately equal to pit diameter ..... pinaphidis = Antenna 21—22 segmented. Mesonotum with dispersed large circular pits (distance 2(1). 653 between pits variable) and coarse rugosities in ithe-distal half @Fig.S)) ..ie22.2. 2% 4(1). Antenna 16-17 segmented. Propodeum with bicolor distinct areola which has a few cross-carinae inside, otherwise almost smooth (Fig. 2) ssi seciict S OAR SEL RSL - ahtanumensis ~ Antenna with 18 or more segments. Propo- deum with the areola bearing a few cross- carinae, or cross-carinated and rugose (Figs. LO. V7) & 6 ae SA repr se tees 3 5(4). Antenna 18-19 or 19-20 segmented. Me- sonotum uniformly feebly rugose, without pits (Figs. 24, 25). Propodeum with areola Nn smooth inside and lateral carinae distinct (Fig. 10), or with irregular carinae to rugos- ities inside and with lateral carinae irregular (Fig. or broadly ovali(Pigsd)) 282s ssa eee 6 - Antenna 22—23 finely granulate-rugose, with a few circular pits (Fig. 16). Propodeum with areola with 29). Ovipositor sheath oval (Fig. 31) segmented. Mesonotum coarse rugosities and numerous cross-cari- nae inside, sometimes longitudinal carinae 17). Ovipositor sheath broadly oval, curved on lower side (Fig. 18) Boks o! Faye Shen Say oy ge bees xanthothera Body coloration generally dark brown to less distinct (Fig. 6(5). black, with yellowish patterns on mesoso- ma. Metasoma ferrugineous. Propodeum (Fig. 10) well-determined complete areola, almost smooth inside. An- tenna 19-20 segmented ...... Juniperaphidis ~ Body coloration prevalently yellow, with with distinct, brownish patterns on mesopleura and me- tanotum. Propodeum (Fig. 29) with the are- ola complete, with irregular carinae to ru- gosities inside, lateral longitudinal carinae irregular to rugose. Antenna I18—19 (17, 20) SESMENEA! 4 cio ceee eo ee Hee columbiana ACKNOWLEDGMENTS We express thanks to the following: D. R. Smith (Systematic Entomology Labora- tory, USDA, Washington, DC) and Paul M. Marsh (same lab, retired) for arranging for the loan of USNM material; R. L. Blinn (Dept. of Entomology, North Carolina State University, Raleigh) for the loan of material from the C. F Smith collection; and the late C. E Smith for his kindness in supplying some material, including paratypes. LITERATURE CITED Gahan, A. B. 1911. Aphidiinae of North America. Maryland Agriculture Experiment Station Bulletin 152: 147-200. 654 Huber, J. T. and M. J. Sharkey. 1993. Structure, pp. 13-59. In Goulet, H. and J. T. Huber, eds. Hy- menoptera of the World: An Identification Guide to Families. Research Branch, Agriculture Cana- da, Ottawa, Ontario, Publication 1894/E, 668 pp. Kiriac, I. 1993. Parasites of the genus Pauesia Quilis (Hymenoptera, Aphidiidae), and description of two new species. Bulletin of the Academy of Sci- ences of the Moldova Republic, Biological and Chemical Sciences, 1993 (No. 4): 40—44. (In Mol- dovian.) Mackauer, M. 1968. Pars 3. Aphidiidae, pp. 1-103. Jn Ferriere, C. and J. van der Vecht, eds. Hymen- opterorum Catalogus (nova editio). Dr. W. Junk, The Hague. Mackauer, M. and P. Stary. 1967. Hym. Ichneumono- idea, World Aphidiidae. Jn Delucchi, V. and G. Remaudiére, eds. Index of Entomophagous In- sects. LeFrancois, Paris, 167 pp. Marsh, P. M. 1979. Aphidiidae, pp. 295-313. Jn Krom- bein, K. V., P. D. Hurd, D. R. Smith, and B. D. Burks, eds. Catalog of Hymenoptera in America North of Mexico. Vol. 1, Symphyta and Apocrita (Parasitica). Smithsonian Institution Press, Wash- ington, DC, 1198 pp. . 1991. A new species of Pauesia (Hymenop- tera: Braconidae: Aphidiinae) from Georgia and introduced into South Africa against the black pine aphid (Homoptera: Aphididae). Journal of Entomological Science 26: 81—84. Pike, K. S. and P. Stary. 1996. New species of Pauesia (Hymenoptera: Braconidae: Aphidiinae) parasit- oids on Cinara (Homoptera: Aphididae: Lachni- nae) associated with conifers in the Pacific North- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON west. Proceedings of the Entomological Society of Washington 98: 324-331. Pike, K. S., P. Stary, R. Miller, D. Allison, L. Boyd- ston, G. Graf, and T. Miller. 1996. New species and host records of aphid parasitoids (Hymenop- tera: Braconidae: Aphidiinae) from the Pacific Northwest, U.S.A. Proceedings of the Entomolog- ical Society of Washington 98: 570-591. Pike, K. S., P. Stary, T. Miller, G. Graf, D. Allison, L. Boydston, and R. Miller. 2000. Aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) of North- west USA. Proceedings of the Entomological So- ciety of Washington 102: 688—740. Sedlag, U. 1971. Strukturelle Anpassungen und evo- lutive Trends in der Gattung Pauesia (Hymenop- tera, Aphidiidae). Proceedings, 13th International Congress of Entomology, Moscow, 1968, pp. 298-299. Sedlag, U. and P. Stary. 1980. Pauesia (Pauesiella) spatulata sp. n., a parasitoid of Cinara-aphids from Central Europe (Hymenoptera, Aphidiidae; Homoptera, Lachnidae). Acta Entomologica Boh- emoslovaca 77: 383-386. Smith, C. EK 1944. The Aphidiinae of North America (Braconidae: Hymenoptera). Ohio State Univer- sity Contributions in Zoology and Entomology No. 6, 154 pp. Stary, P. 1976. External female genitalia of the Aphi- diidae (Hymenoptera). Acta Entomologica Bohe- moslovaca 73: 102-112. Stary, P. and G. Remaudiere. 1982. New genera, spe- cies and host records of aphid parasitoids (Hy- menoptera, Aphidiidae) from Mexico. Annales de la Société Entomologique de France, N.S. 18: 107-127. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 655-662 MOSQUITOES (DIPTERA: CULICIDAE) COLLECTED NEAR THE GREAT DISMAL SWAMP: NEW STATE RECORDS, NOTES ON CERTAIN SPECIES, AND A REVISED CHECKLIST FOR VIRGINIA B. A. HARRISON, P. B. WuitT, S. E. Cope, G. R. PAYNE, S. E. RANKIN, L. J. BOHN, FE M. STELL, AND C. J. NEELY (BAH, PBW) Public Health Pest Management, N. C. Department of Environment and Natural Resources, 585 Waughtown Street, Winston-Salem, NC 27107, U.S.A. (e-mail: bruce.harrison@ncmail.net); (SEC) Preventive Medicine Directorate, Navy Environmental Health Center, 2510 Walmer Avenue, Norfolk, VA 23513, U.S.A.; (GRP) Great Bridge Mosquito Control, 900 Hollowell Lane, Chesapeake, VA 23320, U.S.A.; (SER, FMS) Navy Environmental and Preventive Medicine Unit No. 2, 1887 Powhatan Street, Norfolk, VA 23511, U.S.A.; (LJB) Washington Borough Mosquito Control, 900 Hollowell Lane, Chesapeake, VA 23320, U.S.A.; (CJN) Norfolk Department of Public Health, 2800 Tarrant Street, Norfolk, VA 23509, U.S.A. Abstract.—This paper provides lists of the mosquitoes collected around the North Car- olina and Virginia sides of the Great Dismal Swamp National Wildlife Refuge in 1998; the first collection records for Ochlerotatus aurifer and Oc. j. japonicus in Virginia; notes on Aedes albopictus, Oc. grossbecki, Oc. infirmatus, Oc. thibaulti, Oc. trivittatus, and Wyeomyia smithii; an up-dating of recent publications that affect the Virginia faunal list; and a revised checklist of the mosquitoes of Virginia. The record of Oc. flavescens 1s deleted from the Virginia checklist. Key Words: During 1998 mosquito-borne virus and mosquito faunal surveys were conducted in the vicinity of the Great Dismal Swamp (GDS) in southeastern Virginia and north- eastern North Carolina. The Great Dismal Swamp National Wildlife Refuge (GDSNWR) currently consists of 107,000 acres of forested wetlands. Additional areas of the swamp occur outside the refuge. The entire swamp area in the refuge is in a re- growth phase, as all areas have been lum- bered at least once. Most areas outside the GDSNWR are under agriculture or housing development. The refuge contains one of the most southern large stands of Atlantic white cedar, Chamaecyparis thyoides (L.) new records, mosquitoes, Virginia, North Carolina, checklist BSP. The GDS also serves as a major ref- uge for many mammal, bird, reptile, am- phibian, and plant species. The presence of large numbers of potential bird and mam- mal hosts for viruses and the recent rapid encroachment of humans around the pe- riphery of the swamp was a primary reason for selecting the GDS as a study site. Primary goals of the project included: (1) an assessment of mosquito-borne viruses in the area (which will not be addressed in this paper); (2) a survey of mosquito species that occur in the area; (3) up-dating a checklist of the mosquitoes of Virginia; and (4) training and capacity strengthening for all the collaborating participants. 656 MATERIALS AND METHODS The basic design for this study was to collect adult female mosquitoes in mosquito light traps (Centers for Disease Control) set in diverse habitats around the periphery of the swamp, with eight trap sites in North Carolina and eight trap sites in Virginia. The trap line was 104 miles long and in- volved sites in Camden, Gates, and Pasquo- tank counties in North Carolina and Suffolk and Chesapeake cities in Virginia. Five, two-night collection periods were planned between late April and early October; how- ever, the arrival of Hurricane Bonnie in late August canceled one collection period. Traps, supplemented with CO, (dry ice), were set out between 4:30-5:30 PM and picked up the following morning between 7:30-8:30 AM. Female mosquitoes re- trieved from the traps were returned to the laboratory in Chesapeake, Virginia, anes- thetized with triethylamine (Kramer et al. 1990), identified to species (Darsie and Ward 1981, Slaff and Apperson 1989), sort- ed into pools of up to 50 specimens, and frozen on dry ice for shipment to the Cen- ters for Disease Control and Prevention (CDC, Fort Collins) for virus isolation. Specimens were processed the same day they were captured and removed from the traps. Uncommon species or unusual spec- imens were preserved as voucher speci- mens along with collection data. Species collected in each state during the study are noted in the checklist (Table 1). Use of the genus Ochlerotatus Lynch Ar- ribalzaga and the abbreviation, Oc., follows Reinert (2000). Both Oc. atlanticus (Dyar and Knab) and Oc. tormentor (Dyar and Knab) are documented from Virginia and North Carolina based on larval characters. However, adult females of these two species cannot be separated morphologically with certainty in the mid-Atlantic Region (Har- rison et al. 1998). Therefore we arbitrarily chose to report only the former during this study. Likewise, only Culex pipiens L. is reported even though adult females are in- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON distinguishable from Cx. guinquefasciatus Say, and hybrids, all of which might have been encountered. RESULTS AND DISCUSSION A total of 72,638 mosquito specimens were collected in 110 trap-nights during four two-night collection periods in April, June, July, and October. Of that total, 96.2% of the specimens (69,880) were col- lected during the first two periods and Och- lerotatus c. canadensis (Theobald) made up 97.9% of those specimens. After June, rain- fall was extremely sparse and specimen numbers declined dramatically to only 1,634 in July and 1,124 in October. The paucity of rainfall affected temporary pool species, which probably explains the limit- ed number of Aedes Meigen, Ochlerotatus, and Psorophora Robineau-Desvoidy that were collected. The lack of rainfall also af- fected the pH of the water in the swamp in October (as low as 3.2 to 3.6 in some are- as), Which may have influenced the pres- ence of certain species. A total of 24 spe- cies were collected in both states, of which 21 species came from North Carolina and 23 species came from Virginia (Table 1). Ochlerotatus aurifer (Coquillett), Oc. grossbecki (Dyar and Knab) and Oc. trivit- tatus (Coquillett) were not collected by us in North Carolina. Neither Oc. aurifer nor Oc. grossbecki have ever been collected in North Carolina even though they were found in Virginia 3.9 and 17.7 km, respec- tively, from the state line. Psorophora col- umbiae (Dyar and Knab) was not collected on the Virginia side of the GDS during this study, but is commonly collected by local mosquito control personnel in Chesapeake City. Notes for the new state records and other species are below. NEw STATE RECORDS Ochlerotatus aurifer (Coquillett) Ochlerotatus aurifer females were col- lected in two sites in Suffolk, Virginia. Twenty specimens were collected beside VOLUME 104, NUMBER 3 Jericho Lane at GDS site VA-5, on the edge of the Great Dismal Swamp National Wild- life Refuge (GDSNWR) on: 28 April (4 @); 29 April (12 2); and 19 May (4 @). This trap site was in a mature deciduous forest with a pasture on one side and a road with a ditch filled with clear water on the other, and is approximately 17.7 km from North Carolina. An additional two females were collected in wet deciduous forest next to Desert Road site VA-8 on 29 April (1 9) and 3 June (1 @), and only 3.9 km from North Carolina. Seven specimens collected at VA-5 on 28 and 29 April 1998 are de- posited in the National Museum of Natural History (NMNH), Smithsonian Institution, Washington, DC, and the remaining speci- mens are deposited in the Public Health Pest Management (PHPM) collection in Winston-Salem, NC. Certain characters on these specimens differ from the description of this species provided by Carpenter and LaCasse (1955). Those authors described: (1) the sides of the scutum and most of the prescutellar space with pale golden scales; (2) the posterior pronotum with curved brown scales dorsal- ly, becoming pale golden ventrally; (3) the first abdominal tergum dark with a few in- termixed pale scales; and (4) the femora with dark and pale scales intermixed, darker toward the apices, pale on the posterior sur- face. Our specimens have: (1) the sides of the scutum with bronzy-brown scales ante- riorly, becoming slightly lighter posteriorly over the wing root, and the scales around the prescutellar bare spot bronzy-brown an- teriorly, slightly paler posteriorly; (2) the posterior pronotum covered in curved bronzy-brown scales; (3) the first abdomi- nal tergum with a distinct median posterior patch of creamy scales; and (4) the femora (particularly the forefemur) almost entirely cream colored, except a few very sparse scattered dark scales that become more common and form a dark area near the api- ces. Other than these differences, the spec- imens fit the keys listed above and that of Carpenter and LaCasse (1955). Dr. R. C. 657 Wilkerson, Walter Reed Biosystematics Unit, Walter Reed Army Institute of Re- search, kindly examined specimens of Oc. aurifer in the NMNH mosquito collection and found many had bronzy-brown scales on the sides of the scutum like our speci- mens, instead of pale golden scales as listed by Carpenter and LaCasse (1955). The se- nior author (BAH) collected specimens of 1967 in the Walter Reed Army Institute of Research field study site in the Pocomoke Cypress Swamp in Worchester County, Maryland, only 2.4 km from Virginia. Those specimens had the sides of the scutum pale golden, as de- scribed by Carpenter and LaCasse (1955). The differences noted on the GDS speci- mens should be considered variations. Map number 14 (p. 239) in Darsie and Ward (1981) depicts the distribution of Oc. aurifer across the northern part of Virginia. However, there are no previous published collection records of this species in Virgin- ia. Apparently these authors based that map distribution on a 1945 record of Oc. aurifer collected in Charles County, MD, across the Potomac River from the northern part of Virginia, and a 1937 record collected in the Pocomoke Cypress Swamp in Worchester County, MD, adjacent to Virginia on the eastern shore (Bickley 1957). Accordingly, the above 22 specimens collected during this study represent the first documented evidence of this species in Virginia. These collections of Oc. aurifer just north of the state boundary suggest that it probably oc- curs in northeastern North Carolina. Oc. aurifer in Ochlerotatus japonicus japonicus (Theobald) Ochlerotatus j. japonicus was first rec- ognized in the United States in 1998 and reported from New Jersey and New York (Peyton et al. 1999). Since then it has spread into other states and districts as far south as Virginia. It was first detected in Virginia when three males and one female were collected as larvae in Occoquan, Prince William County, Virginia, on July 658 21, 2000. The reared adults were shipped to the senior author for identification and confirmed as this species. The larvae were collected in stone flower pots and birdbaths by Jim Herendeen and Karrie Trumble un- der the supervision of Benjamin F Mc- Laurin, Jr., Gypsy Moth and Mosquito Con- trol Program, Prince William, Prince Wil- liam County, VA. These specimens repre- sent the first collection of Oc. j. japonicus in Virginia. Two males and one female are deposited in PHPM. COMMENTS ON CERTAIN SPECIES Aedes albopictus (Skuse) Aedes albopictus was first detected in Virginia in 1991 in Virginia Beach and re- ported by Dreda McCreary in the non-ref- ereed Virginia Mosquito Control Associa- tion Newsletter, (Skeeter, Vol. 52, 1992). Since then it has become a major cause for mosquito complaints in many areas of the state. In North Carolina this species is dis- tributed throughout every county and is the primary cause of mosquito complaints, sometimes even in areas with salt marsh. During this study 16 specimens of Ae. al- bopictus were collected in CDC light traps in five of the eight Virginia trap sites around the GDS in Chesapeake and Suffolk cities between 2 June and 7 October. In North Carolina, seven specimens were col- lected in four of the eight traps in Camden, Gates and Pasquotank counties between 2 June and 7 October. Positive sites were those that were closest to human habitation or near discarded tires and containers. Ochlerotatus flavescens (Mueller) Darsie and Ward (1981) recorded Oc. fla- vescens in Virginia based on one sentence in the non-refereed newsletter distributed by the Virginia Mosquito Control Associa- tion (Skeeter, Vol. 23, 1978). This sentence, “Chuck Burr caught a rare mosquito this year, Aedes flavescens.”’ is the only indi- cation for this species in Virginia, as the identification was not confirmed and the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON specimen was not preserved. Ochlerotatus flavescens is considered a Holarctic species that prefers deep temporary pools in mead- ows and marshes on the open plains, and is most abundant in the early spring (Carpen- ter and LaCasse 1955). The nearest pub- lished records of this species to Virginia are in New Jersey (Crans 1970), while Means (1979) lists only three collections from New York. Considering the circumstances surrounding this record and the total ab- sence of specimens collected in Virginia (or even near Virginia) before and since that time, we feel the specimen was probably misidentified. Accordingly, Oc. flavescens is deleted from the checklist of Virginia mosquitoes. Ochlerotatus grossbecki (Dyar and Knab) Gladney and Turner (1969) reported re- cords of this species in Grassymead and Pittsylvania County, Virginia. We have been unable to locate ““Grassymead’’, which apparently came from Dyar (1922). The senior author (BAH) collected speci- mens of this species in a large freshwater (clear) pool on Assateague Island, Accom- ack County, Virginia, in March, 1967. Sev- en females of this species were collected at two sites in Suffolk City and one site in Chesapeake City during the summer of 1998. Three specimens were collected be- side Jericho Lane at GDS site VA-5, on the edge of the Great Dismal Swamp National Wildlife Refuge on 28 April (1 2) and 29 April (2 2). The description of this site is given under Oc. aurifer (above). Two ad- ditional females were collected at the same site on 19 May in a focused attempt to col- lect this species. Two larvae were collected in Emerald Greens, Chesapeake City on 21 May and reared to adults. These last two specimens were collected in a ditch with clear water. In the mid-Atlantic and south- ern states Oc. grossbecki is a late winter- early spring mosquito that is rarely collect- ed, primarily because adults hatch and die before most mosquito collection efforts start. The Jericho Lane site on the edge of VOLUME 104, NUMBER 3 the GDSNWR in Suffolk is only 17.7 km from North Carolina. Collections of Oc. grossbecki from that site suggest that it al- most certainly occurs in North Carolina, where it previously has not been docu- mented (Harrison et al. 1998). A specimen of Oc. grossbecki from trap site VA-5 col- lected on 29 April, has been deposited in NMNH, the remaining six specimens are deposited in PHPM. Ochlerotatus infirmatus (Dyar and Knab) The only previous record of Oc. infir- matus from Virginia is that of three females from Assateague Island reported by Arnell (1976). Nine females of this species were collected at five sites in Virginia during this study. One specimen was collected at GDS site VA-1 on 15 July. This site was in wet deciduous forest beside U.S. highway 17 in Chesapeake City, about | km from North Carolina. Four specimens were collected at GDS trap site VA-2 on 14 July (2 2) and 6 October (2 2). This site was also in wet deciduous forest beside U.S. highway 17 in Chesapeake City, about 4.8 km from North Carolina. One specimen was collected at GDS site VA-6 on 6 October. This site was in mature deciduous forest beside Washing- ton Ditch lane in the GDSNWR in Suffolk. One specimen was collected at GDS site VA-7 on 7 October in a mature pine forest beside Railroad Ditch lane and behind the GDSNWR headquarters. Two specimens were collected at GDS site VA-8 (see site description above under Oc. aurifer) on: 6 October (1 ¢) and 7 October (1 2). One voucher specimen from Virginia (Chesa- peake, highway U.S. 17, VA-1, 15 July 1998) is deposited in PHPM. The remain- ing specimens were pooled for virus isola- tion attempts. Ochlerotatus infirmatus can be very common in the coastal regions of North Carolina and specimens were col- lected during the project in Camden and Gates counties. Ochlerotatus thibaulti (Dyar and Knab) Gladney and Turner (1969) reported only two previous collections of Oc.thibaulti in 659 Virginia (Giles County Beach). During this study one female of Oc. thibaulti was collected in Suffolk, Virginia at GDS site VA-8 (see the site description under Oc. aurifer) on 14 July. Three addi- and Virginia tional specimens were identified during the study period from collections in Virginia Beach (courtesy, Dreda McCreary). Speci- mens of this species were also collected at two sites in North Carolina. One female was collected in Camden County at GDS site NC-2 on 2 June at the North Carolina Welcome Center next to the Dismal Swamp Canal on U.S. highway 17 about 4.8 km from Virginia. The trap site was in mature deciduous forest next to a cultivated field. The second collection (5 ¢, 4 6), on 30 April, was in Merchant’s Mill Pond State Park, Gates County, just southwest of the GDSNWR. One specimen was captured while biting, while the others were aspirat- ed from resting sites in hollow trees and stumps. The nine specimens from Gates County are deposited in PHPM. These two additional records, plus those of Harrison et al. (1998), suggest that this species is prob- ably scattered throughout the coastal plain of North Carolina, wherever there is suit- able old swamp habitat. Ochlerotatus trivittatus (Coquillett) Only three previous Virginia collections of Oc. trivittatus, in Franklin, Pittsylvania and Roanoke counties in the western pied- mont and mountains, have been published (Gladney and Turner 1969). One female of this species was collected in Virginia at GDS site VA-3 on 15 July. This site is lo- cated off George Washington highway in Chesapeake in deciduous forest near a housing development. This record repre- sents the most eastern collection of this spe- cies in Virginia. In North Carolina, Oc. tri- vittatus 1s considered a locally common piedmont and mountain species (Harrison et al. 1998), and has only been collected in one coastal plain county (Robeson). 660 Table 1. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Revised checklist of mosquito species in Virginia.! 1. Aedes aegypti (L.) 2. Aedes albopictus (Skuse)—NV 3. Aedes cinereus Meigen 4. Aedes vexans (Meigen)—NV 5. Anopheles atropos Dyar and Knab 6. Anopheles barberi Coquillett 7. Anopheles bradleyi King 8. Anopheles crucians Wiedemann—NV 9. Anopheles punctipennis (Say)—NV 10. Anopheles quadrimaculatus Say 11. Anopheles smaragdinus Reinert 12. Anopheles walkeri Theobald 13. Coquillettidia perturbans (Walker)—NV 14. Culex erraticus (Dyar and Knab)—NV 15. Culex peccator Dyar and Knab 16. Culex pipiens L—NV 17. Culex quinquefasciatus Say 18. Culex restuans Theobald—NV 19. Culex salinarius Coquillett—NV 20. Culex territans Walker—NV 21. Culiseta inornata (Williston) 22. Culiseta melanura (Coquillett)—NV 23. Ochlerotatus atlanticus (Dyar and Knab)—NV 24. Ochlerotatus atropalpus (Coquillett) 25. Ochlerotatus aurifer (Coquillett)—V 26. Ochlerotatus c. canadensis (Theobald)—NV 27. Ochlerotatus cantator (Coquillett) 28. Ochlerotatus dupreei (Coquillett) 29. Ochlerotatus fulvus pallens (Ross) 30. Ochlerotatus grossbecki (Dyar and Knab)—V 31. Ochlerotatus hendersoni (Cockerell) 32. Ochlerotatus infirmatus (Dyar and Knab)—NV 3. Ochlerotatus j. japonicus (Theobald) 34. Ochlerotatus mitchellae (Dyar) 35. Ochlerotatus sollicitans (Walker) 36. Ochlerotatus sticticus (Meigen) 37. Ochlerotatus stimulans (Walker) 38. Ochlerotatus taeniorhynchus (Wiedemann) 39. Ochlerotatus thibaulti (Dyar and Knab)—NV 40. Ochlerotatus tormentor (Dyar and Knab) 41. Ochlerotatus triseriatus (Say)—NV 42. Ochlerotatus trivittatus (Coquillett)—V 43. Orthopodomyia alba Baker 44. Orthopodomyia signifera (Coquillett)—NV 45. Psorophora ciliata (Fabricius) 46. Psorophora columbiae (Dyar and Knab)—N 47. Psorophora cyanescens (Coquillett) 48. Psorophora discolor (Coquillett) 49. Psorophora ferox (von Humboldt)—NV 50. Psorophora horrida (Dyar and Knab) 51. Psorophora howardii (Coquillett) 52. Psorophora mathesoni Belkin and Heinemann 53. Toxorhynchites rutilus septentrionalis (Dyar and Knab) 54. Uranotaenia sapphirina (Osten Sacken)—NV 55. Wyeomyia smithii (Coquillett) ' Species with a *“‘N’* (= North Carolina) or a ** Vv’ (= Virginia) following the author(s) name(s) are species that were collected around the periphery of the Great Dismal Swamp in 1998. Wyeomyia smithii (Coquillett) Coyne and Hagmann (1970) determined that the species in Virginia was Wyeomyia haynei Dodge, rather than the more north- ern Wy. smithii, and this was reported in Darsie and Ward (1981). Bradshaw and Lounibos (1977), however, found that these names represent variations of a single spe- cies. Based on the work of Bradshaw and Lounibos (1977), Darsie and Morris (1998) sunk Wy. haynei as a junior synonym of Wy. smithii. REVISED CHECKLIST OF MOSQUITO SPECIES IN VIRGINIA The last publications to specifically ad- dress the mosquito fauna of Virginia were Bickley (1957) and Gladney and Turner (1969), who, using the data provided by Dorer et al. (1944) and later collection re- cords, documented 47 species occurring in the state. In subsequent studies an addition- al four species were recognized in Virginia and added to a list of the mosquitoes of Virginia published in Darsie and Ward (1981): Wyeomyia haynei Dodge (Coyne and Hagmann 1970); Ochlerotatus hender- soni (Cockerell) (Zavortink 1972): Ochler- otatus infirmatus (Arnell 1976); and Och- lerotatus flavescens (Skeeter 1978, non-ref- ereed newsletter of the Virginia Mosquito Control Association). Since 1981 additional species have been documented in Virginia. Aedes albopictus was recognized in the state by McCreary (Skeeter 1992, non-ref- ereed newsletter of VMCA). Following the recognition of five sibling species in the Quadrimaculatus Complex (Reinert et al. VOLUME 104, NUMBER 3 1997), Strickman et al. (2000) confirmed the presence of Anopheles quadrimaculatus Say, sensu stricto, and Anopheles smarag- dinus Reinert, in Virginia. Also, Powell and Harrison (2001) provided the first docu- mentation of Oc. tormentor in Virginia. In this study we have documented the occur- rence of Oc. aurifer and Oc. j. japonicus in Virginia. Considering the above additions, our de- letion of Oc. flavescens from the Virginia Checklist, and the recognition of Wy. hay- nei as a junior synonym of Wy. smithii (Darsie and Morris 1998), these actions re- sult in 55 species and subspecies recog- nized in the checklist of the mosquito spe- cies in Virginia (Table 1). ACKNOWLEDGMENTS We gratefully acknowledge L. A. Culp, Jr., Refuge Manager, Great Dismal Swamp National Wildlife Refuge, for permission and assistance in working on the refuge. For assistance and support in establishing the collaborative arrangements, we thank N. H. Newton and A. L. Anderson, Public Health Pest Management, N. C. Department of Environment and Natural Resources, Ra- leigh; R. J. Soderholm, Deep Creek Mos- quito Control Commission, Chesapeake; and A. M. Flemming, Norfolk Department of Public Health, Norfolk. We are most grateful to the following individuals for field and laboratory assistance: K. R. Foley, Sr. NN: Amen, and ‘T.’R. Jones, Great Bridge Mosquito Control Commission, Chesapeake; I. J. Lemnios and E. B. Dan- ganan, Washington Borough Mosquito Control Commission, Chesapeake; L. V. DeMarco and B. Bynum, Norfolk Depart- ment of Public Health, Norfolk; G. L. Wil- son, Deep Creek Mosquito Control Com- mission, Chesapeake; and E. B. Williams, Preventive Medicine Directorate, Navy En- vironmental Health Center, Norfolk. We also thank C. J. Mitchell and N. Karabatsos of the Centers for Disease Control and Pre- vention, Fort Collins, CO, for advice and assistance. 661 LITERATURE CITED Arnell, J. H. 1976. Mosquito Studies (Diptera, Culic- idae). XXXII. A revision of the scapularis group of Aedes (Ochlerotatus). American Entomology 13(3): 1-144. Bickley, W. E. 1957. Notes on the distribution of mos- quitoes in Maryland and Virginia. Mosquito News 17: 22-25. Bradshaw, W. E. and L. P. Lounibos. 1977. Evolution of dormacy and its photoperiodic control in pitch- Contributions of the Institute (Ann Arbor) er-plant mosquitoes. Evolution 31: 546—567. Carpenter, S. J. and W. J. LaCasse. 1955. Mosquitoes of North America (north of Mexico). University of California Press, Berkeley and Los Angeles, 353: pp:, 127 pls: Coyne, G. E. and L. E. Hagmann. 1970. Distribution of Wyeomyia species in New Jersey. Proceedings of Fifty-Seventh Annual Meeting New Jersey Mosquito Extermination Association, pp. 190— 195, Crans, W. J. 1970. The occurrence of Aedes flavescens (Mueller), Psorophora cyanescens (Coquillett) and Culex erraticus (Dyar and Knab) in New Jer- sey. Mosquito News 30: 655. Darsie, R. FE, Jr. and C. D. Morris. 1998. Keys to the adult females and fourth instar larvae of the mos- quitoes of Florida (Diptera, Culicidae). Bulletin of the Florida Mosquito Control Association |: 1— 156. Darsie, R. FE, Jr and R. A. Ward. 1981. Identification and geographical distribution of the mosquitoes of North America, north of Mexico. Mosquito Sys- tematics Supplement |: 1—313. Dorer, R. E., W. E. Bickley, and H. P. Nicholson. 1944. An annotated list of the mosquitoes of Virginia. Mosquito News 4: 48—50. Dyar, H. G. 1922. The mosquitoes of the United States. Proceedings of the United States National Muse- um 62: 1-119. Gladney, W. J. and E. C. Turner, Jr. 1969. The Insects of Virginia: No. 2. Mosquitoes of Virginia (Dip- tera: Culicidae). Virginia Polytechnic Institute Re- search Division Bulletin 49: 1—24. Harrison, B. A., P. B. Whitt, E. E. Powell, and E. Y. Hickman, Jr. 1998. North Carolina mosquito re- cords. I. Uncommon Aedes and Anopheles (Dip- tera: Culicidae). Journal of the American Mos- quito Control Association 14: 165-172. Kramer 5 1D"S4 Bs 7Presser, Es J. Houk, and! Jz 1. Hardy. 1990. Effect of the anesthetizing agent tri- ethylamine on western equine encephalomyelitis and St. Louis encephalitis viral titers in mosqui- toes (Diptera: Culicidae). Journal of Medical En- tomology 27: 1O008—1010. Means, R. G. 1979. Mosquitoes of New York. Part 1. The genus Aedes Meigen with identification keys 662 to genera of Culicidae. New York State Museum Bulletin 430a: 1—221. Peyton, E. L., S. R. Campbell, T. M. Candeletti, M. Romanowski, and W. J. Crans. 1999. Aedes (Fin- laya) japonicus japonicus (Theobald), a new in- troduction into the United States. Journal of the American Mosquito Control Association 15: 238— 241. Powell, E. E. and B. A. Harrison. 2001. Ochlerotatus tormentor (Dyar and Knab), a new mosquito re- cord for Virginia (Diptera: Culicidae). Proceed- ings of the Entomological Society of Washington 103: 1025-1026. Reinert, J. E, P. E. Kaiser, and J. A. Seawright. 1997. Analysis of Anopheles (Anopheles) quadrimacu- latus complex of sibling species (Diptera: Culici- dae) using morphological, cytological, molecular, genetic biochemical and ecological techniques in an integrated approach. Journal of the American Mosquito Control Association 13 (Supplement): 1-102. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 2000. New classification for the composite ge- nus Aedes (Diptera: Culicidae: Aedini), elevation of subgenus Ochlerotatus to generic rank, reclas- sification of the other subgenera, and notes on cer- tain subgenera and species. Journal of the Amer- ican Mosquito Control Association 16: 175-188. Slaff, M. and C. S. Apperson. 1989. A key to the mos- quitoes of North Carolina and the Mid-Atlantic states. North Carolina State University Agriculture Extension Service Publication AG-412: 1-38. Strickman, D., T. Gaffigan, R. A. Wirtz, M. Q. Bene- dict, C. S. Rafferty, R. S. Barwick, and H. A. Williams. 2000. Mosquito collections following local transmission of Plasmodium falciparum ma- laria in Westmoreland County, Virginia. Journal of the American Mosquito Control Association 16: 219-222. Zavortink, T. J. 1972. Mosquito Studies (Diptera, Cu- licidae). XX VIII. The New World species former- ly placed in Aedes (Finlaya). Contributions of the American Entomology Institute (Ann Arbor) 8(3): 1-200. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 663-671 MACROPSYLLA NOVAEHOLLANDIAE (SIPHONAPTERA: HYSTRICHOPSYLLIDAE), A NEW SPECIES OF FLEA FROM TASMANIA MICHAEL W. HASTRITER AND MICHAEL EK WHITING (MWH) Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM, P.O. Box 20200, Provo, UT 84602-0200, U.S.A. (e-mail: hastritermw @ sprintmail.com); (MFW) Department of Zoology, 574 Widtsoe Building, Brigham Young University, Pro- vo, UT 84602, U.S.A. Abstract.—A new species of flea of the family Hystrichopsyllidae, Macropsylla no- vaehollandiae Hastriter, collected from Pseudomys novaehollandiae Waterhouse near Coles Bay, Tasmania, is described. The familial placement of the genus Macropsylla Rothschild and Stephanopsylla Rothschild are discussed. Key Words: Fleas recently collected from small mam- mals in northeastern Tasmania were initial- ly thought to represent Macropsylla her- cules Rothschild. Further study revealed that they represent an undescribed species. Rather common in eastern Australia, the nominate species, M. hercules, was de- scribed from two females, individually col- lected from Mus velutinus = Rattus lutreo- lus (J. E. Gray) and an unidentified species of Mus from Launceston, Tasmania. Hop- kins and Rothschild (1956) later illustrated the male head and genitalia of M. hercules from Emerald, Victoria, Australia and pro- vided a brief description. The purpose of this paper is to describe a new species of Macropsylla and provide supportive evi- dence for the familial placement of this unique genus. MATERIALS AND METHODS The overall body dimensions of males and females were measured from the foremost portion of the frons to the apex of the st. VIII in males and to the posterior border of the sensilium in females. Illustrations were prepared from digitized images prepared flea, Hystrichopsyllidae, Macropsylla, Stephanopsylla, Tasmania with a Zeiss Stemi SV 11 dissecting micro- scope and Dage-MTI digital camera. Ter- minology of morphological structures fol- lows those of Rothschild and Traub (1971). Macropsylla hercules Rothschild 1905 (Figs. 4, 5, 8-12, 14, 16, 17) Material examined.—AUSTRALIA, New South Wales: ex Rattus assimilis = Rattus fuscipes (Waterhouse), 24 6, 32 ¢; ex Rattus ratius (L.), 3vd% 3°25, CxuR: fus- Cipes! 2 OY QO EPR AUTOLUST SNS 52 and ex Melomys sp., 1 °. Northern Terri- tory: ex Melomys littoralis = Melomys bur- toni (Ramsay), | 2. Queensland: ex R. fus- Cipes, WZ Gis, 24S? "ex Ra assimilise 14S 42 2: ex Parameles nasuta E. Geoffroy, 3 °; ex Rattus sp., 2 6, | 2, and Melomys sp.., 1 2. South Australia (Kangaroo Island): ex R. fuscipes, 8 3, 6 &. South Australia (mainland): R. fuscipes, 1 3d. Tasmania: ex R. lutreolus, 20 3, 21 2; Pseudomys hig- ginsi (Trouessart), 1 ¢, 4 2; ex Rattus lu- treolus velutinus = R. lutreolus, 1 3d; ex Mus velutinus = R. lutreolus, 1 @ (lecto- type), and ex Mus sp., 1 2 (paralectotype). West Head (location unknown): ex R. fus- 664 cipes, 1 &. Victoria: ex R. assimilis, 11 6, 4 9; ex Mus assimilis = R. fuscipes, 8 6, 5 2; ex R. lutreolus or R. assimilis, 9 2; and ex R. lutreolus, 1 @. Remarks.—Rothschild (1905) indicated in his original description of M. hercules (based on two females) that the row of mar- ginal spinelets on t. V is interrupted dor- sally. Examination of the lectotype reveals that the dorsal spinelets are not interrupted, but broken off (Figs. 6, 12, black arrows), although those of the paralectotype are in- terrupted (Fig. 9, black arrow). A dorsal in- terruption in the row of marginal spinelets among the 223 specimens (98 6, 125 @) examined was present in only a few speci- mens from three collection localities. Three females from Kosciusko National Park, New South Wales possessed a broad inter- ruption in rows on t. HI-V. No males were examined from that locality. A slight dorsal interruption was also present on t. V of a single male from the Kuipto Forest, just south of Adelaide, South Australia. This male also possesses four deformed setae (much longer than normal spinelets, or combs), two each on one side arising from the base of t. VI and t. VII, respectively. The entire series (8 d, 6 2) from Kangaroo Island (just south of the locality of the spec- imen from Kuipto Forest) possess a dorsal- ly interrupted row on t. V in addition to an interrupted row on t. VI (numbering 5—10 spinelets per side). Excluding the new spe- cies herein described, the rows of marginal spinelets on t-II-V of all specimens exam- ined from Tasmania (22 6, 27 2) were complete, and supernumary setae were not present on t.-VI. Considerable variation ex- ists in the number of marginal spinelets in each row; however, the number of spinelets occurring at, or below the level of the spi- racle on t. V is consistent, ranging from 4— 8 in males and females of M. hercules and O—1 in the new species (Figs. 6—11, white arrows denote position of spiracles). A new unpublished record of M. hercules was examined (National Museum of Natu- ral History, Smithsonian Institution, Wash- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ington, DC) from M. littoralis = M. bur- toni, Arnhem, Smith Point, Caberg Penin- sula, Northern Territory, 25 June 1978, leg. PR. Bavorstock, extending the northern limits of the species to the north central coast of Australia. Rattus fuscipes and R. lutreolus are the usual hosts of this flea, but neither is normally found on the northern coastal areas of Australia [Wilson and Reeder (1993)]. Perhaps these commensal rodents have been introduced by commerce along the coast of the Arnhem region, trans- porting the flea with them. The occurrence of M. hercules on M. burtoni is probably an accidental association with R. fuscipes or R. lutreolus. Macropsylla novaehollandiae Hastriter, new species (Figs: 1—3,.65 7oalSeai>) Type material—AUSTRALIA. ¢ holo- type, 2 allotype, 2 2 paratypes, ex: Pseu- domys novaehollandiae Waterhouse, Coles Bay, Tasmania, 25 November 1998, B. La- zenby; and | 2 paratype, ex: P. novaehol- landiae, Flinder’s Island, Tasmania, 5 Jan- uary 1999, B. Lazenby. Holotype, allotype and two paratypes deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC, and one par- atype in the collection of the senior author. Diagnosis.—Males may be distinguished from M. hercules by the length of the scler- otized inner tube that is obviously longer and wider than the spatulate spiniform of the basimere (Fig. 1), the apico-lateral sur- face of P2 is adorned with 9—10 blunt spi- niform setae (14-19 in M. hercules) (Fig. 4), spinelets of t. V interrupted dorsally (Fig. 6, black arrow), no more than 1—2 marginal spinelets below level of spiracle on t. V (Fig. 6, white arrow), and st. IX is without a distinct sinus in the caudal margin beneath a caudally pointed acute lobe (com- pare Fig. | and 4). Females are very similar to M. hercules, but may be separated by a combination of the following characters: 1) a much weaker lobe along the posterior margin of st. VII, 2) anterior row of setae VOLUME 104, NUMBER 3 Figs. 1-5. 665 Crochet 5 1-3, Macropsylla novaehollandiae (male), Coles Bay, Tasmania. 1, Terminal segments (arrow denotes lack of sinus on sternum IX). 2, Aedeagus (arrows denote dorsal and vental anal lobes). 3, Apex of aedeagus (note relative size of sclerotized inner tube and spatulate spiniform on basimere). 4—5, M. hercules (male), Pearl Beach, New South Wales. 4, Terminal segments with right side and aedeagus removed (P1/P2 = processes | and 2 of basimere; sinus on caudal margin of sternum IX). 5, Terminal aspect of aedeagus (note relative size of sclerotized inner tubes of Figs. 3 and 5). on t. VII much finer and curving strongly caudad below level of spiracle (Fig. 7), 3) dorsal antesensilial bristle #4 or more the length of the middle bristle (only 2 in M. hercules), 4) sclerite in dorsal wall of ovi- duct larger and expanded dorsally (trian- gular vs. rectangular) (Figs. 13—14), and 5) a combination of marginal spinelets on t. V interrupted dorsally with no more than one spinelet extending below level of spiracle (Fig. 7, white arrow). Description.—Head: Clear marginal band running parallel with frons and di- verging dorsad, without setation. Narrow pale marginal band parallel with frons. Six large setae and patch of smaller setae adorn preantennal area in a random pattern in ad- dition to numerous minute scattered setae. Eye well developed, notched ventrally, sit- uated dorsad of angled row of 9 broad spat- ulate genal ctenidia. Anterior arm of ten- torium visible well below eye and anterior PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.2:mm TergunVI 444 0.2 mm triangular cute ~@ rectangular igo ms = 15 ees 17 Figs. 6-17. 6-7, Macrosylla novaehollandiae (male holotype, female allotype, respectively, Coles Bay, Tas- mania). 8-9, M. hercules (female lectotype, female paralectotype, Launceston, Tasmania). 10—11, M. hercules (females), Kangaroo Island, Australia. 12, M. hercules, enlargement of broken teeth noted in Fig. 8. 13, M. novaehollandiae (allotype), sclerite of bursa copulatrix. 14, M. hercules, sclerite of bursa copulatrix. 15, M. novaehollandiae (allotype), spermathecae. 16-17, M. hercules. 16, Spermathecae (lectotype). 17, Spermathecae (Kangaroo Island). VOLUME 104, NUMBER 3 to genal ctenidia, central internal oral tuber arising from oral angle. Antenna of male extending onto prosternum, some setae on pedicel extending about half length of an- tenna. Female antenna not extending to prosternum and longer setae of pedicel ex- tending nearly to apex of antenna. Postan- tennal region with two defined rows of se- tae, with anterior setae somewhat scattered. Row of 10—13 slender setae along dorsal margin of antennal fossa. Distinct suture separating preantennal and postantennal ar- eas. Internal occipital tuber present. Max- illary palpus 5-segmented in both sexes, la- bial palpus 8-segmented in male and 9-seg- mented in female extending just short of apex of coxae. Maxillary lobe sharply acute at apex. Internal apodeme connecting head to thorax extending deep into prothorax. Thorax: Three rows of setae on prothorax and pronotal comb of 15—16 teeth per side in both sexes. Proepisternum with dorsal depression on male to accommodate clavus of antenna, less pronounced in female. Meso- and metanota with 5 rows of setae, each row diminishing in size cephalad. Me- sepimeron and mesepisternum with many scattered setae, pleural rod stout and bifur- cate dorsally. Lateral metanotal area with 7 scattered setae in male and 3 vertical rows (2, 4, 3-4 setae, anterior to posterior) in female. Pleural arch well-developed, dorsal portion heavily sclerotized. Metepisternum bearing 2 moderate setae dorsally, metepis- ternum fusing with that of other side and protruding downward between coxae. Me- tepimeron with 3 vertical rows of setae, posterior stoutest and most dorsal seta be- low level of spiracle. Legs: Lateral surface of forecoxa heavily setose, mesal surface reticulated without setae. Mid- and hind- coxae with sparse small setae along anterior lateral and mesal margins. Oblique sulcus of midcoxa complete. Two femoral pit guard hairs on each femur, each about equal in size on mid- and hindfemora, medial smaller on forefemur (see Remarks). Nu- merous lateral setae found on all femora and only 2—3 setae on mesal surface of 667 each. Each femur with coarse surface retic- ulations. Lateral bristles guarding femoral- tibial joints longest of pair on forefemur and shortest on mid- and hindfemora. Dor- sal margin of foretibia adorned with many heavy long spines (one nearly reaching apex of second tarsal segment) resembling a false comb; mid- and hindtibiae with 7 and 8 dorsal notches, respectively, with ad- ditional setae interspersed between notches. Mesal surface of mid- and hindtibiae sculp- tured with triangular scalelike reticulations. Hindtibia with apical tooth. Segment I of hindtarsus about as long as hindtibia and segment II longer than segments III and IV combined. Fifth tarsal segment of each leg adorned with a complex set of lateral plan- tar bristles; first pair set onto plantar surface positioned between second pair, third and fourth pairs each with two bristles per side. Each plantar surface with many minute se- tae and 2—4 (variable) preapical plantar bristles (some spiniform). Unmodified ab- dominal segments: Chaetotaxy of t. I-VII variable. Only posterior main rows with in- tercalary setae. Tergites I-VI of male bears 3, 4, 4, 4, 4, and 3 rows, respectively and t. VII with a single defined row and nu- merous smaller setae scattered anteriorly. Female with similar pattern, but anterior rows less defined. Both sexes with marginal spinelets on t. H—V with row of spinelets interrupted dorsally on t. V. These total 46, 44, 36, and 30 (male), and 49, 44, 44-45, and 28-31 (female), respectively. Number of spinelets below level of spiracle on t. II— IV 7, 3, and 3 (male) and 6-8, 1—5, and 0— 2 (female). Those situated below level of spiracle on t. V range from O to 2 in both sexes. Both sexes with 3 antesensilial bris- tles, ventral pair of equal length, and dorsal bristle % as long as others. Abdominal st. II with a lateral patch of small setae (more numerous in female), and a single ventral seta per side; male with 5—7 stout setae per side in main rows and numerous scattered setae anterior to main row on st. III—VII, female same for st. II-VI. Modified ab- dominal segments (male): Tergum VIII re- 668 duced, covering little of terminal segments and bearing patch of fine setae across dor- sum. Atrium of eighth spiracle greatly ex- panded, reminiscent of some species of Me- gabothris Jordan 1933. Basimere with two lobes (PI and P2). P2 is highly modified immovable apical process (anatomically in usual position of the telomere) which bears 9—10 blunt spiniforms on mesal surface and single unique spatulate spiniform arising from sinus between the PI and P2 (Figs. 1, 4). Located just below sensilium on basi- mere, 2 structures, which appear as vestigial telomeres, each with several slender setae apically (Fig. 3, arrow). Anterior portion of basimere heavily sclerotized. Sternum VIII with group of 9—10 stout setae per side near caudal margin with numerous smaller setae scattered over surface. Distal arm of st. [IX uniquely fused into single structure (paired in most other genera of fleas) bifurcating at base into usual proximal arms. Apex of fused distal arms adorned with 6 spiniform setae and numerous stout setae. Caudal margin with beak-like process with margin below entire without indication of sinus (Fig. 1, arrow). Dorsal anal lobe studded with blunt spiniforn setae, ventral anal lobe bearing 2 long slender setae (Fig. 2, ar- rows). Many portions of aedeagus more highly sclerotized than those of other fleas, particularly sclerotized inner tube, crescent sclerite, crochet, and medial and _ lateral lobes of fulcrum. Modified abdominal seg- ments (female): Three antesensilial bristles, dorsal bristle about % length of middle bris- tle. Tergum VIII greatly expanded ventrally so as to envelop vestigial (membranous) st. VIII, covered with many setae, and those towards caudal margin more robust than those more anterior. Caudal margin forming angular lobe. Sensilial plate projecting an- terior to caudal margin of t. VII, interrupt- ing dorsal portion of t. VIII. Dorsal anal lobe much larger than inconspicuous re- duced ventral anal lobe. Anal stylet nearly three times as long as wide, bearing single minute seta at base of long apical seta. Ster- num VII without lobes or sinuses and st. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VIII vestigial. Paired spermathecae similar in size and much like those of M. hercules (Figs. 15-17). Hyperdeveloped trianglular sclerite in bursae copulatrix, rectangular in M. hercules (Figs. 13-14). Size: Slide mounted specimens: male, AQMimim @ = 1), female. 45> mmi@e— 2); Alcohol specimens: female, 3.3 mm (n = 2) Etymology.—This species is named after the specific name of the host species, Pseu- domys novaehollandiae, as a noun in ap- position. Remarks.—Nowak and Paradiso (1983) indicated that the New Holland mouse, P. novaehollandiae was limited to a few col- lections early in the 1900s from New South Wales to Mornington Peninsula of southern Victoria and from subfossil remains found 1976 when Hocking (1980) reported a population on the north- eastern coast of Tasmania. Macropsylla in Tasmania until hercules is commonly found throughout eastern Australia from Queensland to South Australia and Tasmania on a variety of mu- rid hosts, none of which include P. novae- hollandiae. The rarity of collecting P. no- vaehollandiae (to include its ectoparasites) probably accounts for the failure to previ- ously find this new species of flea. Addi- tonal collection of this endangered host spe- cies on mainland Australia will predictably yield additional specimens of M. novaehol- landiae. A pit located on the ventral anterior mar- gin of the femora of each leg has previously not been reported. Although the function of this pit is not known, it appears to be pre- sent on most, if not all species of fleas. Dor- sad to the “pit’” are two stout setae that project downward over the “‘pit” so as to shield the underlying “‘pit” structure. The size and arrangement of these setae are var- iable from species to species and may be consistent among different families. The details and significance of this new obser- vation will be published elsewhere. VOLUME 104, NUMBER 3 DISCUSSION Oudemans (1909) placed Macropsylla in a separate subfamily (Macropsyllinae) and family (Macropsyllidae) based on the pres- ence of an internal occipital tuber and Hop- kins and Rothschild (1956), ignoring the subfamily, followed the same scheme and included a second monotypic genus, Stfe- phanopsylla Rothschild, in this family. Smit (1982) astutely did not recognize this ar- rangement and placed Macropsyllinae in the family Hystrichopsyllidae (it is pre- sumed that he also included Stephanopsyl- la, since there was mention of six genera in his family discussion). Lewis (1993, 1998) also followed this arrangement. Several ob- servations substantiate the current system- atic arrangement. Characters commonly shared by Macropsylla and other genera of the Hystrichopsyllidae include: 1) nearly a complete fusion of the distal arms of st. IX from base to apex; 2) presence of dual sper- mathecae; 3) combs or spinelets on one or more abdominal tergites; 4) anterior branch of tentorium visible; 5) apical tibial spur present; 6) mesopleural rod_ bifid (only slightly in Typhloceras Wagner); and 7) common pattern of femoral-tibial joint bris- tles (protibia, outer long, inner short, and meso- and metatibia, outer short, inner long). Smit (1982) indicated the absence of the anterior arm of the tentorium as a char- acter to distinguish genera of the subfamily Macropsyllinae from those of Hystrichop- syllinae. Although the tentorium does not extend as far anteriorly in Macropsylla as it does in Atyphloceras Jordan and Roths- child, Ctenoparia Rothschild, Hystrichop- sylla Taschenberg, and Typhloceras, it is visible. Specimens of Stephanopsylla tho- mast (Rothschild) were not examined; how- ever, illustrations of this monotypic genus appear morphologically similar to Macrop- sylla except for extreme modifications in the structure of the head. These similarities include an angular genal comb, mouth parts placed well behind the oral angle (lesser in Macropsylla), presence of abdominal 669 combs on t. H—V (Macropsylla) and t. I-VI (Stephanopsylla), presence of a dorsal oc- cipital tuber, and a frons composed of a nar- row frontal ridge separated by a narrow band (in lateral view) with a uniform row of minute setae along its posterior margin. These characters distinguish members of the Macropsyllinae from those of Hystri- chopsyllinae. Much emphasis has been placed on the importance of the occipital tuber (Oude- mans 1909, Hopkins and Rothschild 1956). In addition to taxa belonging to Macrop- syllinae, this structure is present in all gen- era of Stephanocircidae (minimally so in Barreropsylla Jordan). Wagner (1934) am- ply (and accurately) described the occipital tuber and other structures within the head capsule, using the stephanocircine Plocop- sylla enderleini Wagner as a model. The na- ture of the occipital tuber can only be un- derstood when viewed from a frontal or dorsal view (dissected specimens). The conventional lateral view of this structure might suggest that the apodeme is project- ing centrally into the middle of the head, which is not the case. The structure is com- prised of a heavily sclerotized ridge which follows the contour of the occiput adding rigidity and strength to the head capsule. The term occipital bridge might be more descriptive than occipital tuber. It is not sur- prising that sephanocircids possess it, since the structure of the head is highly modified to accommodate the helmet found in no other fleas [except the unrelated Smitella thambetosa (Pygiopsyllidae)]. Traub (1968) explained this convergent evolutionary con- dition in detail. Some members of Ischnop- syllidae also possess highly modified in- crassations on the occipital areas of the head. One can only speculate why the oc- cipital tuber occurs in relatively unrelated genera whose basic head _ structures are quite different. The peculiar shape of the frons in Macropsyllinae is reminiscent of some members of the family Ishnopsylli- dae, but this feature is likely an indepen- dently convergent development. Hopkins 670 and Rothschild (1956) suggested the Is- chnopsyllidae (and Stephanocircidae) might well have been descendents of “‘Macrop- syllidae.’’ Phylogenetic analysis of DNA sequence data does not support a close re- lationship between the Ischnopsyllidae and Macropsyllinae. These observations will be discussed in a future paper. Both genera in the subfamily Macrop- syllinae are restricted to the Australian sub- region of the Australian Region. The iso- lated evolution of hystrichopsyllids (Ma- cropsylla and Stephanopsylla) in Australia is puzzling considering the disjunct distri- bution of other hystrichopsyllids (Atyphlo- ceras, Ctenoparia, Hystrichopsylla and Ty- phloceras). With exception of Ctfenoparia, other genera of Hystrichopsyllidae are bo- real inhabitants occurring in the Holarctic Region (Hystrichopsylla, Atyphloceras) and in the Palaearctic Region (7yphloceras). Although Ctenoparia and the two Austra- lian genera share familial characteristics and early common geographical connec- tions (Gonwanaland), they are quite dissim- ilar. It is evident that they were isolated very early or they were introduced by com- mensal murid rodents via human oceanic travels. Jordan (1921) suggested that the differ- ence in sizes of the two spermathecae of Macropsylla might be an evolutionary in- termediate link between those genera pos- sessing dual spermathecae and those with only one. Illustrations (Dunnet and Mardon 1974) of the female of Stephanopsylla tho- masi and all specimens of Macropsylla that we examined demonstrate significant dis- parity in the shape and size of the paired spermathecae, whereas those of other gen- era (excluded from Australia) are consis- tently similar. Traub (1980) provided evi- dence that the Australian Hystrichopsylli- dae (Macropsylla and Stephanopsylla) are the most ancient genera in the family. If this proves to be the case, one would expect the more primitive genera of the Hystrichop- syllidae (Macropsylla and Stephanopsylla) to possibly manifest greater morphological PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON variability than more recently evolved taxa (Atyphloceras, Ctenoparia, Hystrichopsyl- la, and Typhloceras). Other than exteme modification of the head, Stephanopsylla is more similar to other hystrichopsyllids than Macropsylla. These similaries are noted particularly in the general features of t. IX, st. VIII, and the aedeagus. The extreme modification of the head of Stephanopsylla may be an evolutionary adaptation to its marsupial host species (Dasyuridae Gold- fuss and Potoroidae Gray). Stephanopsylla thomasi has a peculiar disjunct distribution in Australia with a rather small number of specimens recorded from Western Australia and from Victoria, Australia. Additional collections are needed to elucidate the geo- graphical distribution of Stephanopsylla thomasi. The apparent preference of Macropsylla for murid rodents (R. fuscipes and R. lu- treolus) is yet another indication of the sys- tematic relationship to the Hystrichopsylli- dae, as other genera in this family all share murid rodents as their preferred hosts. Dun- net and Mardon (1974) reported Australian fleas from 48 different genera of marsupials and only 12 genera of murid rodents. Re- cords of Macropsylla occurring on marsu- pials are undoubtedly accidental associa- tions. Divergence of the ancestral hystri- chopsyllid likely occurred after introduction into Australia on murid rodents, some evolving after a successful transition to marsupial hosts (Stephanopsylla) while oth- ers continued evolving on murids (Macrop- sylla). ACKNOWLEDGMENTS We acknowledge the unselfish assistance of Menna Jones, Division of Botany and Zoology, Canberra, Australia, for coordi- nation of field work and Billie Lazenby and Terrance Pye, School of Zoology, Univer- sity of Tasmania, Hobart, Tasmania, for their efforts in collecting the fleas for which this work would have otherwise not been possible. For their generous loan of speci- mens, we are indebted to Nancy Adams, VOLUME 104, NUMBER 3 National Museum Natural History, Smith- sonian Institution, Washington, DC; Gra- ham Crompton, Manager, Australian Na- tional Insect Collection, Canberra, Austra- lia; Ralph P. Eckerlin, Northern Virginia Community College, Annandale, VA; The- resa Howard, The Natural History Museum, London, England; and John Rawlins, Car- negie Museum of Natural History, Pitts- burgh, PA. Funding for this publication was provided by a grant from the National Sci- ence Foundation, Award Number DEB- 9983195: LITERATURE CITED Dunnet, G. M. and D. K. Mardon. 1974. A monograph of Australian fleas (Siphonaptera). Australian Journal of Zoology, Supplementary Series No. 30, 273 pp. Hocking, G. J. 1980. The occurrence of the New Hol- land mouse Pseudomys novaehollandiae (Water- house), in Tasmania. Australian Wildlife Research 7: 71-77. Hopkins, G. H. E. and M. Rothschild. 1956. An Illus- trated Catalogue of the Rothschild collection of fleas (Siphonaptera) in the British Museum. Vol. II, The British Museum (Natural History), Lon- don, 445 pp. Jordan, K. 1921. A link between the double and single receptacula seminis of Siphonaptera. Ectoparasites 1: 127-128. Lewis, R. E. 1993. Notes on the geographical distri- bution and host preferences in the Order Siphon- aptera. Part 8. New taxa described between 1984 and 1990, with a current classification of the order. Journal of Medical Entomology 30(1): 239-256. 671 . 1998. Résumé of the Siphonaptera (Insecta) of the world. Journal of Medical Entomology 35(4): 377-389. Nowak, R. M. and J. L. Paridiso. 1983. Walker’s mam- mals of the world. 4th Edition. Johns Hopkins University Press, Baltimore 2: 709-713. Oudemans, A. C. 1909. Neue ansichten tiber die mor- phologie des flohkopfes, sowie tiber die ontogen- ie, phylogenie und systematik der fl6he. Novitates Zoologicae 12: 133-158. Rothschild, M. and R. Traub. 1971. A revised glossary of terms used in the taxonomy and morphology of fleas. Trustees of The British Museum (Natural History), London, 85 pp. Rothschild, N. C. 1905. Some new Siphonaptera. Nov- itates Zoologicae 12: 479-491. Smit, EF G. A. M. 1982. Siphonaptera, pp. 557-562. In Parker, S. P., ed. Synopsis and classification of liv- ing organisms, Vol. 2. McGraw Hill, New York, 1,232 pp. Traub, R. 1968. Smitella thambetosa, n. gen. and n. remarkable “‘helmeted”’ flea New Guinea (Siphonaptera, Pygiopsyllidae) with notes sp., a from on convergent evolution. Journal of Medical En- tomology 5(3): 375—404. 1980. The zoogeography and evolution of some fleas, lice and mammals, pp. 93-172. In Traub, R. and H. Starcke, eds. Fleas. Proceedings of the International Conference on Fleas. Ashton Wold/Peterborough, United Kingdom. A.A. Bal- kema, Rotterdam, The Netherlands, 420 pp. Wilson, D. E. and D. M. Reeder. 1993. Mammal spe- cies of the world, a taxonomic and geographic ret- erence. 2nd Edition. Smithsonian Institution Press, Washington, DC, 1,206 pp. Wagner, J. sogenannten “caput fractum”’ bei insecten (ber den kopfbau der ““helmtragenden” fl6he). Zoolo- gischer Anzeiger 106(1/2): 7-15. 1934. Weitere beitrage zur auffassung des PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 672-679 DESCRIPTION OF FEMALES OF STETHOPHOTOPSIS PITTS AND SPHAEROPTHALMA (PHOTOPSIOIDES SCHUSTER) (HYMENOPTERA: MUTILLIDAE) JAMES P. PITTS AND DONALD G. MANLEY (JPP) Department of Entomology, University of Georgia, Athens, GA 30602, U.S.A. (e-mail: jpitts@bugs.ent.uga.edu); (DGM) Department of Entomology, Clemson Univer- sity, Pee Dee Research and Education Center, Florence, SC 29506 Abstract.—The female of Stethophotopsis maculata Pitts, Stethophotopsis cremauranti, n. sp., and Sphaeropthalma (Photopsioides) uro (Blake) are described. These represent the first descriptions of the females of Stethophotopsis Pitts and Photopsioides Schuster. Key Words: Uy Nocturnal mutillids of the southwestern United States have a high diversity, with about 206 known species. Anyone black lighting in the Southwest during the sum- mer is awed by the number of male mutil- lids that are attracted to the lights. Females of these nocturnal species, however, are rarely collected. The females of many of the nocturnal southwestern Mutillidae species remain unknown. Only five species have been described from both male and female specimens. There are an additional 22 spe- cies that are known solely from the females. This leaves 176 species described from males. Some genera and subgenera, such as Acanthophotopsis Schuster, Acrophotopsis Schuster, and Odontophotopsis Viereck, lack a single female association and remain known only from the male. Female associ- ations of these taxa would provide valuable data for generating a phylogenetic hypoth- esis of the sphaeropthalmine mutillids. This paper adds to our knowledge of the females of some of these nocturnal species. A pair of Stethophotopsis maculata Pitts collected in copula was found among the undetermined mutillids in the University of Arizona Collection of Arthropods. The fe- Insecta, Sphaeropthaminae, wasp, velvet ant, new species male represents the first female known for this genus. A second male was found dou- bling the known number of male speci- mens. It was noted that a congeneric species of female also existed. Examination of both species greatly helped in determining fe- male characters of generic level importance. These two species are described, illustrated, and discussed below. During a subsequent collecting trip near Portal, Arizona, a pair of mutillids was col- lected in copula at a black light. The male was identified as Sphaeropthalma (Photop- sioides) uro (Blake). This paper provides the first description of a female of this sub- genus. The female of Photopsioides Schus- ter is described, illustrated and discussed below. MATERIALS AND METHODS We follow Schuster’s (1958) classifica- tion of the sphaeropthalmine genera. Cur- rently, there is no phylogenetic hypothesis available for the subtribe Sphaeropthalmi- na. It is apparent that Sphaeropthalma is not monophyletic (pers. obs.), but reclassi- fication of this group should not be done until more females are known. As the genus VOLUME 104, NUMBER 3 now stands, the subgenus Photopsis Blake includes females that differ in characters that are considered to be of generic impor- tance, such as a defined pygidium and a pet- iolate abdomen. The following acronyms are used for col- lections that were involved in this study: Canadian National Collection of Insects, Ottawa, Canada (CNCI); Department of Entomology Collection, University of Ari- zona, Tucson (UAIC); James P. Pitts collec- tion (JPPC). After Ferguson (1967), we are adopting the following notation for punctation in the order of decreasing coarseness: reticulate, coarse, moderate, small, fine and micro- punctate. Micropunctate refers to punctures that are extremely shallow and do not have vertical walls or sharp margins. We have used the term simple pubescence for hairs that are smooth and do not have barbed sur- faces. Brachyplumose pubescence refers to hairs with barbs that are less than or equal to the diameter of the shaft at the attach- ment of the barb. Plumose pubescence is used for hairs that have longer barbs. T2, iS retcs .arenused)itovidenote the ‘second, third, etc., metasomal tergites while S2, S3, etc., denote the second, third, etc., metaso- mal sternites. Stethophotopsis Pitts (Bigs: 1,4) 7, 9... 10) Type species.—Stethophotopsis macula- ta Pitts. Orig. desig. Female.—Head: Narrower than thorax. Eyes projecting, subcircular, clearly faceted. Clypeus anterior edge truncate, base swol- len medially. Dorsal carina of antenna pre- sent but inconspicuous or absent. Antennal tubercles well developed and subcontig- uous. Antenna 12-segmented. Pedicel and first flagellomere subequal in length (Fig. 1). Mandible with internal tooth on distal one-third (Fig. 4). Ventral margin of man- dible weakly emarginate at basal one-third (Fig. 4). Genal carina absent. Proboscidal furrow triangular, broad, reaching base of mandibles and margined by carinae. Max- 673 illary palpus 6-segmented, labial palpus 4- segmented. Mesosoma: Pyriform, slightly longer than wide (Fig. 7). Mesosoma gradually widening from humerus to anterior spirac- ular tubercles, widest at anterior spiracular tubercles, gradually narrowing from ante- rior spiracular tubercles posteriorly (Fig. 7). Scutellular scale absent (Fig. 7). Propo- deum with separation between dorsal and lateral regions (Fig. 9). Lateral face of pron- otum narrow, punctate, with anterior de- pression. Anterior half of mesopleuron gla- brous. Metapleuron glabrous. Spurs of all tibiae pectinate. Metasoma: First segment distinctly pet- iolate with respect to second (Fig. 10). T1 attached ventrally to midline of T2 in lateral view such that T2 bulges above attachment (Fig. 10). T2 with lateral felt lines (Fig. 10). S2 strongly convex and without lateral felt lines. T6 with pygidial area undefined. Plu- mose pubescence present on apical margin of T2. Coarsely punctate and raised inte- gumental maculations present on T2. Distribution.—Southern Arizona, USA; Sonora, Mexico. Comments.—Females of Stethophotopsis can be distinguished from females of other sphaeropthalmine genera by the combina- tion of presence of plumose pubescence on the metasoma, absence of dorsal carinae on the antennal scrobes, first antennal flagel- lomere as broad as long, absence of genal carinae, first metasomal segment petiolate, condition of the petiolar attachment, pres- ence of coarsely punctate and raised inte- gumental maculations on the second meta- somal segment, and absence of lateral ca- rinae defining pygidium. Stethophotopsis is a distinct genus in the Sphaeropthalmini (subtribe: Sphaeropthal- mina). For males, the unique sternal pro- cesses and the condition of the cuspis of the genitalia, being dilated, slightly spatulate, and elongated (attaining the length of the parameres), are apparently autapomorphic for the genus. It remains unclear at this point what characters, if any, are autapo- 674 morphic for females of this genus. Stetho- photopsis males and females will key to subfamily Sphaeropthalminae without dif- ficulty in existing keys by Brothers (1993, 1995). In Schuster’s (1958) key to the sphaeropthalmine males of the North Amer- ican Southwest, Stethophotopsis terminates at couplet 1, where it can be distinguished by the autapomorphies listed above. Cur- rently, there is no key for the sphaeropthal- mine females. Stethophotopsis maculata Pitts (Piss 1. 4.7, 9:0) Female.—Length: 7.5 mm. Head: Head brownish yellow, clothed with sparse erect and dense decumbent white simple pubescence. Malar space 0.6 x maximum eye width. First flagellomere 1.1 length of pedicel (Fig. 1). Second fla- gellomere slightly longer than first, 1.4 length of pedicel (Fig. 1). Scape and pedicel concolorous with head. Flagellum dark brown. Front with confluent punctation. Vertex narrowly reticulate. Mandible brown distally. Mesosoma: Mesosoma and legs brown- ish yellow, except femora and tibiae dark brown. Pronotum and mesonotum with de- cumbent and short erect dark brown pubes- cence. Metanotum with decumbent white brachyplumose pubescence. Propodeum with white pubescence, somewhat darkened anteriorly, and with long erect and short plumose pubescence posteriorly. Legs with white brachyplumose pubescence. Humeri angulate. Lateral face of pronotum punctate with decumbent white pubescence. Meso- pleuron with posterior half punctate with erect white pubescence; anterior half gla- brous. Propodeum glabrous laterally; retic- ulate on lateral margins. Dorsum of meso- soma reticulated. Metasoma: Yellowish brown. T1 with sparse long erect white brachyplumose and short plumose pubescence. T1 with apical fringe of white plumose pubescence. T2 with two round black integumental macu- lations on anterior fourth with deep punc- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tation, reticulate surface, raised above sur- rounding disk, with thick black simple pu- bescence. T2 felt line 0.2X length of T2. T2-T5 with golden brachyplumose pubes- cence except black in areas of tergal mac- ulations. T2 with golden-white plumose apical fringe. Sternites with white brachy- plumose pubescence. S2 with median lon- gitudinal tumid region on anterior fifth. Material examined.—Mexico, Sonora, 5 mi east of Alamos, 1 2 1 6, 11.VIII.1973, coll. K. Stephan and D. S. Chandler (UAIC); 1 36, Mexico, Sonora, 5 mi east of Alamos, 11.VIII.1973, coll. K. Stephan and D. S. Chandler (UAIC). Comments.—One of the males and the female are mounted on the same pin and have an attached label stating that the pair was collected in copula. The two males dif- fer from the holotype and paratype by being smaller (5 mm versus 7 mm) and having the integument around the ocelli darkened. Stethophotopsis cremauranti Pitts and Manley, new species Female.—Length: 9 mm. Head: WHead orangish brown, clothed with sparse erect and decumbent pale gold- en brachyplumose pubescence. Malar space 0.7X maximum eye width. First flagellom- ere 1.2 length of pedicel. Second flagel- lomere slightly shorter than first, 1.1 length of pedicel. Antenna dark brown. An- tennal scrobe inconspicuously carinate dor- sally. Front and vertex densely punctate. Mandible brown distally. Mesosoma: Mesosoma, trochanters, and coxae orangish brown. Other segments of legs dark brown. Pubescence of pronotum, mesonotum, and propodeum anteriorly pale golden brown; erect pubescence simple; de- cumbent pubescence brachyplumose. Pu- bescence of propodeum anteriorly pale golden and posteriorly white. Pubescence of legs white. Lateral face of prontoum punctate with sparse erect pale golden pu- bescence. Mesopleuron posterior half punc- tate with erect pale golden pubescence. Pro- podeum with posterior lateral half punctate; VOLUME 104, NUMBER 3 Figs. 1-8. nN i 1—3, Apical end of scape and first three flagellomeres of female antenna. 4—6, Mandible of female, dorsal view; arrow indicate ventral emargination. Figs. 7—8, Thorax, dorsal view; arrow indicate widest point. 1, 4, 7, Stethophotopsis maculata. 2, 5, 8, Sphaeropthalma (Photopsioides) uro. 3, 6, Sphaeropthalma (Sphaer- opthalma) pensylvanica. anterior half glabrous. Dorsum of mesoso- ma with coarse punctation. Metasoma: Dark brown. T1 with long erect white brachyplumose and short plu- mose pubescence. Tl with apical fringe of white plumose pubescence. T2 felt line 0.3 length of T2. T2 with two round black integumental maculations on anterior fourth with larger and deeper punctation, appear- ing slightly raised above surrounding disk. Maculations with decumbent and erect bra- chyplumose pubescence. T2—T5 with white brachyplumose pubescence except dark brown in areas of tergal maculations. T2— 676 T3 with sparse white plumose apical fringe. Sternites with white brachyplumose pubes- cence. S2—S3 with sparse white plumose pubescence on lateral apical margin, pubes- cence simple on median three fourths. S2 with median longitudinal tumid region on anterior fifth. Material examined.—Holotype 2, Mex- ico, Chihuahua, 3 mi east of Parral, 5800’, 30.1V.1953, coll. Creighton (CNCI). Para- type, | 2, Mexico, Chihuahua, 21 mi south of Parral, 5600’, 30.1V.1953, coll. Creigh- ton (CNCI). Etymology.—From the Latin cremo meaning “‘to burn to ashes” and auranti meaning “orange” in reference to the color of the head and thorax. Comments.—The paratype is only 7.5 mm in length, but the coloration and punc- tation of both specimens are similar. Steth- ophotopsis cremauranti differs from S. ma- culata in being slightly larger in size. The coloration of the two species is very differ- ent. The head and mesosoma of S. mauranti are slightly darker orange than the head and mesosoma of S. maculata. Steth- ophotopsis cremauranti, however, has the metasoma completely dark brown. The me- tasoma of S. maculata is dark brown only in the areas of the tergal maculations. Fur- thermore, S. cremauranti has an inconspic- uous carina dorsally on the antennal scrobe, which is totally lacking in specimens of S. maculata. Also, the T2 maculations are well developed in S. maculata due not only to the fact that there are deep, raised punc- tures making the maculations appear raised above the surrounding area, but also be- cause there is very thick, black, decumbent pubescence present. In specimens of S. cre- mauranti, neither deep, raised punctation nor thickened pubescence is present. Creé- Sphaeropthalma (Photopsioides Schuster) (Biles5 2 Sapo. elalel) Type species.—Agama uro Blake. Orig. desig. Female.—Head: As wide as thorax. Eyes projecting, subcircular, clearly faceted. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Clypeus anterior edge truncate, base tuber- culate medially. Antennal scrobes carinate dorsally. Antennal tubercles well developed and subcontiguous. Antenna 12-segmented. First flagellomere length 1.8 length of pedicel (Fig. 2). Mandible with strong in- ternal tooth on distal one-third (Fig. 5). Ventral margin of mandible strongly emar- ginate at basal one-third (Fig. 5). Gena well developed, without genal carina. Probosci- dal furrow triangular, broad, margined by carinae, becoming weak near base of man- dibles. Maxillary palpus 6-segmented, la- bial palpus 4-segmented. Mesosoma: Pyriform, slightly longer than wide (Fig. 8). Mesosoma widest an- teriorly, gradually narrowing just posterior to anterior spiracular tubercles (Fig. 8). Scutellular scale absent (Fig. 8). Propo- deum with distinct dorsal and lateral re- gions (Fig. 11). Lateral face of pronotum narrow and punctate. Mesopleuron punc- tate. Metapleuron glabrous. Metapleuron separated from mesopleuron by depression. Tibial spurs pectinate. Metasoma: First segment subsessile with respect to second (Fig. 12). T2 with lateral felt lines (Fig. 12). S2 strongly convex, base prominently raised medially. S2 with- out lateral felt lines (Fig. 12). T6 with py- gidial area undefined. Plumose pubescence present on apical margins of metasomal segments. Distribution.—Southern Arizona, USA. Comments.—Since only one species is known, it is difficult to diagnose the generic level characters. Some characters such as the presence or absence of a dorsal carina on the antennal scrobe may not be valid. Photopsioides is similar to the subgenus Sphaeropthalma by the following: presence of plumose pubescence, presence of dorsal carinae on antennal scrobes, first antennal flagellomere 1.8 as long as pedicel (Figs. 2, 3), and absence of lateral carinae that de- fine the pygidium. The females of Photop- sioides can be distinguished from Sphaer- opthalma by having a slight tooth on the ventral base of the mandible (Fig. 5) (not VOLUME 104, NUMBER 3 Figs. 9-14. opthalma (Sphaeropthalma) pensylvanica. 9, 11, 13, Thorax, lateral, dotted line indicates impunctate area within. 10, 12, Metasoma, lateral. 14, First metasomal tergite and basal portion of second metasomal tergite, lateral. strongly developed as in Sphaeropthalma s.s. (Fig. 6)), by the length of the propo- deum in lateral view being subequal in length to 0.5 the maximum height (Fig. 11) (not ~1X as in Sphaeropthalma s.s. (Fig. 13)), having a petiolar attachment that is more clearly petiolate (Fig. 12) than Sphaeropthalma s.s. (Fig. 14), and having sparse plumose pubescence on the apical margins of metasomal segments (not having 677 9-10, Stethophotopsis maculata. 11-12, Sphaeropthalma (Photopsioides) uro. 13—14, Sphaer- only a median band of dense appressed plu- mose pubescence on the apical margin of metasomal segment one (for Sphaeropthal- ma pensylvanica (Lepeletier)) or segments one and two and a median propodeal stripe (for Sphaeropthalma auripilis (Blake)). Males of Photosioides are similar to those of the subgenus Sphaeropthalma. Both have the pygidium and hypopygium short and truncated, lack sternal felt lines, 678 and the cuspis of the genitalia is dilated and covered with plumose tipped setae (Schus- ter 1958). Males of Photopsioides differ from those of the subgenus Sphaeropthal- ma by being nocturnal, weakly sculptured, and having sparse pubescence on the ven- tral margin of the parameres (Schuster 1958); Sphaeropthalma (Photopsioides) uro (Blake) (Figs. 2; 35,8521, 12) Female.—Length: 9-10 mm. Head: Head orangish brown, clothed with sparse erect pale white brachyplumose pubescence. Malar space 0.6 maximum eye width. First flagellomere 1.8 length of pedicel (Fig. 2). Second flagellomere length equal to length of pedicel (Fig. 2). Antenna orangish brown. Front and vertex with small dense punctures. Mandible reddish black apically. Mesosoma: Mesosoma orangish brown. Legs slightly darker than mesosoma. Pu- bescence of pronotum, mesonotum and me- tanotum pale golden brown. Propodeum with white brachyplumose pubescence, plu- mose posteriorly. Humeri angulate. Lateral face of prontum punctate with sparse erect pale white pubescence. Mesopleuron punc- tate with erect pale white pubescence. Me- tapleuron glabrous. Propodeum_ punctate laterally. Dorsum of mesosoma_ coarsely punctate. Propodeum surface narrowly re- ticulate posteriorly. Metasoma: Orangish brown. T3—T6 slightly darker than Tl and T2. Tl with erect brachyplumose pubescence and short- er erect white plumose pubescence. T1 with apical fringe of white plumose pubescence. T2 felt line 0.3X length of T2. T2 with brown erect subplumose pubescence, white on apical margin. T3—T5 with white pubes- cence. T2—T4 and S2-—S5 with dense white plumose apical fringe. S2—S5 with simple white pubescence. T2 surface reticulate an- teriorly, coarsely punctate posteriorly. T3— T6 finely punctate. SI coarsely punctate with white plumose pubescence. S2 with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON median longitudinal tumid region on ante- rior fifth, coarsely punctate. S3—S6 finely punctate. Material examined.—Arizona, Cochise Cony 3ani “north ‘of yPRortaly4 lice gee 3.VIIE.1999, coll. J.P. Pitts (JPPC); Cochise Co., 2 mi north of Portal, San Simon Road, 1 oe Vit 2000; colif-P.Ritts GPPG@)xGo- chise €o., Portal, 1 2,°23.°V1.1969; Collny; D. Roth (UAIC); Pima County, 8 mi north of Vail, 1 2, 30.VIII.1962, coll. E Werner (UAIC). Comments.—The specimens differ in size slightly. One specimen is darker be- neath the felt lines. ACKNOWLEDGMENTS We are indebted to Carl Olsen (UAIC) and John Huber (CNCI) for loan of mate- rial. The following institutions are hereby acknowledged for funding (JPP): the CNCI for a CanaColl grant allowing JPP to travel to the CNCI, Ottawa, Canada, in 1999 and the American Museum of Natural History for the Theodore Roosevelt Memorial Fund for travel to the Southwestern Research Sta- tion, Portal, Arizona, during 2000. We thank Wade Sherbrooke of the Southwest- ern Research Station for use of their facil- ities. We thank Joseph McHugh (University of Georgia), Cecil Smith (University of Georgia), and Theresa Pitts-Singer (Univer- sity of Georgia) for critically reviewing the manuscript. LITERATURE CITED Brothers, D. J. 1975. Phylogeny and classification of the aculeate Hymenoptera, with special reference to Mutillidae. University of Kansas Science Bul- letin 50: 483-648. 1993. Family Mutillidae, pp. 188-203. In Goulet, H. and J. T. Huber, eds. Hymenoptera of the World: An Identification Guide to the Fami- lies. Centre for Land and Biological Resources Research, Ottawa, Ontario. Research Branch, Ag- riculture Canada, Publication 1894/E. 668 pp. . 1995. Mutillidae, pp. 541-548. /n Hanson, P. E. and I. D. Gauld, eds. The Hymenoptera of Cos- ta Rica, Oxford University Press, Oxford, 893 pp. . 1999. Phylogeny and evolution of wasps, ants, VOLUME 104, NUMBER 3 and bees (Hymenoptera, Chrysidoidea, Vespoidea, and Apoidea). Zoologica Scripta 28: 233-249. Ferguson, W. E. 1967. Male sphaeropthalmine mutillid wasps of the Nevada Test Site. Brigham Young Uni- versity Science Bulletin, Biological Series 8: 1—26. Lelej, A. S. and P. G. Nemkov. 1997. Phylogeny, evo- lution and classification of Mutillidae (Hymenop- tera). Far Eastern Entomologist 46: 1—24. 679 Pitts, J. PB and J. V. McHugh. 2000. Stethophotopsis, a new genus of Sphaeropthalmini (Mutillidae: Sphaeropthalminae) with a brachypterous male from Arizona. Journal of Hymenoptera Research 9: 29-33. Schuster, R. M. 1958. A revision of the sphaeropthal- mine Mutillidae of America north of Mexico. II. Entomologica Americana 37: 1-130. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 680-686 REDESCRIPTION OF TWO SPECIES OF OMMATIUS WIEDEMANN, WITH LECTOTYPE AND PARALECTOTYPE DESIGNATIONS FOR OMMATIUS TENELLUS VAN DER WULP AND RANGE EXTENSION, AND A REPLACEMENT NAME FOR OMMATIUS TIBIALIS RICARDO (DIPTERA: ASILIDAE) A. G. SCARBROUGH Department of Biology, Towson University, 8000 York Road, Baltimore, MD 21252, U.S.A. (e-mail: ascarbrough @towson.edu) Abstract.—The type series of Ommatius tenellus van der Wulp and O. tibialis Ricardo were examined and redescriptions are given for both species. A lectotype and paralecto- types are designated for O. tenellus, new records extend the range of the species across northern Africa, and illustrations of the terminalia are presented. Ommatius abdelkurien- sis Scarbrough (new name) is proposed as a replacement name for O. tibialis. Key Words: As a part of my revisionary studies of the African species of the tribe Ommatiini (Scarbrough and Marascia 1996, 2000), this paper clarifies and stabilizes the identities of Ommatius tenellus van der Wuilp (1899) and O. tibialis Ricardo (1903). Both species were originally reported from present day Yemen, O. tenellus from two sites on the mainland and O. tibialis from the island of Socotra. Yet specimens labeled incorrectly as O. tenellus are frequently found in mu- seums. Furthermore, the species was re- ported from southern India (Joseph and Pa- rui 1998) and was listed in the recent Ori- ental and Afrotropical catalogues (Oldroyd 1975). Until recently these reports have not been verified (Scarbrough and Hill 2000). Material examined herein extends the range of O. tenellus westward from Yemen and Israel across the Sahara desert to Senegal. However, O. tenellus is not yet found east of Yemen. A new name for Ommatius tib- ialis, a junior homoym of O. tibialis Say (1823), is proposed. Diptera, Asilidae, Afrotropical, Ommatius, redescriptions METHODS Terminology, measurements, and ratios follow that of previous work (Scarbrough and Marascia 1996, 2000; Scarbrough and Hill 2000). Museum acronyms follow Ar- nett et al. (1993). Data labels associated with each syntype are numbered (1), (2), (3), etc., indicating the sequence of labels on the pin from top to bottom. Brackets provide further information not present on type labels. Ommatius tenellus van der Wulp (Figs. 1-10) Ommatius tenellus Wulp 1899: 97. Type lo- cality: Arabia [= Yemen], Leiji & Haith- alhim; 6 syntypes OXUM, BMNH; Old- royd 1975: 135 (catalogue); 1980: 348 (catalogue). Redescription.—Male: Black body, gray to yellowish-gray tomentose, pale yellow to whitish vestiture and yellow legs. Length, body 7.0-8.0 mm; wing 5.3—5.7 mm. Head: Face scattered short setae dorsally, VOLUME 104, NUMBER 3 681 Wi €°0 6 0.5mm Figs. 1-6. Ommatius tenellus, male terminalia. 1—3, Lateral, dorsal, and ventral views. 4, Gonostylus. 5—6, Aedeagus, dorsal and lateral views. Abbreviations: Ce = cercus, Ep = epandrium, Ge = gonocoxite, Hy = hypandrium. 682 abundant, longer setae ventrally; 2—4 thick, ventromedial bristles and 2 rows of long, thin bristles present; FHWR_ 1.0:6.5—1.0: 6.6. Flagellum slightly longer than scape; FWLR 1.0:1.1. Occiput with 10—12 short, thick, postocular bristles extending to mid- lateral margin of eye, none proclinate. Thorax: Pronotum gray tomentose, white setose; bristles white. Mesonotum mostly dense tomentose, glabrous median stripe, often divided anteriorly with narrow line of tomentum, and 2 lateral spots pre- sent; setae sparse dorsally, 2 dorsocentral bristles present, about as thick and as long as scutellar bristles; anterior notopleural bristle black. Scutellum without a subapical groove, 2 marginal bristles and sparse thin, setae present. Anepimeral bristle present. Halter yellow. Wing: Hyaline, surface slightly corru- gated. Dense microtrichia present narrowly along apical margin. Cell rl long, ending just before costal vein, petiole extremely short. Cell r4 with base just beyond apex of cell d, sides beyond base narrow, parallel. Crossvein r-m just before to just beyond middle of cell d, slightly shorter than basal petiole of cell m3. Base of cell m, wide, WR 1.0:2.0:1.9. Cell m3 with apex near base of cell ml. Leg: Fore and middle coxae with stout bristles. Fore and middle trochanters yel- lowish brown, hind trochanter brown. Fem- ora largely yellow, brown variable anteri- orly, ranging from light brownish yellow basally, light to brown streaks, to exten- sively brown, narrow apex of middle and hind femora brown in type series only; bris- tles yellow. Fore femur with | stout, ven- trobasal bristle; middle femur with | anter- oventral and 3—4 posteroventral bristles. Hind femur with 5, wide spaced, anterov- entral bristles; 5 posteroventral bristles pre- sent on basal third only; HFWLR 1.0:5.5. Tibiae yellow, narrow apex of middle tibia usually and hind tibia always brown. Basal tarsomere of all tarsi largely yellow, narrow apex and remaining tarsomeres of each tar- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sus brown; all tarsi with few to several yel- low bristles. Abdomen: Slightly clavate apically. Ground color mostly black to dark brown, apical margin of most segments lighter, api- cal 1 to 2 segments lighter, grading to red or yellow; tomentum mostly gray, light brownish-yellow to reddish-brown spot with brown setae on most tergites dorsally; setae mostly yellow. Terminalia: Brownish yellow to yellow. Epandrium wide basally, abruptly narrowed and digitate apically. Aedeagus with sheath concave in lateral view, apically with abun- dant setae, thin in lateral view, thicker in dorsal view. Gonocoxal process, unusually long and wide, triangular. Hypandrium about as long as wide, slightly emarginated apically. Female: Differs from male as follows. Body 8.3 mm; wing 7.7 mm; FHWR 1.0: 5.7; -FWIER. 1.0:1.0;-mI/WER PEO:2 3-2: HFWLR 1.0:49. Thorax: Glabrous stripes and/or spots, narrow/small, sometimes ab- sent. Leg: Hind femur with only long, thin posteroventral setae on basal third. Abdo- men: Apical 2—3 segments reddish yellow to yellow. Tergite 9 long laterally, broadly emarginate posteriorly. Spermatheca wide basally, strongly narrowed and recurved apically. Genital fork slender. Specimens examined.—ALGERIA: 1 ¢, Hoggar Tit Oued Anded, 21.VIII.1987, Tanurix A. Pauly Réc., (MRAC). CHAD: 3 2. Korrom, 800 m, 30 Aout 1959, TIBES- TI, Piedmont, SW Romeau de Mire, (MNHN); | 6, Tchad: N’Gouri, distr. De Kanem VIII-1958, P. Renaud, (MRAC). IS- RABE 12> We Watir 5-1 1975.- AS ereid- berg, (LAUD? 1 2, Ehot GIxeIoyA Freidberg, (TAUI). SUDAN: 1 &, Khar- toun, (NMSA); 1 d, Soudan Kartoun, 21— 26. VII.1927, W. Wismeijer, (RMNH). SEN- EGAL: 2 6,2 2, 25—25 km Sud de Richard Toll piege malaise, 14.8.1989, Leg H. v. d. Volk c.s. (NMSA). YEMEN: Syntypes, 1 6; A0Ote sex ?; (OXUM); syntypewiliid® (BMNH). Distribution.—North Africa, Israel, and VOLUME 104, NUMBER 3 0.5mm 7 9 Figs. 7-10. fork. Abbreviations: Ce = cercus, T8 = tergite 8, T9 = Yemen. Active in March and July through September. Remarks.—This desert species is recog- nized by its small size (7-8 mm); black body with gray tomentum and white to yel- lowish vestiture; clavate abdomen; presence 683 ww ZO wi 70 10 Ommatius tenellus, female terminalia. 7-8, Dorsal and ventral views. 9, Spermatheca. 10, Genital tergite 9, S8 = sternite 8. of ventral bristles on the femora and coxae; black glabrous stripe and two lateral spots per side on the mesonotum; and characters of the terminalia (Figs. 1-10). Superficially, it is similar to O. aegyptius (Scarbrough and Marascia 2000) but differs in the much 684 smaller size, presence of marginal scutellar bristles, a row of anteroventral bristles be- low the hind femur, and the combined char- acters of the wing and terminalia. Van der Wulp (1899) lists only a male and female following the species name but additional comments following the descrip- tion suggest other specimens were in his type series; that is ‘Fairly common. Several specimens from Haithalhim and Lahe}.’ A search of collections in the Netherlands and England produced 6 specimens with Wulp’s labels [see below]. As a holotype was not designated, all of the specimens must be considered syntypes. To fix the current in- terpretation of this name and to ensure sta- bility, I here designate the male in OXUM lectotype and the five remaining specimens in OXUM and BMNH as paralectotypes. Label data and condition of the lectotype and paralectotypes follow. Lectotype d, by present designation, (1) [red lined label with the word] ‘Type’ (2) [a note with] ‘v. d. Wulp, Trans. ENT. Soc. 1899, pages 97— 8, pl. i, figs. 14-15,’ (3) [data label with] S. W. Arabia/19 m. Fr. Aden/Haithalhim/ capt. Mar. 25.95/pres. 1899 by J. W. Yer- bury; (4) [data label with] 1899 [above] iol.) ilepe Ent. Type *#% 280, 4/6, (OXUM) [excellent condition]. Paralecto- types, by present designation, | d, (1) [data label] ‘Type’, (2) Ommatius tenellus vy. d. Wulp, (30 Lahej, Arabia/9.5.95, [good con- dition], (BMNH); 4 @, | (sex), same labels as lectotype except: (4) 1899 [above] 7753, (5) Hope Ent. Type # 280 1/6 [excellent condition]; 1899 [above] 7750, Hope Ent. Type # 280 2/6 [poor condition with ab- domen glued to round label, entire fore tar- sus and apical tarsomeres of remaining tarsi absent; (4) 1899 [above] 7755, (5) Hope Ent. Type # 280 3/6 [excellent condition]; (4) 1899 [above] 7751, (5) Hope Ent. Type # 280 5/6 [excellent condition]; (4) 1899 [above] 7754, (5) Hope Ent.Type # 280 6/ 6) [poor condition; abdomen, wings and left hind leg absent], (OXUM). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ommatius abdelkuriensis Scarbrough, new name Ommatius tibialis Ricardo 1903: 375. Type locality: Sokotra, Ab-dul-Kuri; plate xxii, figs. 10, 10a (primary homonym of Om- matius tibialis Say 1823: 49); 2 holo- type, BMNH; Oldroyd, 1980: 348 (cata- logue). Redescription.—Female: Blackish. Body 9.7 mm; wing, 7.3 mm. Head: Gray tomen- tose with mostly yellow setae. Face evenly contoured, slightly protruding beginning just below antenna; setae, sparse dorsally, more abundant, longer ventrally; 2 vertical rows of 4 black bristles present; FHWR 1.0: 5.0. Frons with 5—6 short, thin, white setae laterally. Antenna yellowish-brown tomen- tose, blackish setose, pedicel ventrally with 1 long, thick seta, at least twice as long as remaining setae; FWLR 1.0:1.3. Ocellar tu- bercle with 2 long, black setae posteriorly, 2—3 short, yellow setae anteriorly. Occiput whitish setose, postocular bristles mostly yellow, 4—5 dorsal bristles thin and brown, longest strongly proclinate. Thorax: Mesonotum dorsally subshiny with sparse grayish tomentum, tomentum dense laterally and posteriorly; setae sparse, whitish; 4 lateral bristles black; dorsocen- tral setae thin, mostly short, 4 longer setae posteriorly, about two-thirds as long as pos- talar bristle, | black seta present. Scutellum grayish tomentose, with numerous, long se- tae, setae about half as long as 2 marginal bristles; marginal bristles yellow, about as long as and thick as posterior dorsocentral setae; preapical groove absent. Pleuron grayish tomentose, white setose; anepimeral bristle absent. Halter base yellow, knob red- dish. Wing: Hyaline, dense microtrichia pre- sent apically. Crossvein r-m at apical third of cell d. Base of cell r4 beyond apex of cell d; base of vein M, strongly angled for- ward, cell ml wide apically; ml1WR 1.0: 3.4:2.8. Leg: Fore and middle coxae grayish to- mentose, whitish setose; hind coxa mostly VOLUME 104, NUMBER 3 brownish tomentose. Femora shiny black, mostly or all whitish setose, ventral setae long. Fore femur with a row of long, thin setae, basal 3—4 setae about 1.5 times as long as greatest width of fore femur. Middle femur with anterior bristles black; anterov- entral bristles absent, setae short, poster- oventral setae twice as long as anteroventral setae. Hind femur anteriorly with | black and | yellowish bristle; all ventral vestiture yellowish, anteroventral setae mostly short and thin, 2—3 medially contrastingly thicker than most; posteroventral setae much longer and thinner than anteroventral setae, medi- ally 3—4 thickest and longest; HFWLR 1.0: 6.2. Tibiae yellow with narrow apex brown; vestiture mostly blackish, each tibia with 2— 3 thin yellow bristles; apical third of hind tibia anteriorly brown to brownish yellow. Tarsi brown except basal tarsomere of fore and middle tarsi lighter, reddish brown; bristles black. Abdomen: Black, mostly gray tomen- tose, pale yellow setose; tergites 1—6 dor- sally reddish-brown tomentose, apical 3—4 tergites with black setae; apex of most ter- gites with black, thin setae medially, thick- er, longer, pale yellowish bristles laterally: tergites 7 and 8 sparse gray tomentose, api- cal bristles thicker than preceding tergites; sternite 6 with a black bristle in apical cor- ner, sternite 7 with 6—7 bristles. Sternite 8 flat, apical margin broadly produced medi- ally; 4 prominent black bristles and a long, low, median carina present, carina extends posteriorly from apical margin to posterior most bristle. Cercus narrow, twice as long as wide. Type material—Holotype 2, Sokotra/ 1916-1975; Holotype [round label]; Om- matius tibialis Ricardo/det J. E. Chainey, 1986, [BMNH]. Distribution.—Known only from the Is- land of Socotra, Yemen. Active in Febru- ary. Remarks.—This species is recognized by the blackish body; shiny black femora, and gray tomentum and mostly light vestiture of 685 the body; thinly tomentose mesonotum; abundant long setae on the scutellum and below the femora; and strongly produced apical margin and four prominent bristles on sternite 8. It is further characterized by the presence of thin dorsocentral setae and scutellar bristles, both yellow, and in the absence of an anepimeral bristle. Unknown to Ricardo, the binominal, Ommatius tibialis, had been used earlier for a North American species (Say 1823, Bul- lington and Lavigne 1984). Here I propose a replacement name, Ommatius abdelku- riensis Scarbrough, in reference to the type locality. ACKNOWLEDGMENTS Many thanks to the following curators for providing specimens for this paper: John Chainey, The Natural History Muse- um, London, BMNH; Jason Londt, Natal Museum of South Africa, Pietermaritzburg, NMSA; E. De Coninck and Jos Debacker, Musee Royal de L Africa Centrale, Tervur- en, RMAC; Erik J. van Nieukerken, Na- tional Museum of Natural History/Natural- is, Leiden, RMNH; Léonidas Tsacas, Mu- séum National D’ Historie Naturelle, Paris, MNHN; Darren Mann and I. Lansbury, Ox- ford University Museum of Natural Histo- ry, Oxford, OXUM; Amnon Freidberg, Tel Aviv Museum of Zoology, Tel Aviv, TAUI. Special thanks to John Chainey, Darren Mann, and I. Lansbury for kindly providing the types of Ommatius tibialis Ricardo and O. tenellus v. d. Wulp, and to Darren Mann, Erik J. van Nieukerken, and Ben Brugge, Zoblogisch Museum, Amsterdam, for their assistance and hospitality during my visit to study type material at their museums. Drs. William L. Grogan, Salisbury University, Salisbury, MD, and Eric M. Fisher, Cali- fornia Department of Food and Agriculture, Sacramento, CA, provided helpful com- ments on this manuscript. LITERATURE CITED Arnett, R. H., G. A. Samuelson, and G. M. Nishida. 1993. The insect and spider collections of the 686 world, 2nd Ed. Sandhill Crane Press, Gainesville, FL, 310 pp. Bullington, S. and R. Lavigne. 1984. Review of the genus Ommatius Wiedemann (Diptera: Asilidae) in Eastern United States with descriptions of five new species. Annals of the Entomological Society of America 77: 372-392. Joseph, A. N. T. and P. Parui. 1998. Fauna of India and the adjacent countries. Diptera (Asilidae) (Part 1) General introduction and tribes Leptogas- terini, Laphriini, Atomosini, Stichopogonini and Ommatini. Zoological Survey of India, Calcutta, 278 pp. Oldroyd, H. 1975. Family Asilidae, pp. 99-156. In Delfindo, M. D. and E. Hardy, eds. A catalog of Diptera of the Oriental Region Volume 2. Subor- der Brachycera through Division Aschiza, Subor- der Cyclorrhapha. University Press of Hawai, Honolulu. . 1980. Family Asilidae, pp. 334—374. In Cros- skey, R. W., ed. Catalogue of the Diptera of the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Afrotropical Region. British Museum (Natural History), London. Ricardo, G. 1903. Asilidae, pp. 362-375, pl. 22, figs. 7—11. In Ricardo and Theobald, in Forbes: Natural History of Socotra and Abd-el-Kuri. II. The flies of Abd-el-Kuri. Liverpool, xiv + 598 pp. Say, T. 1823. Description of the dipterous insects of the United States. Journal of the Academy of Sci- ence Philadelphia 3: 9-54; 73-104. Scarbrough, A. G. and C. G. Marascia. 1996. Status of the genus Emphysomera Schiner, 1866 (Dip- tera: Asilidae), with synopsis of Afrotropical spe- cies. Annals of the Natal Museum 37: 191-213. 2000. Revision of Ommatius Wiedemann (Diptera: Asilidae) subgenus Metommatius Hull, 1962. Annals of the Natal Museum 41: 157-179. Scarbrough, A. G. and Hilary N. Hill. 2000. Om- matiinae Robber flies (Diptera: Asilidae) from Sri Lanka. Oriental Insects 34: 341—407. Wulp, FE D. v. d. 1899. Asilidae from Aden and its neighborhood. Transactions of the Entomological Society of London. Part 1, pp. 81—98, pls. 2-3. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 687-69 | LEPTOYPHA ELLIPTICA MCATEE AND L. ILICIS DRAKE (HEMIPTERA: TINGIDAE): NEW DISTRIBUTION RECORDS OF SELDOM-COLLECTED LACE BUGS, WITH CLARIFICATION OF HOST-PLANT RELATIONSHIPS A. G. WHEELER, JR. Department of Entomology, Clemson University, Clemson, SC 29634, U.S.A. (e-mail: awhlr@clemson.edu) Abstract.—Previously published information on the host plant of Leptoypha elliptica McAtee and L. ilicis Drake—holly, /lex sp. (Aquifoliaceae)—is based on misidentification of the actual host. Both lace bugs specialize on shrubs of the genus Forestiera (Oleaceae) and, thus, develop on oleaceous plants like nearly all other species of Leptoypha. In addition to a clarification of host associations for these little-known tingids, new distri- bution records and biological notes are provided. Key Words: Holly, /lex sp. (Aquifoliaceae), is the only recorded host plant of the lace bug Leptoypha ilicis Drake; the specific epithet reflects this tingid’s collection on a shrub presumed to be a species of //ex. A second lace bug, L. elliptica McAtee, also was tak- en on “‘Jlex’’ sp. at the type locality of L. ilicis in Georgia (Drake 1919, McAtee 1919). My 1985 collection of both species on “holly” in Tennessee supported an ab- errant host association in a genus that oth- erwise develops on members of the Ole- aceae. My eventual discovery that the host plant in Tennessee was not a holly but an olea- ceous shrub, glade privet (Forestiera ligus- trina [Michaux] Poiret), suggested that the ‘“Tlex’’ in Georgia had been similarly mis- identified in 1917 when L. ilicis was first collected. A 1991 trip to Stone Mountain, Ga., the type locality of L. ilicis, plus ad- ditional fieldwork, confirmed my hypothe- sis that L. ilicis, and the often syntopic L. elliptica, develop on species of Forestiera. Here, in addition to clarifying host rela- insect distribution, Forestiera spp., host-plant relationships tionships for both tingids, I provide new distribution records and biological notes. Voucher material has been deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. Leptoypha elliptica McAtee (Fig._1) Described from ‘‘Texas” without host- plant data by McAtee (1917), this tingid has since been reported from Florida and Geor- gia (Drake 1918), Indiana (Blatchley 1926), Missouri (Froeschner 1944), and Tennessee (Drake and Ruhoff 1965). It was included in a list of the Tingidae of Oklahoma (Drew and Arnold 1977) because the recorded dis- tribution suggested its eventual collection in that state. Drake’s (1919) record of holly as the host on Stone Mountain, Ga., was based on the miridologist H.H. Knight’s apparent misidentification of F. ligustrina. The Texas record from “‘swamp bush” (Drake 1918) almost certainly refers to swamp privet (F. acuminata; see Discussion). New collection records (* = new state 688 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-2. 1, Leptophya elliptica. 2, L. ilicis. record)—-GEORGIA: DeKalb Co., Stone Mountain, southwestern slope ca. 400 m, 3 May 1991. *ILLINOIS: Johnson Co., Belk- nap-Karnak Rd., SW. of Belknap, 8 June 1991. INDIANA: Knox Co., co. rd. 1300- S nr. Swan Pond, 13 km NW. of Decker, 7 June 1991. *KENTUCKY: Ballard Co., Rt. 51 NE. of Wickcliffe, 9 June 1991. MIS- SOURI: Mississippi Co., Birds Point, 9 June 1991 (nymphal exuviae only); New Madrid Co., New Madrid, 9 June 1991. *SOUTH CAROLINA: Aiken Co., Savan- nah River Bluffs Heritage Preserve, ca. 5 km NW. of North Augusta, 18 April 1992 (nymphs only, reared to adulthood), 12 May 2001. TENNESSEE: Davidson Co., Couchville Glade, NNE. of LaVergne, 10 June 1997; Long Hunter State Recreation Area, 29 May 1985, 19 April 1991; Mt. View Rd. Cedar Glade, NNW. of LaVergne, 12 June 1997, 13 May 2000; Dyer Co., Rt. 78, 1 km N. of Obion River, 1.5 km S. of Bogota, 2 June 1985, 9 June 1991; Ruth- erford Co., Flat Rock Cedar Glades & Bar- rens State Natural Area, 8.5 km NNE. of Murfreesboro, 11 June 1997, 14 May 2000; Sunnybell Glade, SW. of Mona, 12 June 1997; Wilson Co., Lane Farm Glade, NNW. of Silver Hill, 10 June 1997. Host plants.—Forestiera ligustrina was the host for collections in Georgia and South Carolina, plus Davidson, Rutherford, and Wilson counties in Tennessee. In other cases the host was F. acuminata (Michaux) Poiret. Leptoypha ilicis Drake (Fig. 2) Drake (1919) described L. ilicis from Stone Mountain, Ga., mentioning its simi- larity to L. mutica (Say). The type series was collected in June 1917 on “‘holly”’ but, as noted above, this supposed host associ- ation is erroneous. Additional records of this species have been few: Florida (Blatch- ley 1926, 1928), Oklahoma, and Texas (Hurd 1946). Hurd’s (1946) New Hamp- shire record, which Bailey (1951) said re- quired verification, has been excluded from VOLUME 104, NUMBER 3 subsequent catalogs of the family (Drake and Ruhoff 1965, Froeschner 1988). New collection records (* = new state record; unless cited completely, dates and specific localities are the same as those pro- vided for L. elliptica) GEORGIA: Co- lumbia Co., Heggies Rock, E. of Appling, 5 April 1997; DeKalb Co., Pleasant Hill outcrop, N. of Lithonia, 7 July 1996, and Stone Mountain. *ILLINOIS: Johnson Co. *INDIANA: Knox Co. *KENTUCKY: Bal- lard Co. *MISSOURI: Mississippi and New Madrid counties. “SOUTH CAROLINA: Aiken Co. (12 May 2001 only). *TENNES- SEE: Davidson, Dyer, Rutherford, and Wil- son counties. Host plants.—Because this species gen- erally co-occurred with L. elliptica, its hosts are the same as listed under that species: F. ligustrina (Georgia and South Carolina, plus Davidson, Rutherford, and Wilson counties in Tennessee) and F. acuminata (all other localities). DISCUSSION Froeschner (1944) reported adults and nymphs of L. elliptica from Missouri but did not mention a host plant. The actual host relationships have gone unrecorded for both L. elliptica and L. ilicis since the orig- inal descriptions of these lace bugs ap- peared more than 80 years ago. My recent fieldwork now demonstrates that the specif- ic epithet L. ilicis is a misnomer based on misidentification of the oleaceous shrub F. ligustrina at the type locality. Even this lace bug’s other known host, F. acuminata, can be confused with holly (Stephens 1973). Both L. elliptica and L. ilicis specialize on species of Forestiera, and they often are syntopic. The two species are easily distin- guished; the adults of L. elliptica are broad- ly elliptical, whereas those of L. ilicis are narrowly oblong and subparallel (see Figs. 1, 2). The collection of L. ilicis from Vac- cinium sp. (Blatchley 1928) and in “palm jungle sweepings” (Hurd 1946) should be considered accidental occurrences. All records of these lace bugs fall within 689 the known distributions of their principal hosts, fF. acuminata and F. ligustrina. A\- though these plants do not range throughout the eastern United States (they do not occur north or east of central Kentucky and Ten- nessee and southern South Carolina), they are the most widely distributed members of this small New World genus [ca. 15 spp. (Everett 1981)]. That most entomologists are unfamiliar with these inconspicuous plants likely has contributed to the scarcity of both lace bugs in collections. Neither tin- gid was taken during surveys of the lace bugs of Georgia (Beshear et al. 1976, Be- shear 1981) and Oklahoma (Drew and Ar- nold 1977), despite historical records of both species from Georgia and of L. ilicis from Oklahoma (Drake and Ruhoff 1965). Forestiera acuminata occurs sporadically along stream and river banks and in bottom- land swamps and sloughs from southern In- diana and central Illinois, west to south- eastern Kansas, eastern Oklahoma, and Texas, and eastward to southern South Car- olina, Georgia, and Florida (Little 1977, 1980; Godfrey 1988; Gleason and Cron- quist 1991). This weak, leaning, straggly, and shade-tolerant shrub or small tree often grows under larger trees (Deam 1932, Brown and Kirkman 1990). Most abundant in Arkansas, Missouri, and Texas, swamp privet generally attains its largest size in Louisiana (Sargent 1922). This likely was the plant on which Froeschner (1944) found nymphs of L. elliptica in Missouri. Forestiera ligustrina, a deciduous shrub of limestone outcrops and sandy or rocky soil, has a more restricted range than F. ac- uminata. Glade privet is found irregularly from central Kentucky and Tennessee to eastern Texas, southern South Carolina, southwestern Georgia, southern Alabama, and northern Florida (Godfrey 1988, Glea- son and Cronquist 1991). It is a character- istic plant of shrub thickets adjacent to most eastern limestone (cedar) glades (e.g., Mey- er 1937, Quarterman et al. 1993, Baskin et al. 1995, Baskin and Baskin 1999). Both lace bugs, like other members of 690 the genus, are mesophyll feeders; they col- onize lower leaf surfaces of Forestiera spe- cies and cause chlorosis on upper surfaces. Foliar chlorosis usually is light and patchily distributed on host plants. Assuming that adults overwinter, which is typical of other Leptoypha species (Mead 1975, Sheeley and Yonke 1977), they become active in early spring. In the present study, late in- stars of L. elliptica were observed in mid- April in Tennessee and South Carolina. The collection of fifth instars of L. elliptica in Tennessee in late May and of L. ilicis in Georgia in early July likely represented a second generation. Other Leptoypha_ spe- cies, such as L. mutica and L. costata Par- shley, are bivoltine or trivoltine (Dickerson and Weiss 1916, Sheeley and Yonke 1977). In addition to the need to clarify details of their life histories, other biological as- pects of these lace bugs warrant attention. Do L. elliptica and L. ilicis use other Fo- restiera species as hosts? Does the ash- feeding L. mutica include F. acuminata in its host range, as suggested by its collection on this plant in Texas (Drake 1918), or was the presence of adults on swamp privet strictly accidental? Two apparent problems that were suggested by Mead (1975)—the taxonomic status of L. i/icis and puzzling degree of intraspecific variation in L. mu- tica—will be addressed in a forthcoming review of Leptoypha by Thomas J. Henry. ACKNOWLEDGMENTS I am grateful to T. J. Henry (Systematic Entomology Laboratory, USDA, % Nation- al Museum of Natural History, Washington, DC) for identifying the tingids and for help- ing collect the tingids in Tennessee in 1985; G. R. Carner (Dept. of Entomology, Clem- son University) for photographing the tin- gids; J. R. Allison and T. S. Patrick (Geor- gia Dept. of Natural Resources, Social Circle) for accompanying me in the field at Stone Mountain; R. L. Brown (Dept. of En- tomology and Plant Pathology, Mississippi State University) for inviting me to partic- ipate in the Mississippi Entomological Mu- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON seum’s William H. Cross Expedition to the Tennessee cedar glades in 1997 and for se- curing collecting permits; the Palmetto In- sect Naturalists for allowing me to accom- pany them on their 1997 field trip to Heg- gies Rock in Georgia and obtaining per- mission to collect insects; R. S. Jensen (Tennessee Dept. of Conservation, Nash- ville) for granting permission to collect at Long Hunter State Recreation Area; G. K. Call and S. Rollins (The Nature Conservan- cy of Tennessee, Nashville) for permission to collect insects at the Mt. View Road Ce- dar Glade and the Flat Rock Cedar Glades; S. H. Bennett (South Carolina Dept. of Nat- ural Resources, Columbia) for permission to collect at Savannah River Bluffs Heri- tage Preserve; M. A. Homoya (Indiana Dept. of Natural Resources, Indianapolis), L. Johnson (Indiana University, Blooming- ton), A. Koelling (Illinois State Museum, Springfield), J. B. Nelson (University of South Carolina, Columbia), and P. S. Som- ers (formerly Tennessee Dept. of Conser- vation, Nashville) for providing herbarium records of Forestiera spp. and/or directions to plant colonies; E. R. Hoebeke (Dept. of Entomology, Cornell University, Ithaca, NY) for providing copies of several papers; and P. H. Adler (Dept. of Entomology, Clemson University) for reviewing the manuscript. LITERATURE CITED Bailey, N. S. 1951. The Tingoidea of New England and their biology. Entomologica Americana 31: 1-140. Baskin, J. M. and C. C. Baskin. 1999. Cedar glades of the southeastern United States, pp. 206-219. In Anderson, R. C., J. S. Fralish, and J. M. Baskin, eds. Savannas, Barrens, and Rock Outcrop Com- munities of North America. Cambridge University Press, Cambridge. Baskin, J. M., D. H. Webb, and C. C. Baskin. 1995. A floristic plant ecology study of the limestone glades of northern Alabama. Bulletin of the Torrey Beshear, R. J. 1981. New records of Georgia lace bugs (Homoptera [sic]: Tingidae). Journal of the Geor- gia Entomological Society 16: 266. Beshear, R. J., H. H. Tippins, and J. O. Howell. 1976. The lace bugs (Tingidae) of Georgia. University VOLUME 104, NUMBER 3 of Georgia Agricultural Experiment Station Re- search Bulletin 188: 1—29. Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America, with Especial Reference to the Faunas of Indiana and Florida. Nature Pub- lishing, Indianapolis, Ind. 1,116 pp. 1928. Notes on the Heteroptera of eastern North America with descriptions of new species, I. Journal of the New York Entomological Society 36: 1—23. Brown, C. L. and L. K. Kirkman. 1990. Trees of Geor- gia and Adjacent States. Timber Press, Portland, Ore. 292 pp. Deam, C. C. 1932. Trees of Indiana (2nd rev. ed.). Fort Wayne Printing, Fort Wayne, Ind. 326 pp. Dickerson, E. L. and H. B. Weiss. 1916. Notes on Lep- toypha mutica Say (Hemip.). Entomological News 27: 308-310. Drake, C. J. 1918. Notes on North American Tingidae (Hem.-Het.). Bulletin of the Brooklyn Entomolog- ical Society 13: 86-88. 1919. On some North American Tingidae (Hemip.). Ohio Journal of Science 19: 417—421. Drake, C. J. and E A. Ruhoff. 1965. Lacebugs of the World: A Catalog (Hemiptera: Tingidae). Smith- sonian Institution, Washington, D.C. 634 pp. Drew, W. A. and D. C. Arnold. 1977. Tingoidea of Oklahoma (Hemiptera). Proceedings of the Oklahoma Academy of Science 57: 29-31. Everett, T. H. 1981. The New York Botanical Garden Hlustrated Encyclopedia of Horticulture. Vol. 4, Di-Fu. Garland Publishing, New York. pp. 1059— 1422. Froeschner, R. C. 1944. Contributions to a synopsis of the Hemiptera of Missouri, Pt. II. Lygaeidae, Pyr- rhocoridae, Piesmidae, Tingidae, Enicocephalidae, Phymatidae, Ploiariidae, Reduviidae, Nabidae. American Midland Naturalist 31: 638-683. . 1988. Family Tingidae Laporte, 1807 (=Tin- gididae; Tingitidae). The lace bugs, pp. 708—733. In Henry, T. J. and R. C. Froeschner, eds. Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden. Gleason, H. A. and A. Cronquist. 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd ed. New York Botanical Garden, Bronx, N.Y. 910 pp. 691 Godfrey, R. K. 1988. Trees, Shrubs, and Woody Vines of Northern Florida and Adjacent Georgia and A|- abama. University of Georgia Press, Athens. 734 pp. Hurd, M. P. 1946. American Tingoidea (Hemiptera-Heteroptera). Iowa State College Journal of Science 20: 429— 492. Little, E. L., Jr. 1977. Atlas of United States trees, Vol. 4. Minor eastern hardwoods. U.S. Department of Agriculture Forest Service Miscellaneous Publi- cation 1342. 17 pp. + 230 maps. . 1980. The Audubon Society Field Guide to North American Trees: Eastern Region. Alfred A. Knopf, New York. 714 pp. McAtee, W. L. 1917. Key to the Nearctic species of Leptoypha and Leptostyla (Heteroptera Tingidae). Bulletin of the Brooklyn Entomological Society 12: 55-64. . 1919. Corrections and additions to an article on Leptoypha and Leptostyla (Heteroptera; Tin- gidae). Bulletin of the Brooklyn Entomological Society 14: 142-144. Mead, F W. 1975. The fringetree lace bug, Leptoypha mutica (Say) (Hemiptera: Tingidae). Florida De- partment of Agriculture and Consumer Services, Division of Plant Industry, Entomology Circular 161. 2 pp. Meyer, A. M. 1937. An ecological study of cedar glade invertebrates near Nashville, Tennessee. Ecologi- cal Monographs 7: 403—443. Quarterman, E., M. P. Burbanck, and D. J. Shure. 1993. Rock outcrop communities: limestone, sandstone, and granite, pp. 35-86. Jn Martin, W. H., S. G. Boyce, and A. C. Echternacht, eds. Bio- diversity of the Southeastern United States: Up- land Terrestrial Communities. John Wiley, New York. Sargent, C. S. 1922. Manual of the Trees of North America (Exclusive of Mexico). Houghton Mif- flin, Boston. 910 pp. Sheeley, R. D. and T. R. Yonke. 1977. Immature stages and biology of Atheas austroriparius and Leptoy- pha costata (Hemiptera: Tingidae). Annals of the Entomological Society of America 70: 603-614. Stephens, H. A. 1973. Woody Plants of the North Cen- tral Plains. University Press of Kansas, Lawrence. 530 pp. Generic classification of North PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 692-701 TAXONOMIC NOTES ON STRONGYLOGASTERINI (HYMENOPTERA: TENTHREDINIDAE) STEPHAN M. BLANK Deutsches Entomologisches Institut im ZALE SchicklerstraBe 5, 16225 Eberswalde, Germany (e-mail: blank @zalf.de) Abstract.—The western Palaearctic members of the Strongylogaster multifasciata group, S. multifasciata (Geoffroy 1785) and S. xanthocera (Stephens 1835), are keyed, and the status of the included nominal taxa is discussed in view of taxonomical and zoogeographical aspects. Lectotypes are designated for Strongylogaster desbrochersi Ko- now 1892 and S. desbrochersi var. lepticus Konow 1902, which are new synonyms of S. xanthocera (Stephens 1835). Strongylogaster lineata cypria Benson 1954 is a new syn- onym of S. multifasciata. The striking morphological variability among West Palaearctic S. xanthocera is caused by the polycentric, both Mediterranean and Eurosiberian, origin of the population. Hemitaxonus Ashmead 1898 is proposed as a new synonym of Thrinax Konow 1885, and 22 of 23 species involved are new combinations with 7hrinax. Key Words: Strongylogaster, a genus of Selandriinae (Tenthredinidae), is represented in the Hol- arctic region by some 40 species. The lar- vae are external feeders on ferns. The mem- bers of the S. multifasciata-group share the distally widened and elongate valvulae 3 of the ovipositor sheath as an apomorphy. Nai- to (1996) considered six species of this group, two of which, S. multifasciata (Geoffroy) and S. xanthocera (Stephens), are distributed also in the western Palaearc- tic. In Europe and northern Africa, adults and larvae of these species sometimes occur numerously on the fronds of the fern named bracken (Preridium aquilinum (L.) Kuhn). This fern, a cosmopolitan pest of pastures, is poisonous to grazers and often harmful to the autochthonous plant cover. It is one, in the Mediterranean possibly the main, lar- val host of S. multifasciata and S. xantho- cera (see Taeger et al. 1998 for list of hosts). Throughout the Mediterranean area S. Strongylogaster, Hemitaxonus, Thrinax, Selandriinae multifasciata and S. xanthocera seem to produce local forms, and several of them have been named in the past. The present taxonomic results partly contradict Naito’s (1996) concept. A key for the S. multifas- ciata-group in the West Palaearctic realm is presented because the published keys (Nai- to 1996, Blank 1998) are not appropriate for the identification of males and of North African specimens. Zoogeographical as- pects are discussed to construe morpholog- ical variability. The second part of this work treats the proper use of the name Thrinax, which was so far applied either for a subdivision of Strongylogaster or for a separate genus close to Strongylogaster. MATERIAL AND METHODS The following abbreviations are used for museum collections: DEI = Deutsches En- tomologisches Institut, Eberswalde, Ger- many; MCSN = Museo Civico de Storia VOLUME 104, NUMBER 3 1 2 Female hypopygium. |, Strongylogaster YE §YLO;: Figs. 1-2. multifasciata. 2, S. xanthocera. Naturale ““Giacomo Doria’, Genoa, Italy; MZLU = Museum of Zoology, Lund Uni- versity, Sweden; NHM = The Natural His- tory Museum, London, Great Britain; ZSM = Zoologische Staatssammlung, Mtinchen, Germany. Before dissection, the male genitalia were treated with 3% potassium hydroxide solution for 6-8 hours at ca 50°C to dis- solve tissue from chitinous structures, and then rinsed with water. The dissected penis valve was mounted in glycerine for photo- graphing. Digital photographs were taken with a Olympus C-3030 camera and pro- cessed with PhotoImpact 4.2 and Corel- DRAW 9. In the list of Thrinax species, the occurrence of each species in a zoogeo- graphical region is indicated by [WP] for West Palaearctic, [EP] for East Palaearctic, [N] for Nearctic, and [O] for Oriental. KEY TO WESTERN PALAEARCTIC SPECIES OF THE STRONGYLOGASTER MULTIFASCIATA GROUP ile Female 2 - Niall cieereewroney 2502 oe PS re beyh aha op Gils AR 3) 2(1). Appendix of hypopygium hardly protruding beyond distal margin of hypopygium, ba- sally with a deep depression confined by a distinct raised edge (Fig. 1). Pterostigma usually dark brown anteriorly and_ pale brown posteriorly. Antenna black, usually scape, pedicel and ventral side of antennom- ere. Sred= = 2 = : Strongylogaster multifasciata (Geoffroy 1785) = Appendix of hypopygium distinctly protrud- ing beyond distal margin of hypopygium, without basal depression or distinct edge (Fig. 2). In European specimens, usually an- tennomeres |—4(—9) red and pterostigma un- icolorous pale, in North African specimens antenna mostly black and pterostigma dark bowl = st sos == Strongylogaster xanthocera (Stephens 1835) 693 3(1). Valviceps 1.85—2.05 times longer than wide, dorsal and ventral edge almost straight, dis- tally with an almost straight dorso-distal and ventro-distal outline, medial sclerotization clearly extending up to distal edge of val- viceps (Figs. 3—4). Pterostigma usually an- teriorly dark brown and posteriorly pale brown. Fore and hind coxae pale except for a narrow line along base. Abdominal seg- ments 3—9 (always?) without black pattern Strongylogaster multifasciata (Geottroy 1785) - Valviceps 1.55—1.70 times longer than wide, ventral edge sinuate, distal edge evenly rounded, medial sclerotization distally fad- ing and not clearly extending up to distal edge of valviceps (Figs. 5—8). Pterostigma unicolorous brown to piceous, seldom bi- colorous. Fore coxa with black line along base about as wide as ocellar diameter, hind coxa predominantly black at least laterally. Abdominal segments 4—9 red, in central and northern European specimens, anteriorly more or less black on segments 4—9 Strongylogaster xanthocera (Stephens 1835) Strongylogaster Dahlbom 1835 Strongylogaster Dahlbom 1835: 4, 13. Type species: Tenthredo multifasciata Geot- froy 1785, subsequent designation under the plenary power (ICZN 2000: Opinion 1953; see Blank et al. 1999) Pseudotaxonus A. Costa 1894: 157. Type species: Tenthredo (Allantus) filicis Klug 1817, by monotypy. Synonymy by Ross (1937) Polystichophagus Ashmead 1898: 310. Type species: Tenthredo (Allantus) filicis Klug 1817, by monotypy. Synonymy by Enslin (1913) Prototaxonus Rohwer 1910c: 49. Type spe- cies: Prototaxonus typicus Rohwer 1910 [= Strongylogaster tibialis Cresson 1880], by original designation. Synony- my by Ross (1937) Thrinax auct. nec Konow 1885; = Stron- gyvlogaster subgen. Thrinax auct. Strongylogaster multifasciata (Geoffroy 1785) (Figs. 1, 3, 4) Tenthredo spec. Geoffroy 1762: 278, Nr. 14, @. Tenthredo multi-fasciata [recte: multifas- 694 ciata| Geoffroy in Fourcroy 1785: 368, Nr. 14, 2, type locality: France, Paris. Tenthredo lineata Christ 1791: 450, °, type locality: ?; = Strongylogaster lineata (Christ 1791). Synonymy by Blank (1998). Tenthredo alternans Serville 1823: 17, 2, type locality: France, environs of Paris. Synonymy by Dalla Torre (1894). Tenthredo alternans Lepeletier 1823: 73, 2, type locality: France, environs of Par- is. Synonymy by Dalla Torre (1894). Tenthredo (Allantus) linearis Klug 1817: 217-218, d, type locality: Austria, Carin- thia, preoccupied in Tenthredo by Schrank 1781: 343. Synonymy by Hartig (1837) and Dalla Torre (1894). Strongylogaster cretensis Konow, 1887: 26, 2 6, type locality: Greece, Crete. Syn- onymy by Naito (1996). Strongylogaster lineata cypria Benson 1954: 276, 2, type locality: Cyprus, near Platania Forest Station. New synonym. Tenthredo cingulata auct. nec Fabricius 1793: See Blank etal: (1999). Type material.—Strongylogaster creten- sis. Lectotype d (designated by Schedl LOS) AS eEretay =) Coll’ -Konow > [red=] “Typus”; [label with red margin:] ““Lecto- typus Strongylogaster cretensis Konow °”’; “Strongylogaster multifasciata (Geoffroy, 1785) det. S. M. Blank 2001”. Both anten- nal flagella and parts of the left fore wing and of the legs missing. Paralectotype: 1 d from Crete, in poor condition. DEI. Strongylogaster lineata cypria. Holotype 2: [round label with red edge:] ““Type’’; “Cyprus Nr. Platania Forest Stn. 2,500-— 4,000’ 7.v.1945 G. A. Mavomoustakis”’; “near Platania Forest Station 3,500—4,000 ft 7.5.45”; “Holotype Strongylogaster li- neata cypria subsp. n. det. 2 R.B. Benson 19537 >) “B._Mr Type: Hyme 1677 2° ‘Stron- gylogaster multifasciata (Geoffroy, 1785) det. S. M. Blank 2001’. In perfect condi- tion. Paratypes: 5 2, only 1 @ checked from “Lebanon, Falouka, 17.v.1953’’, leg. Mavromoustakis. NHM. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Remarks.—The typical form of S. mul- tifasciata is widely distributed throughout the western Palaearctic Region. According to Naito (1996) the same species also oc- curs in the eastern Palaearctic (Sibiria, Chi- na, Korea, Japan, Taiwan). At least in Cen- tral Europe males are comparatively rare, and possibly part of the population repro- duces parthenogenetically. Strongylogaster cretensis is a valid species according to Sched] (1981), who checked the types and designated a lectotype. Naito (1996) syn- onymized S. cretensis with S. lineata (= S. multifasciata). The hypopygium of the fe- male lectotype agrees with S. multifasciata. In both males and females the abdomen is red except for tergum 1 and more or less tergum 2, and the legs are predominantly red. Such specimens are only known from western Crete (Sched! 1981, 1993) and form an allopatric local population of S. multifasciata (see Naito 1996). The form S. lineata cypria described by Benson (1954) for material from Cyprus and Lebanon has entirely pale hind femora and completely black basal antennal seg- ments. It agrees well with the concept of S. multifasciata in the shape of the hypopy- gium and the bicolorous pterostigma. The name Falonka given by Benson is a type error for the Lebanese collecting locality Falouka (= Falugha, Falougha). Naito (1996) omitted cypria, which is hereby treated as a synonym with multifasciata. The subapical claw is usually larger in S. xanthocera than in S. multifasciata, but this character underlies considerable variability. The following characters, which were ap- plied by former authors (see e.g., Muche 1969, Naito 1996), are according to my ex- perience not diagnostic for the separation of the two species: front lobe of mesonotum with/without coarse punctures, density of macro- and microsculpture on other parts of the thorax; relative length of antennal seg- ments; shape of underside of sawsheath near base. The geographic variability of fur- ther characters will be discussed under S. xanthocera below. VOLUME 104, NUMBER 3 695 IIASA ah Soin Pele etter, ite cre Figs. 3-8. Geographic variability of penis valve. 3, Strongylogaster multifasciata from Germany, Lower Bavaria, Bairmannsried. 4, S. multifasciata from Greece, Crete, Khania. 5, S. xanthocera from Germany, Berlin- Friedrichshagen. 6, S. xanthocera from Japan, Honshu, Sasayama. 7, S. xanthocera from Algeria, Theniet el Had. 8, S. xanthocera from Tunisia, Nafzah. Strongylogaster xanthocera germanland and Skane. Synonymy by (Stephens 1835) Konow (1905). (Figs. 2, 5-8) Strongylogaster desbrochersi Konow 1892: Tenthredo xanthocera Stephens 1835: 337, 214-214, @ 6, type locality: Algeria, 2 6, type locality: England, environs of Theniet el Had. New synonym. London. Strongylogaster desbrochersi var. lepticus Strongylogaster geniculata C. G. Thomson Konow 1902: 390, ¢ 6, type locality: 1871: 243, 2, type locality: Sweden, An- Tunisia, Ain Draham; Strongylogaster 696 desbrochersi var. lepidus: Forsius (1930), type error. New synonym. Type material.—Strongylogaster des- brochersi. Lectotype ° (here designated in order to ensure the name’s proper and con- sistent application): ““Alger.”’; “Coll. Ko- now sulired:| ~ Lype -2 cabinet labels “Strongylogaster Desbrochersi Knw. A\- ger.”’; [red:] ““Lectotypus 2 Strongylogas- ter desbrochersi Konow, 1892 des. S. M. Blank 1998”; “‘Strongylogaster multifas- ciata (Geoffroy, 1785) det. S. M. Blank 2001°’. Head and front legs missing. Para- lectotypes: | d *‘Teniet-el-Haad [leg.] Des- brochers 1889”; 1 ¢ “*Teniet el Haad AI- genav; 1 6. Algena: (DEI Strongylogaster desbrochersi var. lepti- cus. Lectotype ¢ (here designated in order to ensure the name’s proper and consistent application): ‘“‘Tunisie Ain Draham, 4.5.[18]98 [leg.] de Gaulle’; “Coll. Ko- now’; [red:] ““Lectotypus d Strongylogas- ter desbrochersi var. lepticus Konow, 1902 des. S. M. Blank 1998”; “‘Strongylogaster multifasciata (Geoffroy, 1785) det. S. M. Blank 2001”. In perfect condition. Paralec- totypes: 2 d “Tunisie Ain Draham, 9.5.98 de Gaulle?. DEI. Remarks.—Konow (1892) described S. desbrochersi from several males and fe- males collected in Algeria, which he char- acterized by the predominantly red abdo- men and the body length ranging between 7.5 and 10.0 mm. The only available female of the syntype series is hereby designated as the lectotype. Naito (1996) synonymized S. desbrochersi and S. lineata (= S. multi- fasciata). This is incorrect because the hy- popygium of the S. desbrochersi lectotype is similar to S. xanthocera (Fig. 2) and not to S. multifasciata. Konow (1902) described Strongylogaster desbrochersi var. lepticus from Tunisia due to the more or less darkened abdomen and legs. Naito (1996) omitted lepticus, but La- court (1999) listed it in synonymy with S. multifasciata. The female syntype(s), men- tioned in the original description, are lost. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON One of the available males is hereby des- ignated as the lectotype. Konow’s original name is deemed to be of subspecific rank, because he stated ““Tunis’? (= Tunisia) as the particular geographical area where the material was collected. Strongylogaster xanthocera is widely distributed throughout the Palaearctic, and in the East it occurs among others in the Russian Far East and in Japan (Zhelochov- tsev 1988, Naito 1996). The most southern European records are from the southern parts of the Balkans and Sicily (Hellén 1967, Zombori 1994, Blank 1993). From Tunisia and Algeria two taxa, lepticus (= lepidus, type error) and desbrochersi, have been described which shall occur there sympatrically. It is impossible to reconfirm this opinion which is based on Forsius’ (1930) checklist of the North African saw- flies, as this lacks original data for his state- ment, and the whereabouts of concerning material is unknown. The North African populations are geographically isolated from the European and particularly those from Tunisia differ strikingly from the Cen- tral and North European. The following character variability is perceptible among West Palaearctic S. xanthocera: 1. The serrulae on the dorsal edge of the valviceps are little larger in males from the southern Mediterranean (Tunisia and Alge- ria; Figs. 7-8) and Greece than in males from Central Europe (Fig. 5), Corsica and Japan (Honshu, Sasayama, 3.5.1963, leg. T. Naito, Fig. 6; DEI). 2. Tunisian specimens (2 6.5—7.5 mm, d 5.5-7.0 mm) are smaller than Algerian (2 ca 9.0 mm, ¢ 7.5—-8.5 mm), specimens from Bulgaria, Corsica, Greece, Sicily, Central and North Europe, and Japan are the largest (2 9.5-11.5 mm, ¢ 7.5—10 mm). 3. Southern Mediterranean specimens tend to have a dark pterostigma, whereas the pterostigma is pale in northern Medi- terranean, Central and North European and Japanese specimens. 4. The amount of pale color on the fe- VOLUME 104, NUMBER 3 male antenna increases in northern direc- tion: in the females from Tunisia and Al- geria the antenna is black and rarely little pale on scape and pedicel; a female from Sicily with scape and pedicel distally little pale (Messina, leg. E Vitale, MCSN; re- ported by Zombori [1984] as S. desbroch- ersi); a Bulgarian female has the scape dis- tally, the pedicel completely and antennom- ere 3 basally red (southern Pirin mountains, Popina Laka, 1,000 m, 14.6.1989, leg. Zer- che & Behne, DEI); females from Greece, Corsica and Central Europe have anten- nomeres 1—4 red; in many females from northeastern Germany the distal antennom- eres are brown instead of black (Berlin- Friedrichshagen; DEI); three females from southern Finland have completely pale an- tennae (Karislojo; DEI). 5. Females with completely red abdom- inal segments 4—10 were observed in the southern Mediterranean realm, where they may occur syntopically (Tunisia; Blank 2002) or at least sympatrically with the pre- dominantly dark form (red form S. des- brochersi, type locality: Teniet el Haad [= Theniet el Had] ca 135 km SW Algiers; dark form: Algeria, Massif des Mouzaia [= Djebel Mouzaia ca 50 km SW Algiers], 14.5.1936, leg. Peyerimhoff, MCSN, see Zombori 1984). The red form is unknown from the northern Mediterranean realm and from Central and North Europe. 6. The abdominal segments 4—9 are red in males from the Mediterranean realm, but the anterior portions of segments 4—9 are extensively darkened towards the abdomi- nal tip in Central European and Japanese males. The geographic pattern of these charac- ters matches a polycentric species, which had its refugia in the Mediterranean and in eastern Asia during the glacial times. The strains characterized above do not cover the feature of lepticus, desbrochersi and xan- thocera as recognized prior to Naito (1996). The abdominal coloration in males and the size of the dorsal serrulae of the valviceps roughly distinguishes the populations with 697 Mediterranean and Eurosiberian origin (zoogeographic terminology in accordance with de Latin 1967). I shall abstain from ranking any of them at subspecific level, because males from Corse cannot be clas- sified to either of these groups. They have the abdominal segments 4—9 completely red like other Mediterranean males, but their valviceps bears only small serrulae like in Central European males. Among the Med- iterranean material those specimens from North Africa are characterized by their dark pterostigma and smaller body size. The oth- er characters seem either to follow a (cli- matic?) gradient (antennal color in females) or to occur arbitrarily (females with red ab- domen within local populations of the Med- iterranean region). NOMENCLATURAL NOTES ON HEMITAXONUS AND THRINAX The species of this genus chiefly occur in East Asia and in North America (Naito 1971). The only representative of the West- ern Palaearctic, 7. contigua, is rare in Eu- rope and usually misinterpreted in the Eu- ropean faunistic literature. Konow (1885) described 7. “‘contigua m. (= mixta [sensu] Thoms.)”’ and repeated part of the charac- ters, which Thomson (1871) listed for 7. mixta (i.e., length of body and antennae, shape of ovipositor sheath). Accordingly, it is apparent that Konow had no material of his own on hand, but he interpreted Thom- son’s description and named the taxon. Type material is not present in Konow’s collection (coll. DEI). The specimen, which Malaise (1933) cited as ““Thomson’s type’’, is the lectotype of 7. contigua (type local- ity: northern Sweden; cf. Blank 1998). Mal- aise (1933) identified this specimen as ‘*struthiopteridis Fors.’ [= Sahlbergia struthiopteridis Forsius, 1910]. S. struthiop- teridis has widely been accepted as a mem- ber of Hemitaxonus (e.g., Enslin 1914, Zhelochovtsev 1951, Naito 1971). Conse- quently, the name Thrinax is a senior syn- onym of Hemitaxonus and not available for a subdivision of Strongylogaster s.l. as it 698 was understood by most authors so far. It should be pointed out that Zhelochovtsev (1988) has already combined the name Thrinax with struthiopteridis Forsius, but without giving an explanation. The change of generic name involves the following taxa: Thrinax Konow 1885 Thrinax Konow 1885: 19, 22—23. Type spe- cies: Thrinax contigua Konow 1885 [= Sahlbergia struthiopteridis Forsius 1910], subsequent designation § by MacGillivray (1908), = Strongylogaster (Thrinax) = Hemitaxonus Ashmead 1898: 311. Type species: Taxonus dubitatus Norton 1868, by original designation. New synonym. = Epitaxonus MacGillivray 1908: 365— 366. Type species: Taxonus albidopictus Norton 1869, by original designation. Synonymy by Rohwer (1911) = Sahlbergia Forsius 1910: 49-50. Type species: Sahlbergia struthiopteridis For- sius 1910, designation by monotypy. = Sahlberghia, type error. Synonymy by Enslin (1913) = Canonarea Malaise 1947: 37-38. Type species: Canonarea albooralis Malaise 1947, by original designation. Synonymy by Naito (1990) = Trearea Malaise 1947: 2, 35-36. Type species: Trearea compressicornis Mal- aise 1947, by original designation. Syn- onymy by Naito (1990) Thrinax albidopicta (Norton 1868), n. comb. [N]; = TVaxonus albido-pictus Norton 1868 [nec 1869]; = Taxonus ami- cus Norton 1868 [nec 1869]; = Hemitax- onus rufopectus Rohwer 1910b: 204 Thrinax albooralis (Malaise 1947), n. comb. [O]: = Canonarea albooralis Malaise 1947: 38 Thrinax angustata (Zhelochovtsev 1951), n. comb. [EP]; = Hemitaxonus angus- tatus Zhelochovtsev 1951: 145-146 Thrinax athyrii (Naito 1971), n. comb. [EP]; = Hemitaxonus athyrii Naito 1971: 20-21, 26-27 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Thrinax compressicornis (Malaise 1947), n. comb. [O]; = Trearea compressicornis Malaise 1947: 36 Thrinax contigua Konow 1885 [WP, EP]; = Hemitaxonus contigua (Konow 1885); = Sahlbergia struthiopteridis Forsius 1910: 50-51 Thrinax dubitata (Norton 1862), n. comb. [N]; = Taxonus dubitatus Norton 1862: MS Thrinax flexa (Lee & Ryu 1996), n. comb. [EP]; = Hemitaxonus flexus Lee & Ryu 1996 Thrinax formosana (Takeuchi 1928), n. comb. [EP]; = Hemitaxonus formosanus Takeuchi 1928: 43 Thrinax goniata (Wei 1997), n. comb. [EP or O]; = Hemitaxonus goniatus Wei 1997: 1575-1576, 1606—1607 Thrinax japonica (Rohwer 1910), n. comb. [EP]; = Hemitaxonus japonicus Rohwer 1910ar 112 Thrinax kamtchatica (Malaise 1931), n. comb. [EP]; = Hemitaxonus kamtchati- cus Malaise 1931: 22 Thrinax melanogyne (Naito 1971), n. comb. [EP]; = Hemitaxonus melanogyne Naito 1971: 20, 22-23 Thrinax minomensis (Takeuchi 1929), n. comb. [EP]: = Hemitaxonus minomensis Takeuchi 1929: 511-512 Thrinax multicincta (Hall 1918), n. comb. [N]; = Hemitaxonus mutlticinctus Hall 1918 [nec Rohwer, nec 1917]: 28 Thrinax nigrooralis (Malaise 1947), n. comb. [O]; = Canonarea_ nigrooralis Malaise 1947: 39 Thrinax paucipunctata (Malaise 1931), n. comb. [EP]; = Hemitaxonus paucipunc- tatus Malaise 1931: 139-140 Thrinax primaria (D. R. Smith 1966), n. comb. [N]; = Hemitaxonus primarius D. R. Smith 1966: 114, 119-120 Thrinax rufoclypeus (Wei 1998), n. comb. [EP]; = Hemitaxonus rufoclypeus Wei in Nie & Wei 1998: 126 Thrinax sasayamensis (Okutani 1954), n. comb. [EP]; = Hemitaxonus sasayamen- sis Okutani 1954: 76-77 VOLUME 104, NUMBER 3 Thrinax takeuchii (Naito 1971), n. comb. [EP]; Hemitaxonus takeuchii Naito,1971: 20-21, 27-28 Thrinax tokunagai (Takeuchi 1941), n. comb. [EP]: Hemitaxonus tokunagai Takeuchi 1941: 248—249 Thrinax weni (Wei 1998), n. comb. [EP]; = Canonarea weni Wei in Nie & Wei 1998: 125-126 ACKNOWLEDGMENTS Many thanks to the following colleagues for the loan or donation of material: J.-L. Boevé (Bruxelles), R. Danielsson (Lund), E. Diller and J. Schuberth (Munich), R. Eck (Dresden), M. Kraus (Niirnberg), R. Poggi (Genoa) and C. Taylor (London). T. Naito (Kobe), D. R. Smith (Washington), A. Tae- ger (Eberswalde) and an unknown reviewer helpfully commented on all or parts of the manuscript. B. Ewald and C. Kutzscher (Eberswalde) kindly supported me prepar- ing the illustrations, and A. D. Liston (Frontenhausen) checked the English of an earlier version. LITERATURE CITED Ashmead, W. H. 1898. Classification of the Horntails and Sawflies, or the Sub-Order Phytophaga (Paper No. 7.—Conclusion). The Canadian Entomologist 30: 305-316. Benson, R. B. 1954. Some sawflies of the European Alps and the Mediterranean region (Hymenoptera: Symphyta). Bulletin of the British Museum (Nat- ural History), Entomology series, London 3(7): 267-295. Blank, S. M. 1993. A contribution to the sawfly fauna of northern Greece (Hymenoptera, Symphyta). Beitrige zur Entomologie, Berlin 43(2): 431—439. . 1998. Die mittel- und nordeuropiischen Se- landriinae (Hymenoptera: Tenthredinidae), pp. 207-224. In Taeger, A. and S. M. Blank, eds. Pflanzenwespen Deutschlands (Hymenoptera, Symphyta). Kommentierte Bestandsaufnahme. Keltern, 364 + 3 pp. . 2002. Sawflies (Hym., Symphyta) from Tu- nisia. Entomologist’s Monthly Magazine, Bright- wood 138: 68—70. Blank, S. M., A. Taeger, and T. Naito. 1999. 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Insects of the Three Gorge Reservoir Area of Yangtze River. Chongqing. Zhelochovtsey, A. N. 1951. Obsor palearktitscheskich pililschtschikoy podsemejstva. Selandriinae (Hym., Tenthr.). Sbornik Trudov Zoologicheskogo Muzeya MGU, Moskva 7: 123-153. 1988. Pereponchatokrylye, pp. 3-237. In Medvedey, G. S., ed. Opredelitel’ Nasekomikh Evropeyskoy Chasti SSSR. Nauka, 3(6): 271 pp. Zombori, L. 1984. The Symphyta of the Dodero col- lection. 3. The list of species (Hymenoptera). Bol- letino della Societa Entomologica Italiana, Geno- va 116(4—7): 105-120. Leningrad PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 702-715 LIFE HISTORY AND DESCRIPTION OF ADULTS AND IMMATURE STAGES OF GOEDENIA STENOPARIA (STEYSKAL) (DIPTERA: TEPHRITIDAE) ON GUTIERREZIA CALIFORNICA (DE CANDOLLE) TORREY AND A. GRAY AND SOLIDAGO CALIFORNICA NUTTALL (ASTERACEAE) IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e- mail: richard.goeden @ucr.edu) Abstract.—Goedenia stenoparia (Steyskal) is an oligophagous, multivoltine fruit fly (Diptera: Tephritidae) that feeds in the flower heads of hosts belonging to the family Asteraceae, tribe Astereae, and subtribe Solidagininae, like all other known hosts of Goe- denia spp. Newly reported hosts are Ericameria parishii (E. Greene) H. M. Hall, Gutier- rezia californica (de Candolle) Torrey and A. Gray, Hazardia squarrosa (Hooker and Arnott) E. Greene, Lessingia glandulifera A. Gray, and Solidago californica Nuttall. Var- iations in taxonomic characters of adults are described. The third-instar larvae and puparia are described and figured, and selected characteristics of these stages are compared with the same stages of three other species of Goedenia. The prothorax and gnathocephalon of the third instar is smooth and mostly free of the minute acanthae that circumscribe most other body segments. The third instar of G. stenoparia lacks oral ridges, as do the third instars of three other congeners studied to date. The anterior thoracic spiracle bears two papillae. Minute acanthae cover the center of the truncated, posteriorly sclerotized, caudal segment, that also is perforated by scattered pores, and this central area is ringed by concentric, incomplete series of shallow, elliptical depressions. The life cycle is of the aggregative type. Overwintering limitedly occurs as sexually immature adults, but mainly in dead flower heads as prepuparial third instars and puparia in apically open, vasiform cells consisting of floret and achene fragments glued together with dried liquid feces and sap. Overwintered puparia of G. stenoparia were parasitized by chalcidoid Hymenoptera in the genera Eurytoma (Eurytomidae), Pteromalus (Pteromalidae), and Torymus (Tory- midae) as probable, primary, solitary, larval-pupal endoparasitoids. Key Words: Insecta, Ericameria, Goedenia, Gutierrezia, Hazardia, Lessingia, Solidago, Asteraceae, nonfrugivorous Tephritidae, biology, taxonomy of adults and immature stages, flower-head feeding, aggregative life cycle, seed predation, parasitoids Most indigenous, western North Ameri- can Myopitini (Diptera: Tephritidae: Te- phritinae) formerly assigned to the Palearc- tic genus Urophora Robineau-Desvoidy were redesignated as Goedenia by Freid- berg and Norrbom (1999). To date, the life history and immature stages of three of the eight known species of Goedenia have been described in detail, i.e., G. timberlakei (Blanc and Foote), G. rufipes (Curran) and G. setosa (Foote) by Goeden et al. (1995) and Goeden (2002a, b), respectively. This VOLUME 104, NUMBER 3 paper describes the life history and selected immature stages of a fourth species, G. stenoparia (Steyskal). MATERIALS AND METHODS The present study utilized specimens of adults reared from 1-liter samples of mature flower heads of Gutierrezia californica (de Candolle) Torrey and A. Gray collected south of Banner along Chariot Canyon Road at 940 m elevation and Solidago cal- ifornica Nuttall at Banner at 820 m eleva- tion in northeastern San Diego County, Cal- ifornia. The life history study and descrip- tion of the immature stages of Goedenia stenoparia were based in large part on dis- sections of samples of live mature and dead overwintered flower heads of both hosts collected during 1994—1997. One-liter sam- ples of flower heads containing the larvae and puparia were transported in cold-chests in an air-conditioned vehicle to the labora- tory and stored under refrigeration for sub- sequent dissection, photography, descrip- tion, and measurement. Twenty-six, third- instar larvae and four puparia dissected from flower heads of S. californica were preserved in 70% EtOH for scanning elec- tron microscopy (SEM). Prepuparia and pu- paria were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temper- ature for adult and parasitoid emergence. Specimens for SEM were hydraied to dis- tilled water in a decreasing series of acid- ulated EtOH. They were osmicated for 24 h, dehydrated through an increasing series of acidulated EtOH and two, 1-h immer- sions in hexamethyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, studied and digitally photographed with a Philips XL-30 scan- ning electron microscope in the Institute of Geophysics and Planetary Physics, Univer- sity of California, Riverside. Adults reared from isolated prepuparia and puparia were individually caged in 850- ml, clear-plastic, screened-top cages with a cotton wick and basal water reservoir and 703 provisioned with a strip of paper toweling impregnated with yeast hydrolyzate and su- crose. These cages were used for studies of longevity in the insectary of the Depart- ment of Entomology, University of Califor- nia, Riverside, at 25 + 1°C and 14/10 (L/ D) photoperiod. Five pairs of virgin males and females obtained from G. californica and 10 pairs obtained from S. californica in emergence cages were held in each of 15, clear-plastic, petri dishes each provisioned with a flattened, water-moistened pad of ab- sorbant cotton spotted with honey (Head- rick and Goeden 1994) for observations of courtship and copulation behavior. Plant names used in this paper follow Hickman (1993) and Bremer (1994); te- phritid names follow Foote et al. (1993). Terminology and telegraphic format used to describe the immature stages follow Goe- den (2001 a, b;:c; 2002a; b)y Goedenvetial: (1993), Goeden and Headrick (1992), Goe- den and Norrbom (2001), Goeden and Teer- ink (1997), Headrick et al. (1996), Teerink and Goeden (1999), and our earlier works cited therein; Means: vSE.careaused throughout this paper. Digitized photo- graphs used to construct text figures were processed with Adobe Photoshop® Version 6. RESULTS AND DISCUSSION TAXONOMY Adult.—Goedenia stenoparia was de- scribed in the genus Urophora by Steyskal (1979) from a single male collected by E. I. Schlinger from Glendale in Los Angeles, California, on 30.vii.1955. Freidberg and Norrbom (1999) reclassified it along with most other indigenous, western North American species formerly placed in Uro- phora as Goedenia. Steyskal (1979) provid- ed a drawing of the wing and Foote et al. (1993) figured the wing pattern and the head and thorax in lateral view. A total of 211 specimens of adults reared from five hosts, i.e., 72 from Gutierrezia californica (de Candolle) Torrey and A. 704 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Gray, four from G. microcephala (de Can- dolle) A. Gray, 16 from G. sarothrae (Pursh) Britton and Rusby, 93 from Hazar- dia squarrosa (Hooker and Arnott) E. Greene, and 26 from Solidago californica, were examined to estimate variation in the main characters used to distinguish Goe- denia stenoparia. Foote et al. (1993) stated that “‘From all native Urophora [i.e., Goe- denia| species with a distinctively banded wing pattern and primarily yellow scutel- lum, sfenoparia is distinguished by the nar- row wing bands, the failure of the discal band to attain the hind margin of the wing , and the nearly complete break in the subapical band along vein M....” One third of the 211 specimens examined had a black scutellum; the central third or half of the scutellum of the remaining flies were yellow or tawny yellow. The discal band attained the hind margin in 78 (37%) of the 211 flies, but the part posterior to vein Cu, is faint posteriorly in all but 15 (5%) of the 211 flies, i.e., seven males and eight females. It is noteworthy that the dis- cal band was broken in cell dm in 71 (34%) flies, 1.e., 57 males and 14 females, and was narrow and/or faint in cell dm in an addi- tional 38 (18%) flies, including 20 males and 18 females. In addition, the discal band im 1 390f the 7 1! fies, 1.6, nine males and four females, also was broken in cell r,,; or was faint and/or narrow in r,,,; In seven more flies. Finally, one male and two fe- males (0.1%) had the discal band intact in cell dm, but broken in cell r,,;. The presence of a nearly complete break in the subapical band along vein M as a diagnostic character also showed variation. Only 74 (36 %) of 207 adults, i.e., 43 males and 31 females, examined for this character showed a complete or ‘“‘nearly complete” break. Another distinguishing character ascribed to G. stenoparia by Steyskal (1979) and Foote et al. (1993) is the notopleuron yel- low or tawny at the base of the posterior notopleural seta. This character was empha- sized in Figure 507 in Foote et al. (1993). In 64 (30%) of the 211 flies, i.e., 31 males and 33 females, the area at the base of this seta was dark brown or black. Foote et al. (1993) described the legs as yellowish, with only the hind femur slightly darkened basally. The legs of all but one of 210 flies examined for this character would better be described as tawny; moreover, 45 (21%) of these flies, i.e., 34 males and 11 females, had a more extensively darkened hind femur, including some with a substan- tial basal darkening of all femora, and two males with a dark basal part of the tibia. Finally, only 31 (15%) of the 209 flies examined had the gena not more than 1/5 as high as the eye as suggested by Foote et al. (1993), the remaining flies showing ra- tios of 0.20—0.27 times the height of eye. Thus, this character definitely is too vari- able to use to distinguish G. stenoparia. Although adults of G. stenoparia show considerable variation, they usually can be separated from G. caurina by a combina- tion of the reduced and fragmented wing pattern and the extensively darkened legs. Likewise, specimens of G. formosa are readily distinguished by their wider and complete discal and subapical bands that extend to the hind wing margin. A hyaline spot often present in the apical band in cell r,,; also distinguishes many specimens of G. formosa (my unpublished data). Fur- thermore, the host plants of G. caurina and G. formosa in southern California differ from those of G. stenoparia, as reported be- low. Immature stages.—Egg: Twenty-nine eggs dissected from immature inflorescenc- es of Solidago californica (Figs. 4A, B) were white, smooth, elongate-ellipsoidal, 0.56 + 0.03 (range, 0.52—0.58) mm in length and 0.16 + 0.01 (range: 0.14—0.16) mm in width, with a 0.02-mm, peglike ped- icel at tapered, anterior end. Anterior end smoothly rounded. The eggs of G. stenoparia on average were slightly longer and wider than the ova of G. timberlakei (Goeden et al. 1993) and only slightly longer and narrower than VOLUME 104, NUMBER 3 those of G. rufipes (Goeden 2002a), but otherwise generally agreed with published descriptions of those species. Third instar larva: Elongate-ellipsoidal, roundly tapered anteriorly, bluntly truncat- ed posteriorly (Fig. 1A), integument white, but venters of meso-, metathorax, and ab- dominal segments Al—A4 with dark brown to black infuscation (Fig. 4C); caudal seg- ment dark brown or black; minute acanthae, apically pointed or rounded, conical and posteriorly directed (Figs. 1B-1 D-2; 2A-2) or hemispheroidal (Figs. 1B-2, 2D-1, 3B-1, C-2, D-2), circumscribe anterior fifth of meso- and metathorax, all but pleura and posterior fifth of abdominal segment A-1, posterior four-fifths of abdominal segments A2-A7, excluding pleura of A7, and cover all eight venters thereof, and posterior, trun- cated surface of caudal segment (Fig. 4A- 1); prothorax smooth, but venter with flat- tened, posteriorly directed minute acanthae (Fig. 1D-1), and circumscribed by inner, medial ring of verruciform sensilla (Fig. 1C-1) and outer, incomplete ring of un- paired, verruciform sensilla (Fig. 1C-2); gnathocephalon (Figs. 1C-3, ID) conical and medially divided by vertical cleft (Fig. 1D-2), dorsal sensory organ well-defined, hemispherical (Figs. 1D-3, E-1); pores above and dorsolateral of each dorsal sen- sory organ (Fig. ID-4), anterior sensory lobe (Figs. 1D-5, E-2) bears terminal sen- sory organ (Figs. 1D-6, E-3); lateral sen- sory organ (Fig. 1E-4), supralateral sensory organ (Fig. IE-5), and pit sensory organ (Fig. 1E-6); two medial, integumental pet- als (Fig. E-7); four, papilliform, lateral in- tegumental petals (Figs. 1E-8, F-1) above each mouthhook (Figs. 1D-7, F-2), stomal sense organ (Figs. 1D-8, F-3) ventrolaterad of anterior sensory lobe; mouthhook biden- tate (Figs. 1D-7, F-2), anterior tooth con- cave posteriorly (Fig. 1F-4); median oral lobe laterally compressed, apically pointed (Figs. 1D-10, F-5), separated from labial lobe (Fig. 1F-6); anterior thoracic spiracle on posterior margin of prothorax bears two doliform papillae (Figs. 1C-4, 2A-1, B-1); 705 mesothoracic, lateral spiracular complex with six verruciform sensilla in vertical se- ries (Fig. 2B-2), mesothoracic spiracle not seen; metathoracic lateral spiracular com- plex with nearly closed, lateral spiracle (Figs. 2B-3, C-1) and four verruciform sen- silla in vertical series (Fig. 2B-4); lateral spiracular complex of first abdominal seg- ment with partially closed spiracle (Figs. 2B-5, D-1) and five verruciform sensilla in vertical series (Fig. 2B-6); caudal segment with pair of posterior spiracular plates (Figs. 3A-1, B) surrounded by hemispheri- cal minute acanthae (Figs. 3A-2, B-1, C-2, D-2) interspersed dorsally and medially with open pores (Fig. 3A-3, B-4), these structures ringed by two incomplete, con- centric series of shallow, elliptical depres- sions (Fig. 3A-4), with two tapered stelex sensilla (Figs. 3A-5, C-1) and a verruciform sensillum (Figs. 3A-6, D-1); dorsolateral to each posterior spiracular plate; posterior spiracular plate (Figs. 3A-1, B) bears three smoothly flattened rimae (Fig. 3B-2), ca. 0.01 mm in length, and four spinose, inter- spiracular processes, each ca. 0.005 mm long (Fig. 3B-3). The habitus of the third instar of G. sten- oparia (Fig. 1A) resembles those of G. tim- berlakei (Goeden et al. 1995), G. rufipes (Goeden 2002a) and G. setosa (Goeden 2002b). In all four species, the venters of the thorax, anterior abdominal segments, and the caudal segment are darkly pig- mented (Figs. 5D, E; Goeden et al. 1995; Goeden 2002a, b) and minute acanthae cir- cumscribe the meso- and metathorax and abdomen, and especially noteworthy, the central, posterior surface of the caudal seg- ment, which also is dotted with scattered pores (Figs. 3A-2, B-4; Goeden et al. 1995; Goeden 2002b). This central area is ringed by concentric series of shallow, elliptical depressions in all four species (Figs. 3A-2, B-4; Goeden et al. 1995; Goeden 2002b). The prothorax and gnathocephalon of G. timberlakei and G. rufipes are smooth and free of minute acanthae; whereas, the pro- thoracic venters of G. setosa (Goeden 706 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Third instar of Goedenia stenoparia: (A) habitus, anterior to left; (B) minute acanthae on venter of abdominal segment A6, anterior to left; (C) gnathocephalon and prothorax, frontal view, 1—verruciform sensilla of inner ring, 2—verruciform sensilla of incomplete, outer ring, 3—gnathocephalon, 4—anterior spiracle; (D) gnathocephalon, frontolateral view, 1—minute acanthae, 2—vertical, medial cleft, 3—dorsal sensory organ, 4— pores, 5—anterior sensory lobe, 6—terminal sensory organ, 7—mouthhooks, 8—stomal sense organ, 9—median oral lobe; (E) 1—dorsal sensory organ, 2—anterior sensory lobe, 3—terminal sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ, 6—pit sensory organ, 7—medial integumental petals, 8—lateral integu- mental petals; (F) oral cavity, ventral view, anterior at top, 1—lateral integumental petals, 2—mouthhook, 3— stomal sense organ, 4—posterior concavity on anterior tooth of mouthhook, 5—median oral lobe, 6—labial lobe. VOLUME 104, NUMBER 3 707 Fig. 2. Third instar of Goedenia stenoparia, continued: (A) |1—anterior spiracle, 2—minute acanthae, (B) lateral spiracular complexes, anterior to right, 1—anterior spiracle, 2—verrucitorm sensilla on mesothorax, 3— lateral spiracle on metathorax, 4—verruciform sensilla on metathorax, 5—lateral spiracle on first abdominal segment, 6— verruciform sensilla on first abdominal segment; (C) close-up of lateral spiracle on metathorax, 1—spiracle, 2—minute acanthae; (D) close-up of lateral spiracle on first abdominal segment, 1—spiracle, 2— minute acanthae. 2002b) and G. stenoparia anteriorly bear a few minute acanthae (Figs. 1D-1). Two me- dial and four lateral integumental petals are present in G. stenoparia (Figs. 1E-7, 8, F- 1), as in G. rufipes (Goeden 2002a); where- as, G. timberlakei has six lateral integu- mental petals (Goeden et al. 1995, unpub- lished data) and G. setosa has five (Goeden 2002b). The lateral-most integumental petal is separated from the stomal sense organ in all four species (Fig. 1F-3; Goeden et al. 1995; Goeden 2002a, b). The third instars of all four species of Goedenia studied to date lack oral ridges on either side of the mouth opening, and ventral or ventrolateral to the stomal sense organ (Bigs, [DES E Gocdencetal> 1995; Goeden 2002a, b, and unpublished data). The mouthhooks of the third instars of G. stenoparia (Figs. 1D-7, F-2), like those of G. timberlakei (Goeden et al. 1995), G. rufipes (Goeden 2002a) and G. setosa (Goeden 2002b), are bidentate. Moreover, a ventral view of the oral cavity (Fig. 1F), like that figured and described for G. rufi- pes (Goeden 2002a), but not obtained for either G. timberlakei (Goeden et al. 1995, unpublished data) nor G. seftosa (Goeden 2002b), showed the concavely scalloped, posterior surface of the anterior tooth (Fig. 1F-4). The anterior spiracle of all four Goedenia 708 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. species bears only two papillae (Figs. 1C- 4, 2A-1; Goeden et al. 1995, Goeden 2002a, b). The lateral spiracular complex of the me- sothorax of G. stenoparia includes six ver- ruciform sensilla in a vertical series (Fig. 2B-2) like that of G. rufipes (Goeden 2002a); whereas, in G. timberlakei, this same complex includes only two verruci- form sensilla (Goeden et al. 1995). Like- wise, the metathoracic lateral spiracular complex of G. stenoparia rufipes includes four verruciform sensilla (Fig. 2B-4), again like G. rufipes (Goeden 2002a); whereas, only two such sensilla occur in G. timber- lakei (Goeden et al. 1995). Finally, five ver- ruciform sensilla in a vertical series com- prise the lateral spiracular complex of the first abdominal segment of G. stenoparia Third instar of Goedenia stenoparia, continued: (A) caudal segment, 1—posterior spiracular plates, 2—minute acanthae, 3—pores, 4—shallow, elliptical depressions, 5—stelex sensilla, 6— verruciform sensillum; (B) posterior spiracular plate, 1—minute acanthae, 2—rimae, 3—interspiracular processes, 4—pores; (C) 1— stelex sensillum, 2—minute acanthae; (D) 1—verruciform sensillum, 2—minute acanthae. (Fig. 2B-6), three such sensilla occur in G. rufipes (Goeden 2002a), but only one ver- ruciform sensillum is found on this segment in G. timberlakei (Goeden et al. 1995). Puparium: Light to dark, reddish brown with dark brown to black, anterior stripe or series of spots on venters of meso- and metathorax and abdominal segments A-1 to A-4 (Fig. 5E), and similarly dark, caudal segment, elongate-ellipsoidal, with smooth- ly rounded anterior end, and truncated pos- terior end (Fig. 4A). Anterior end bears in- vagination scar (Fig. 4B-1) and raised, bi- lobed, anterior thoracic spiracles (Fig. 4B- 2). Flattened posterior end of caudal segment studded with smoothly rounded, hemispherical, minute acanthae (Figs. 4C- 1, D-1) interspersed with open pores (Figs. 4C-2, D-2). A pair of raised, hemisphero- VOLUME 104, NUMBER 3 idal, posterior spiracular plates (Fig. 4C-3) each bear three elliptical rimae interspersed with four peg-like, interspiracular process- es. These structures are ringed by shallow, elliptical depressions (Fig. 4C-4). Twenty- three puparia dissected from flower heads of Gutierrezia californica and Solidago cal- ifornica averaged 2.66 + 0.07 (range, 1.92— 327) mm an-leneth: 116 = 0:02. ange, 0.99—1.42) mm in width. DISTRIBUTION AND Hosts Foote et al. (1993) mapped the distribu- tion of Goedenia stenoparia as southwest- ern California along with one location in southwestern Utah, which suggests a wide- spread distribution for this tephritid in west- ern North America north of Mexico. The collective distributions of the host plants listed below also would indicate such a wide distribution for G. stenoparia, which probably also encompasses northern Mexi- co (Hickman 1993). Goedenia stenoparia was reported as reared from G. microcephala, Gutierrezia sarothrae and Ericameria cuneata (as Hap- lopappus cuneatus) (A. Gray) McClatchie by Goeden (1987). Foote et al. (1993) re- ported these rearing records along with a sweep record from Hymenoclea_ salsola Torrey and Gray, a non-host (Goeden and Ricker 1986), which Goeden (1987) used to illustrate the often misleading or useless na- ture of sweep records—no matter how ac- curately the plant and tephritid are identi- fied! Gutierrezia californica and Solidago californica were newly reported in this pa- per as hosts, as are E. parishii (E. Greene) H. M. Hall, Hazardia squarrosa, and Les- singia glandulifera A. Gray. All of these new host records for Goedenia stenoparia are from southern California. All of the aforementioned hosts belong to the family Asteraceae, tribe Astereae, subtribe Soli- dagininae (Bremer 1994), as do all other reported and confirmed hosts of Goedenia spp. (Goeden 1987, Freidberg and Norrbom 1999). Accordingly, G. stenoparia is classed as narrowly oligophagous (on eight 709 Fig. 4. Puparium of Goedenia stenoparia: (A) habitus, anterior to right, (B) anterior end, |—invagi- nation scar, 2—anterior spiracles; (C) caudal segment, |—minute acanthae, 2—pores, 3—posterior spiracular plates, 4—shallow, elliptical depressions. species of one host subtribe in the Astera- ceae) (Goeden 1987, Headrick and Goeden 1998). As noted above, their host plants also PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 104, NUMBER 3 serve to distinguish Goedenia formosa and G. caurina from G. stenoparia, at least in southern California. My current opinion is that G. formosa is now known only from Isocoma menziesii (Hooker and Arnott) G. Nesom, formerly treated by Munz (1974) as two varieties of Haplopappus venetus (von Humboldt) Blake, and reported as such as hosts by Goeden (1987). Additional hosts for G. formosa reported by Wasbauer (1972) and Goeden (1987) should refer to either G. sfenoparia, as noted above, to G. caurina, or to an undescribed Goedenia from Grindelia spp. (my unpublished data). Consequently, G. caurina currently is rec- ognized by me as reared from Chrysotham- nus parryi (A. Gray) E. Greene, C. visci- diflorus (Hooker) Nuttall, Ericameria eri- coides (Lessing) Jepson, and E. palmeri (A. Gray) H. M. Hall (Goeden 1987, unpub- lished data). Unfortunately, my coworkers and I did not study G. caurina or G. for- mosa in the aforementioned hosts, as their taxonomic distinction from G. stenoparia was only recently better clarified, as re- ported above. BIOLOGY Egg.—No flower head of Gutierrezia californica containing an egg of Goedenia stenoparia was sampled; however, it 1s as- sumed that a single egg is inserted into each young preblossom flower head based upon dissections of flower heads each containing a single larva as reported below. The few, solitary eggs observed in very young, closed preblossom heads of Solidago cali- fornica supported this assumption (Fig. 5A); however, a cluster of three eggs found within the apical leaves covering a young, compound, terminal infloresence of S. cal- eo Bigg: a ifornica, also containing very small, pre- blossom flower heads (buds) (Fig. 5B) in- dicated an alternative mode of oviposition for this tephritid. None of these eggs were inserted within plant tissues, observations supported by empty chorions found adja- cent to newly eclosed first instars observed within a few other stem terminals. Larva. Upon eclosion from the egg, each first instar commenced to feed sepa- rately within a fast-developing, preblossom flower head on the florets therein. The re- ceptacle was neither abraded or pitted by such feeding. Single second instars were found feeding on ovules in eight, separate, closed, pre- blossom flower heads of Gutierrezia cali- fornica (Fig. 5C). They usually fed within an ovule with their bodies perpendicular to, but always above, the receptacles (Fig. 4A). Receptacles of the eight flower heads con- taining second instars averaged 0.62 + 0.03 (range, 0.57—0.71) mm in diameter. These larvae had damaged an average of 3.0 + 0.7 (range, 2—4) ovules, or about 46% of the average total of 6.5 + 0.4 (range, 5—8) ovules counted within each of the eight flower heads. However, 516 flower heads were individually dissected in order to lo- cate these eight flower heads infested with second instars (Fig. 5C). Likewise, single second instars were found feeding on ovules in 11 separate, closed, preblossom flower heads of S. cal- ifornica (Fig. 5C). All were found within adjacent flower heads on several compound infloresences, again indicative of oviposi- tion of eggs in clusters. They, too, usually fed on ovules with their bodies perpendic- ular to, but always above, the receptacles (Fig. i5C)s\ Receptacles of) the: Id) filower Life stages of Goedenia stenoparia in flower heads of Gutierrezia californica and Solidago califor- nica: (A) single egg (arrow) in S. californica; (B) three eggs (arrows) in S. californica; (C) second instar (arrow) in S. californica, (D) third instar feeding in receptacle having consumed other contents of flower head of G. californica (note dark infuscation on venter); (E) third instar in S. californica; (F) puparium in flower head of G. californica; (G) mating pair, dorsal view; (H) mating pair, lateral view; (1) unreceptive female depressing Oviscape to avoid coitus. Lines = | mm. TW heads containing second instars averaged 1.00 + 0.04 (range, 0.85—1.14) mm in di- ameter. These larvae had damaged an av- erage of 2.1 + 0.3 (range, 1—4) ovules, or about 46% of the average total of 10.6 + 1.3 (range, 8-15) ovules counted within each of the 11 flower heads. Third instars fed with their long axes ori- ented perpendicular to the receptacles, and with their mouthparts directed towards the receptacles, which they scored or pitted, lightly when young to deeply when fully grown, in all of 38 infested flower heads examined of G. californica (Fig. 5D). The 38 flower heads (10 closed preblossom; 28 dead, overwintered) each contained a single third instar (Fig. 5D). These 38 flower heads averaged 0.75 + 0.03 (range, 0.85— 1.42) mm in diameter and contained an av- erage of 8.9 + 0.2 (range, 4—11) ovules/ achenes, all of which were damaged or completely destroyed by the time of pre- pupariation (Fig. 5D). Similarly, third instars fed as above in all of 50 infested flower heads examined of S. californica (Fig. 5E). The 50 flower heads (7 closed preblossom or 43 at least partly open, postblossom and senescent or dead and overwintered) each contained a single third instar. These 50 flower heads averaged 1.14 + 0.02 (range, 0.42—0.85) mm in di- ameter and contained an average of 17.2 + 0.5 (range, 12—22) ovules/achenes, all of which were damaged or completely de- stroyed by the time of prepupariation (Fig. SE)! Pitted receptacles in both of the above host species (Figs. 5D, E) suggest that sap constitutes at least part of the diet of third instars of G. stenoparia, as reported also for G. timberlakei (Goeden et al. 1995), G. ru- fipes (Goeden 2002a), and G. setosa (Goe- den 2002b). Goeden (1988), Headrick and Goeden (1990), Goeden and Headrick (1992), Goeden et al. (1995), Headrick et al. (1996), and Goeden and Teerink (1997) also first noted, described, and discussed sap feeding by florivorous species of Te- phritidae in the genera Trupanea, Paracan- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tha, Neaspilota, Tephritis, Dioxyna, and Xenochaeta, respectively. Upon completing feeding in flower heads of Gutierrezia cal- ifornica, the larva constructed a vasiform cell consisting of ovule/achene/floret frag- ments impregnated with and hardened by dried, liquid feces and sap. The wall of these cells in S. californica was noticably thinner than in G. californica. The flattened, sclerotized caudal segment of the third in- star nicely serves as a plug that tightly clos- es the mouth of the cell, yet allows respi- ration, and thus apparently also may serve to shield the larva from arthropod predators (but certainly not parasitoids, see below) during overwintering (Goeden 2002b). Pri- or to pupariation the prepupa turns 180° and orients with its anterior end away from the receptacle, retracts its mouthparts, and forms a puparium (Figs. 5F). Pupa.—Infested, overwintered flower heads each contained a single puparium (Figs. 5F). The posterior of the puparium faced the receptacle, rested in the cuplike base of the cell, and all but the anterior part of the puparium tightly adhered to the inner wall of the cell in G. californica (Fig. 5F). However, puparia in S. californica flower- heads were only loosely confined. Adult.—Under insectary conditions, 45 males of G. stenoparia lived an average of 28 + 2.5 (range, 7—77) days, and 31 fe- males lived an average of 25 + 2.6 (range, 6—53). These longevities were less than the mean longevities reported for G. timberlak- ei (Goeden et al. 1995) and G. setosa (Goe- den 2002b), but still are commensurate with the aggregative type of life histories pos- sessed by all three of these tephritids (Headrick and Goeden 1994, 1998; Goeden etal) 1995): The premating and mating behaviors of G. stenoparia were not studied in the field, but were observed for five pairs (one male and one female each) reared from Gutier- rezia californica and 10 pairs from S. cal- ifornica in separate petri dish arenas. These arenas were of the type found to be useful in studying mating behaviors of many other VOLUME 104, NUMBER 3 nonfrugivorous, tephritid species (Headrick and Goeden 1994). Both sexes exhibited wing hamation (Headrick and Goeden 1994) throughout the day concurrent with other behaviors, i.e., grooming, resting, and feeding; this also was the most common wing movement reported for G. timberlake (Goeden et al. 1995), G. rufipes (Goeden 2002a), and G. setosa (Goeden 2002b). Pre- mating behaviors observed with G. steno- paria included males and females tracking individuals of the opposite sex, during which males sometimes swayed and usually exhibited abdominal pleural distension. During mating, the wings of the male were overlapped (Fig. 5G) or parted at 10—60°, the wings of the female were parted at 60— 90° (Fig. 5G), with both pairs of parted wings centered over the midline of each fly (Hiss? 5G)> The foretarsi “of ‘the’ male grasped the dorsum of the abdomen of the female laterally at the thoracic juncture, the midtarsi grasped the abdomen laterally or the oviscape at its base, and the hindtarsi crossed under the oviscape (Fig. 5H). The bodies of both flies paralleled the substrate with the oviscape elevated about 45° (Fig. 5H). In arenas, the flies mated at least once on successive days at different times during daylight; six pairs were observed to mate twice in one day. Another especially active pair was observed mating 28 times on 21 consecutive days, after which, the female died. A total of nine matings were observed that lasted an average of 128 (range, 38-— 312) min. Mating females were observed to walk about the arenas, to groom, drink, and form droplets (Headrick and Goeden 1994, 1998). Females became restless before ter- mination of mating and pushed against the males with their hind tarsi, they also lofted their wings so as to push against the males and fully extended their aculei. The male in turn countered this agonistic behavior with copulatory induction behavior (CIB), 1.e., rubbed his hind tarsi along the oviscape, grasped the female tightly, rocked from side to side to regain purchase or to avoid the female’s pummeling, and sometimes rapid- 7fite: ly vibrated his wings, all of which appeared to calm the female and allow coitus to con- tinue. During postcoital separation, the male turned and rapidly walked off and away from the female while pulling his genitalia free, a process lasting just 5 and 8 Ss in two cases. Nonreceptive females avoid- ed matings by decamping, by deflexing their Oviscapes to the substrate (Fig. 51) or otherwise by physically preventing males from gaining purchase by means of agonis- tic behaviors described above. Seasonal history.—The life cycle of Goe- denia stenoparia in southern California fol- lows an aggregative pattern (Headrick and Goeden 1994, 1998) in which the prepu- paria, puparia, and a few adults variously are the overwintering stages. Some adults emerged from a few puparia formed in late- fall (October-November) and these unmat- ed, sexually immature adults overwinter. The remaining prepuparia and puparia over- winter in cells in dead flower heads that re- main attached to dead inflorescences of Gu- tierrezia californica, S. californica, and presumably the other hosts of Goedenia stenoparia. These overwintered individuals emerge as adults in spring (April-May) and pass the following summer (June—Septem- ber), probably as non-reproductive individ- uals in riparian habitats. They eventually aggregate on preblossom, fall-blooming, host plants, to mate, and subsequently to oviposit in or alongside of the small, newly- formed, closed, preblossom flower heads. Natural enemies.—Overwintered puparia of G. stenoparia were parasitized by chal- cidoid Hymenoptera in the genera Euryto- ma (Eurytomidae), Pteromalus (Pteromali- dae), and Torymus (Torymidae) as probable primary, 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- 714 tioned in this paper. Krassimer Bozhilov in the Institute of Geophysics and Planetary Physics, University of California, River- side, greatly facilitated my scanning elec- tron microscopy. I also am grateful to Jeff Teerink for his technical assistance and to David Headrick for his helpful comments on an earlier draft of this paper. LITERATURE CITED Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc. Portland, Oregon. Foote, R. H., EL. Blanc; and A: L.’Norrbom. 11993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York. Freidberg, A. and A. L. Norrbom. 1999. A generic reclassification and phylogeny of the Tribe My- opitini (Tephritinae). Chapter 23. /n Aluja, M. and A. L. Norrbom, eds. Fruit Flies (Tephritidae): Phy- logeny and Evolution of Behavior. CRC Press, Boca Raton, Florida. 944 pp. Goeden, R. D. 1987. Host-plant relations of native Urophora spp. (Diptera: Tephritidae) in southern California. Proceedings of the Entomological So- ciety of Washington 89: 269-274. 1988. Life history of Trupanea imperfecta (Coquillett) on Bebbia juncea (Bentham) Greene in the Colorado Desert of southern California (Diptera: Tephritidae). Pan-Pacific Entomologist 64: 345-351. . 2001la. Life history and description on im- mature stages of Neaspilota footei Freidberg and Mathis (Diptera: Tephritidae) on Aster occidental- is (Nuttall) Torrey and A. Gray (Asteraceae) in southern California. Proceedings of the Entomo- logical Society of Washington 103: 191-206. . 2001b. Life history and description on im- mature stages of Tephritis joanae Goeden (Dip- tera: Tephritidae) on Ericameria pinifolia (A. Gray) H. M. Hall (Asteraceae) in southern Cali- fornia. Proceedings of the Entomological Society of Washington 103: 586—600. . 2001c. Life history and description of im- mature stages of Tephritis teerinki Goeden (Dip- tera: Tephritidae) on Hulsea vestita A. Gray (As- teraceae) in southern California. Proceedings of the Entomological Society of Washington 103: 807-825. . 2002a. Life history and description of im- mature stages of Goedenia rufipes (Curran) (Dip- tera: Tephritidae) on JIsocoma acradenia (E. Greene) E. Greene in southern California. Pro- ceedings of the Entomological Society of Wash- ington 104: 576—S88. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 2002b. Life history and description of im- mature stages of Goedenia setosa (Foote) (Dip- tera: Tephritidae) on Ericameria brachylepis (A. Gray) H. M. Hall in southern California. Proceed- ings of the Entomological Society of Washington 104: 629-639. 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. Goeden, R. D., D. H. Headrick and J. A. Teerink. 1993. Life history and descriptions of immature stages of Tephritis arizonaensis Quisenberry (Diptera: Tephritidae) on Baccharis sarothroides Gray in southern California. Proceedings of the Entomological Society of Washington 95: 210— yp) . 1995. Life history and description of imma- ture stages of Urophora timberlakei Blanc and Foote (Diptera: Tephritidae) on native Asteraceae in southern California. Proceedings of the Ento- mological Society of Washington 97: 779-790. Goeden, R. D. and A. L. Norrbom. 2001. Life history and description of adults and immature stages of Procecidochares blanci n. sp. (Diptera: Tephriti- dae) on Isocoma acradenia (E. Greene) E. Greene (Asteraceae) in southern California. Proceedings of the Entomological Society of Washington 103: 517-540. Goeden, R. D. and D. W. Ricker. 1986. The phytoph- agous insect fauna of the desert shrub, Hymeno- clea salsola, in southern California. Annals of the Entomological Society of America 79: 39-47. Goeden, R. D. and J. A. Teerink. 1997. Life history and description of immature stages of Xenochaeta albiflorum Hooker in central and southern Cali- fornia. Proceedings of the Entomological Society of Washington 99: 597-607. Headrick, D. H. and R. D. Goeden. 1990. Resource utilization by larvae of Paracantha gentilis (Dip- tera: Tephritidae) in capitula of Cirsium califor- nicum and C. proteanum (Asteraceae) in southern California. Proceedings of the Entomological So- ciety of Washington 92: 512—520. 1994. Reproductive behavior of California fruit flies and the classification and evolution of Tephritidae (Diptera) mating systems. Studia Dip- terologica 1(2): 194—252. . 1998. The biology of nonfrugivorous tephritid fruit flies. Annual Review of Entomology 43: 217-241. Headrick, D. H., R. D. Goeden and J. A. Teerink. 1996. Life history and description of immature stages of Dioxyna picciola (Bigot) (Diptera: Te- phritidae) on Coreopsis spp. (Asteraceae) in VOLUME 104, NUMBER 3 aS southern California. Proceedings of the Entomo- — Steyskal, G. C. 1979. Taxonomic studies on fruit flies logical Society of Washington 98: 332-349. of the genus Urophora (Diptera: Tephritidae). Hickman, J. C., ed. 1993. The Jepson Manual. Uni- Special Publication, Entomological Society of versity of California Press. Berkeley and Los An- Washington. 61 pp. geles. Teerink, J. A. and R. D. Goeden. 1999. Description of Munz. P. A. 1974. A Flora of Southern California. Uni- the immature stages of Trupanea imperfecta (Co- versity of California Press, Berkeley and Los An- quillett). Proceedings of the Entomological Soci- geles. ety of Washington 101: 75-85. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 716-720 RANGE EXTENSIONS OF NEW WORLD TORTOISE BEETLES (COLEOPTERA: CHRYSOMELIDAE: CASSIDINAE) CAROLINE S. CHABOO Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, U.S.A. (e-mail: chaboo@amnh.org) Abstract.—New country records with localities are provided for 63 species of New World Cassidinae. Data are based on specimens in the National Museum of Natural His- tory, Smithsonian Instituion, Washington, D.C., U.S.A. Host plant data from specimen labels, including several new records, are reported for some species. Resumen.—Se sumistran nuevos registros para algunos paises con localidades para 63 especies de escarabajos cassidines americanos. Los datos estan basados en la colecci6n del Museo Nacional de Historia Natural (E.U.). Se sumistran datos de plantas hospeda- doras a partir de etiquetas de algunos especimenes, incluyendo nuevos registros. Key Words: The subfamily Cassidinae was recently catalogued by Borowiec (1999). By updat- ing taxonomic names and providing known host records, distributional data, and a bib- lography, Borowiec established a baseline for further research on the Cassidinae. The large collection of Cassidinae at the Nation- al Museum of Natural History, Smithsonian Institution, Washington, DC, was not per- sonally examined by Borowiec in the prep- aration of the catalogue. The strength of this collection is in its representation of New World material. Based on my exami- nation of specimens, I here provide data that extend the distributions of 63 New World species in 15 genera and give avail- able associated host plant information. METHODS Names follow Borowiec (1999) with genera and species listed in alphabetical or- der. Data are given as on labels, with the month of collection in roman numerals and the number of specimens in parentheses. Chrysomelidae, Cassidinae, distribution, host plants Plant data from specimen labels are indi- cated and discussed. Data are limited to species which I could identify. Dissections or type examinations were not done to con- firm names. New Range Extensions in the Subfamily Cassidinae Agroiconota propinqua (Boheman).— Known distribution: Colombia, Costa Rica, Cuba, Nicaragua, Panama. New records: Dominican Republic: San_ Cristobal, 26.VIL 1917 coll] EL Morisson’ (2) Nor San Cristobal, at Blanton Mine, 26. VII.1917, coll. H. Morisson (54); Santo Domingo, 1O:VE1I934—) coll eS, Danforth (@)aea Toma, N. of San Cristobal, 9-10.V1I.1969, colls. Flint & Gomez (1); S. Domingo, Guaiamati, VII.1925, coll. H.E. Box (2); Haiti: Pt. Au Prince, coll. R.J. Crew (12); Pt. Au Prince, VIII.1920, coll. G.N. Wol- cott. Jamaica: Chapinand, 13.VIII.1935, coll. Blackwelder (20); Kingston, IV.1906, coll. V. Duzee (2); Green Hills, 28.X1.1961, VOLUME 104, NUMBER 3 coll. A.B. Gurney (1); Cockpit City, 13- 20.X11.1966, coll. J. Maldonado (2); Span- ish Town, on prison farm, ex sweet potato hkeonvolvulaceae|5: VILLI? (C1). “USA: Puerto Rico: Penuelas, [X.1960, coll. M. Santiago (4). Venezuela: El] Limon, AR 450 m: 18.VII.1951, coll. E Fernandez (1). Agroiconota subtriangularis Spaeth.— Known distribution: Peru. New record: Bo- livia: Coroico (1). Agroiconota tristriata (Fabricius).— Known distribution: Argentina, Brazil, French Guiana, Paraguay, Peru, Suriname. New records: Venezuela: Suapure, Caura, 21.XII.1899, coll. E.A. Klages (1); Orino- co, Caura River, coll. S.M. Klages (1). Charidotella (Xenocassis) puella (Bohe- man).—Known distribution: Belize, Co- lombia, Costa Rica, Ecuador, Mexico, Nic- aragua. New records: Peru: Omaguas, coll. F Knab (16). Honduras: Tela, coll. W.M. Mann (1), in banana thrash, 10.X.1935, colin R-Wweiinee (193 ka ‘Ceibas:22:X1.1915, col Eye Dyerath)svka ‘Ceiba, 191V.A1911, colly Fy-gbDyer (1); Tegucigalpa; 3:1V.1917 @)tesucigalpar5:1V.1917;, coll) FJ nDyer (2); Tela, ex sweet potato [Convolvula- ceae], coll. W.M. Mann. Discomorpha (Discomorpha) biplagiata (Guérin).—Known distribution: Brazil, Co- lombia, Mexico. The record from Mexico is questionable (Borowiec 1999). New rec- ords: Ecuador: coll. E Campos (13): on floor of banana hold, 16.1V.1936, coll. Scott (1); Chiquinita (2); El Salado, coll. EF Cam- pos (10); Guayaquil, coll. EF Campos (2). Venezuela: Suapure; + «Cauray ‘River, 1.VII.1899, coll. E.A. Klages (10); Suapu- re, Caura River, 12.V.1900, coll. E.A. Kla- gesi@l): Discomorpha (Discomorpha) metallica (Guérin).—Known distribution: Bolivia, Brazil. New record: Peru: Satipo, [V.1944 @): Discomorpha (Discomorpha) miniata (Boheman).—Known distribution: Colom- bia. New record: Trinidad: coll. Wells (1): Siparia, 1.1936 (4). Discomorpha (Discomorpha) variegata 717 (Linnaeus).—Known distribution: Caribbe- an, French Guiana, Suriname. New records: Guyana: Georgetown (1): Blairmont Plan- tation, coll. H.E. Box (3). Discomorpha (Vulpia) fryi (Spaeth). Known distribution: Peru. New record: Bo- livia: [no additional data] (1). Dorynota (Akantaka) bivittipennis (Boh- eman).—Known distribution: Brazil. New records: Peru: Huanuco, X.19°? (pin ob- scures data), coll. A. Miller (1); Tarapato, V-VIII.1886, coll. M. de Mathan (1). Dorynota (Akantaka) distincta (Baly).— Known distribution: Ecuador. New records: Bolivia: Chapare, coll. Zischka (1); Santa Cruz, Buena Vista, Tax, 10.11.1951, coll. A. Martinez (1). Dorynota (Akantaka) viridisignata (Boh- eman).—Known distribution: Argentina, Bolivia, Brazil. New record: Paraguay: Alto Paraguay, Emboscada (1). Dorynota (Dorynota) nodosa (Bohe- man).—Known distribution: Colombia, Panama. New records: Venezuela: Suapure, Caura River, 18.1.1900, coll. E.A. Klages (1); Cua, Estado Miranda, coll. C.H. Ballou @):; Dorynota (Dorynota) pubescens Blake.—Known distribution: Dominican Republic. New records: Haiti: 3.111.1950, coll. G.N. Wolcott (11); Constanza, 3—4000 ft, VIII.1938 (1). Echoma (Echoma) bonfilsit§ (Bohe- man).—Known distribution: Ecuador, Peru. New record: Brazil: [no additional data] (1). Echoma (Echoma) clypeata (Panzer).— Known distribution: Bolivia, Brazil, Ecua- dor, French Guiana, Guyana, Paraguay, Peru. New records: Venezuela: Suapure, Caura River, 4.VIII.1899, coll. E.A. Klages (1); Suapure, Caura River, 22.1X.1899, coll. E.A. Klages (1); Suapure, Caura River, 6- 12.1X.1899, coll. E.A. Klages (1); Cojedes, San Carlos DEFORSA, 150 m, 7.X.1997, colls. C.J. Rosales, S. Kiefer & J. Segovia (3); Cojedes, San Carlos DEFORSA, 150 m, 9.X.1997, coll. C.J. Rosales (3). Echoma (Echoma) quadristillata (Bohe- man).—Known distribution: Costa Rica, 718 Nicaragua. New record: Guatemala: Puerto Barrios, 24.11.1905 (4). Eutheria piperata (Burmeister).— Known distribution: Uruguay. New record: Argentina: Buenos Aires, Biolujan Tigre, coll. Martinez (1). Gratiana insculpta (Boheman).—Known distribution: Mexico. New records: Pana- ma: Canal Zone, coll. G. Irenao (1); Canal Zone, rd to Corozal, [X.1912, coll. H.G. Ir- eneo (2). Metrionella angularis (Champion).— Known distribution: Costa Rica, Nicaragua. New records: Panama: Bocas del Toro, 3.VII.1908, ex Ipomoea sp. [Convolvula- caeae], coll. W. Robinson (1). This is the first host plant record for this genus of 12 species. Microctenochira aberrata (Weise).— Known distribution: Costa Rica, Guatema- la, Nicaragua. New records: Honduras: La Ceiba, Rio Congrejal, south, 3.1X.1979, coll. G.V. Manley (1); La Ceiba, Rio Con- grejal, south, 11.1X.1979, coll. G.V. Manley (1). Microctenochira aciculata (Bohe- man).—Known distribution: Brazil. New records: Argentina: Misiones, San Ignacio, coll. Baden (1). Paraguay: Caagualu, Paso Yabai dis X1s195)1> coll) Foerster (1): Microctenochira aspersa (Champion).— Known distribution: Costa Rica, Panama. New records: Colombia: Nicanse, in banana debriss 72x 1936; collx iE Charles (1): Guatemala: Quirigua, 7.V.1926, coll. J.M. Aldrich. Mexico: in banana _ debris, SC EIS3 6s "colle iG: Cn Martin (2); Oax: Tuxtepex, [V.1934 (6); San Antonio, 5.H1.1968, ex palm fronds [Aracaceae], coll. D. Johnson. Palms are probably an un- reliable host plant record since records for 10 of the 104 Microctenochira indicate only dicotyledonous host plants. Also, the tribe Cassidini is not known to be a palm- feeding group. Microctenochira bonvouloiri (Bohe- man).—Known distribution: Guatemala, Mexicomm New iereconds: US Acailexas, Brownsville, coll. Townsville (1); coll. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Wickham (3); Esper. Rch. (1); 24.V1I.1938 (15). Microctenochira_ diffinis (Boheman).— Known distribution: Bolivia, Colombia, Ec- uador, French Guiana, Peru. New records: Guyana: Demerara, coll. R.J. Crew (1). Su- riname: Kwakoegron, Saramacca R., 135ViIEI927 (1); Brokopondos District: Brownsberg Naturpark, Mazaroni Plateau, 4-500 m, 19-25.VIHII.1982, Malaise trap, coll. W.E. Steiner (1). Microctenochira faimairei (Boheman).— Known distribution: Bolivia, Colombia, Ec- uador, Panama, Peru. New record: Brazil: Prov. Mato Grosso, 1866, coll. P. Germain (1). Microctenochira plebeja (Boheman).— Known distribution: Nicaragua, Mexico. New record: Guatemala: with Dracaena cuttings, det. at Miami, 26.VIII.1969, coll. A.S. Mills (1). Omaspides (Omaspides) clathrata (Lin- naeus).—Known distribution: French Guy- ana, Guyana, Suriname. New record: Bra- zil: Para, Cachimbo, IX.1954, coll. M.A. Ivarenga (1). Omaspides (Omaspides) ellipsigera Spaeth.—Known distribution: Brazil. New record: Peru: Achinamiza;, 10: VIR1925@); Omaspides (Omaspides) specularis Er- ichson.—Known distribution: Colombia, Ecuador, Peru. New record: Brazil: Hyutan- ahan, Rio Purus, coll. S.M. Klages (1). Omocerus (Nebroma) scabrosus (Bohe- man).—Known distribution: Argentina, Brazil. New record: Bolivia: Isiamus, XII, coll. W.M. Mann (1). Omocerus (Omocerus) daeruginosus (Boheman).—Known distribution: Brazil, Suriname, Venezuela. New records: Boliv- ia: Caranavi, coll. Denier (1); Nueva Moka, Gutierrez, XI.1951, coll. Prosen (2). Guy- ana: Blairmont Plantation, coll. H.E. Box @): Omocerus (Omocerus) bicornis (Linnae- us)—Known distribution: Brazil, French Guyana, Suriname. New record: Venezuela: Suapure, Caura River, 24.VI.1900, coll. E.A. Klages (1). VOLUME 104, NUMBER 3 Omocerus (Omocerus ) relucens (Spaeth).—Known distribution: Colombia. New record: Guyana: Blairmont Plantation, coll: ELE. Box (). Omocerus (Omocerus) taurus (Bohe- man).—Known distribution: Colombia, Cuba, Brazil, French Guiana, Guyana, Trin- idad, Venezuela. New record: Jamaica: [no additional data] (1). Omocerus (Paratauroma) casta (Bohe- man).—Known distribution: Costa Rica, Ecuador, Nicaragua, Panama, Venezuela. New record: Colombia: [no additional data] (Gl): Omocerus (Paratauroma) caeruleopunc- tatus (Boheman).—Known distribution: Nicaragua. New record: Panama: [no addi- tional data] (2). Omocerus (Paratauroma) smaragdinus (Boheman).—Known distribution: Brazil. New records: Bolivia: Rurrenabaque, Beni, VII.1925, coll. G.L. Harrington (1); Rurren- abaque, Beni, X, coll. W.M. Mann (1). Co- lombia: Villavincencio, Meta, 18.VII.1938 (1). Omocerus (Platytauroma) truncatus (Boheman).—Known distribution: Argenti- na, Bolivia, Brazil, Colombia, Paraguay. New record: French Guiana: Cayenne (1). Orexita dolorosa Spaeth.—Known dis- tribution: Amazonas. New record: Peru: Sa- tipo, 11.1944 (2). Orexita picta (Boheman).—Known dis- tribution: Brazil. New records: Argentina: Misiones, 1947, coll. Heritier (1). Paraguay: Caaguazu, Paso Yabai, 15.XI.1951, coll. J. Foerster (4); Caaguazu, Paso Yabai, 20X10 1951; coll: J. Foerster (1); Hogenau, X.1940, coll. Schade (1). Orexita sp. New record: Guyana: [no ad- ditional data] (2). This is the first record of the genus in Guyana. Spaethiella miniata (Boheman).— Known distribution: Colombia. New re- cord: Panama: Barro Colorado Is., Canal Zone, 29.XII.1928 (1). Spaethiella costipennis (Boheman).— Known distribution: Brazil, Ecuador, Peru. New record: Bolivia: Santa Cruz, Buena 719 Vista, Tacui, 10.11.1951, ex Marantaceae, coll. A. Martinez (1). This is probably a valid host plant since other Spaethiella spe- cies and closely related tribes are monocot feeders. Host data is known for only 3 of the 33 Spaethiella species and these are in the Arecaceae. Spaethiella erhardti (Boheman).— Known distribution: French Guiana, Suri- name. New record: Peru: Amazon Center for Education and Environmental Research, 28.11.1999, colls. M. Lawson & D. Krabill A). Spaethiella laevicollis (Spaeth)—Known distribution: Colombia. New record: Vene- zuela: Caracas Valley, 20.I1V.1922 (2). Stolas acuta (Boheman).—Known distri- bution: Brazil. New record: Paraguay: 1900, coll. Cosset (1). Stolas aenea (Olivier).—Known distri- bution: Brazil, French Guiana. New re- cords: Paraguay: Sapucay, coll. W.T. Foster (2); San Bernardino, coll. K. Fiebrig (1). Stolas consanguinea (Kirsch).—Known distribution: Peru. New record: Ecuador: Chiquinola (1). Stolas conspersa (Germar).—Known dis- tribution: Brazil. New records: Paraguay: Bernardino, coll. K. Fiebrig (6); Villarica, II.1922, coll. P. Jorgensen (1). Stolas festiva (Klug).—Known distribu- tion: Argentina, Bolivia, Brazil, Uruguay. New records: Paraguay: Col. Independen- cia, 23.01.1951, collyFoerster (2); CCaagua- zu, Paso Yabai, 15.X1I.1951, coll. J. Foerster (2); S. Estanislao, coll. Schade (1, USNM); Hohenau, III.1953, coll. EK Walz (3); Hoh- enau, coll. Schade (2): Hohenau, coll. Pod- tiaguin (2); P. Lopez, coll. Podtiaguin (5); Colonia, San Lazaro, Rio Apa, coll. Podtia- guin (1). Stolas ignita (Boheman).—Known distri- bution: Brazil. New record: Argentina: Mi- siones, [X.1947, colls. S. Javier, P. Londero & Monros (1). Stolas implexa (Boheman).—Known dis- tribution: Brazil. New record: Uruguay: Cerro Largo, Frail Muer. to Flossdorf (1). Stolas inaequalis (Linnaeus).—Known 720 distribution: Brazil, Suriname. New re- cords: Guyana: Georgetown (1). Venezuela: El] Dorado Bo, 100 m, 25.VIII.1954, coll. GJ Rosalesi@)! Stolas isthmica (Champion).—First host plant data: ex Ipomoea sp. (3); ex [Ipomoea purga (1) (Convolvulaceae); and ex Xan- thosoma sagittifolium (L.) Schott and Endl. (Araceae) (3). Ipomoea is probably a valid host since it is a common one for many cassidines. Among the 170 Stolas species, host data is known for only 14 species and 8 of these are recorded from /pomoea spp. On the other hand, Xanthosoma is probably not a valid host record since derived cas- sidine genera like Stolas tend to feed on dicotyledonous host plants. The Araceae have not been recorded as a host for any cassidines. Stolas lacordairei (Boheman).—Known distribution: Argentina, Brazil, Paraguay, Uruguay. New record: Chile: Sta. Lucia, San Josew19357 coll PiReed"@): Stolas quatuordecimsignata (Bohe- man).—Known distribution: Bolivia. New record: Peru: Sta. Rosa Convencion, 1936, coll. J. Soukoup (6). Stolas subcaudata (Spaeth).—Known distribution: Paraguay. New records: Ar- gentina: Salta, II.1941, colls. Oran, Tabillas & Martinez (1). Bolivia: Trinidad, Rio Iba- re, 3.1.1948, coll. Kuschel (1). Stolas submetallica (Weise).—Known distribution: Peru. New record: Bolivia: Chapare, 400 m, coll. Zitschka (1). Stolas punicea (Boheman).—Known dis- tribution: Belize, Guatemala, Mexico, Nic- aragua. New records: Honduras: Loncotilla, S311:1928;> coll’ P-C. Standley (); La Ceiba; 23-50. VAISTS. coll GVe ‘Stanley “(4)e Ka Ceiba, 1O°VE1978- coll? -G:V_ Stanley "©): a Ceiba,, 1916, coll Eu: Dyer -(): Stolas tristigma (Boheman).—Known distribution: Belize, Brazil, Guatemala, Honduras, Mexico, Nicaragua. New re- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cords: Argentina: Chaco, Resistencia, coll. Monros (1). Paraguay: [no additional data] (1); Chaco Dpto., Boqueron, 18.XII.1959, coll. Foerster (2); Hohenau, Col. Indepen- dencia, 23.1V. 19518 cols) Foerstesu@): Puerto Pinosco, coll. Podtiaguin (2); Caa- guazu, Paso Yobai, 13.X.1951, coll. Foers- (were (IL). Syngambria andreae (Boheman).— Known distribution: Bolivia, Brazil. New record: Paraguay: Hohenau, X.1940, coll. Schade (4). Tapinaspis wesmaeli (Boheman).— Known distribution: Colombia, Costa Rica, Guatemala, Mexico. New record: El Sal- vador: Lake Coatepeque; 19.VII, colls. G.F & S. Hevel (1). DISCUSSION The records reported here document ex- tended distributions of these species. Bo- rowiec’s (1999) catalogue is an invaluable reference and serves as a basis for further systematic and biological work on the Cas- sidinae. Additional range extensions and other biological data are expected for cas- sidines as existing collections are examined and new collections are made. ACKNOWLEDGMENTS I thank Alex Konstantinov for access to collections in his care; Quentin Wheeler, Dave Furth and Steve Lingafelter for their hospitality during my visit; Allan Pardo- Smith and Fernando Merino for help with Spanish translations; and two anonymous reviewers for helpful comments. Funding for my research comes from the Graduate Training Program in Arthropod Systematics between Cornell University and the Amer- ican Museum of Natural History. LITERATURE CITED Borowiec, L. 1999. A world catalogue of the Cassi- dinae (Coleoptera: Chrysomelidae). Biologica Si- lesiae, Wroclaw. 476 pp. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 721—784 THE NEW WORLD TRIBES AND GENERA OF HISPINES (COLEOPTERA: CHRYSOMELIDAE: CASSIDINAE) C. L. STAINES Collaborating scientist, Instituto Nacional de Biodiversidad, Apartado Postal 23-3100, Santa Domingo, Heredia, Costa Rica; current address: 3302 Decker Place, Edgewater, MD 21037, U.S.A. (e-mail: staines.charles@nmnh.si.edu) Abstract.—The tribes and genera of New World hispines are revised and a key to the 98 genera is presented. The traditional division of hispines and cassidines is rejected as inadequate. The name of the subfamily is Cassidinae. The tribe Imatidiini Hincks 1952 is synonymized with Cephaloleiini Chapuis 1875 (new synonym). Oediopalpini Monr6és and Viana 1947 is synonymized with Spilophorini Chapuis 1875 (new synonym). The tribes Delocrariini Hincks 1952 and Hemisphaerotini Hincks 1952 are treated since they are often confused with hispines. Ocnosispa Weise 1910 and Pseudispa Chapuis 1875 are elevated to full generic status. Plicatopalpa Pic is synonymized with Uroplata Chevrolat (new synonym). Serratispa quadricosta, n. gen., n. sp. from Brazil is described and placed in the tribe Sceloenoplini. Key Words: Cassidinae, hispines, genera Hispines (Coleoptera: Chrysomelidae: Cassidinae) are a group of approximately 3,000 species worldwide (Uhmann 1957b, 1958, 1964; Seeno and Wilcox 1982). Cur- rently they are placed in 170 genera and 24 tribes (Seeno and Wilcox 1982). According to the present classification, there are no tribes or genera indigenous in both the Old and New Worlds. Weise (1911) published the most recent key to the New World genera, covering the 49 genera described by that date. He (Weise 1910, 1911) also proposed the tribal clas- sification that is still in use. There has been little change in the higher classification since then. In proposing his tribal classifi- cation Weise latinized the names proposed by Chapuis (1875) but did not attribute the names to Chapuis. Subsequent authors have attributed the tribal names to Weise (1910, 1911) but the author is Chapuis [ICZN Ar- ticle 11(f) (1999)]. Uhmann (1957b, 1964) compiled the most recent catalog, contain- ing 82 genera and 1,391 species. Seeno and Wilcox (1982) recorded 83 genera from the New World. There have been several regional revi- sions of New World hispines. The most comprehensive of these are Monros and Vi- ana (1947) for Argentina and Staines (1996b) for Nicaragua. Other works are Blaisdell (1939), Butte (1968a, b, c, 1969), Clark (1983) and Riley (1985) on certain genera in America north of Mexico; San- derson (1967) on the West Indies hispines; Uhmann (1930a, b, 1934, 1935c, 1937c) on Costa Rican species; Ramos (1998) on Bra- zilian Oxychalepus, and Staines (1996a) the Central American and West Indian Cephal- oleia. Fourteen genera have been treated for the entire New World: Fischer (1935) re- vised Coraliomela, Mecistomela, Alurnus, and Pseudocalaspidea; Spaeth (1938) re- vised Imatidium (now divided into Aslam- 722 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON idium, Imatidium, Parimatidium, and Rhod- imatidium), Staines revised Brachycoryna (1986), Acritispa (1988), Octotoma (1989b, 1994a), Anisostena (1993a, 1994b, c, d), and Physocoryna (1999); and Ramos (1996) revised Clinocarispa. HISTORY OF HIGHER CLASSIFICATION There is some difficulty in clearly defin- ing hispines. While there are a combination of characters to separate them from cassi- dines, various intermediate genera connect the two groups. The line dividing them have been differently placed by different systematists. For example, the genus /ma- tidium Fabricius has been placed in both groups at one time or another. Weise (1910) suggested that the tribes Oediopalpini and Cephaloleiini may belong in the cassidines rather than the hispines, based on life his- tory studies. Barber (1946) described Cub- ispa and placed it in the Hispinae; Monros (1954) argued that it was a Eumolpinae. While the intent of this article is to pro- vide a workable key to the tribes and genera normally placed in the hispines, a short re- view of the literature on the higher classi- fication of the Hispinae-Cassidinae is in or- der. Latreille (1802, 1803, 1807, 1829) pro- posed Hispe as part of the tribe Cassidaires in the family Cycliques. Gyllenhal (1813) proposed the group Cassideae for the genus Cassida Linnaeus and Hispoideae for the genus Hispa Linnaeus. Stephens (1829) proposed the family Hispidae. Westwood (1839) placed the group in the Cassididae in the subsection Phytophaga. Lacordaire (1845—1848) proposed the Hispides in Tribe 4. Chapuis and Candeze (1853) placed the group as the tribe Hispides in Division III. Chapuis (1875) placed the Hispinae and Cassidinae in Section [V—Cryptostomes. Jacoby (1908) followed the same system making Cryptostoma Division V. Jacoby’s basic system has been followed by subse- quent workers, some using family desig- nations for the two groups and others leav- ing them as subfamilies. Chen (1940) cre- ated the family Cassididae with Hispinae and Cassidinae as subfamilies, but most workers have not followed this system. Crowson (1955) included the Cassidinae as part of the Hispinae. Monros (1959) pro- posed the group Cassidiformes with the two subfamilies. Chen (1964) then proposed the family Hispidae with the subfamilies Ani- soderinae, Hispinae, and Cassidinae. Chen (1973) also proposed the new superfamily Cassidoidea with four families: 1. Cassididae (= Cassidinae of the older system). 2. Anisoderidae consisting of the tribes Anisoderini, Cryptonychini, Eurispini, Pro- sopodontini, Alurnini, Pharangispini, Coe- laenomenoderini, Promecothecini, Cephal- oleini, Arescini, Hysbosispini, and Gono- phorini. 3. Hispidae consisting of the tribes Ex- othispini, Chaeridionini, Oncocephalini, Hispini, Sceloenoplini, Hispoleptini, Chal- epini, and Uroplatini. 4. Callispidae consisting of the tribes Callispini, Leptispini, and Oediopalpini. The tribes Hemisphaerotini, Spilophorini, Delocranini, and Imatidiini were not placed in a family by Chen (1973). Suzuki (1985, 1988, 1994) considered the subfamily Cassidinae with two tribes Hispini and Cassidini. Chen et al. (1986) further refined the 1973 classification. Bo- rowiec (1995) considered the subfamily Hispinae and proposed a tribal classification of the genera normally considered Cassidi- nae; one of the proposed changes was the synonymization of the Cassidinae tribe Im- atidiini with the hispine tribe Cephaloleiini. Reid (1995) looked at chrysomelid sub- families using cladistic methods. He com- bined hispines and cassidines into the sub- family Hispinae before he did his analysis. This treatment avoided addressing the ques- tion of monophyly and relative rank of the taxa. Reid stated that there is not enough data available for a resolution of relation- ships without considerable massage of the data; his system is to be considered tenta- tive. Lawrence and Newton (1995) considered VOLUME 104, NUMBER 3 hispines and cassidines as the subfamily Hispinae. Verma (1996) treated Hispinae and Cassidinae as separate subfamilies based on the male genital system. Suzuki (1996) proposed a classification of the sub- family Cassidinae with the tribes Hispini and Cassidini. Farrell (1998) treated Cas- sidinae and Hispinae as separate subfami- lies based on DNA analysis. Hsiao and Windsor (1999) found that there was a complete separation of cassoid and hispoid groups based on DNA analysis of 48 spe- cies. This work however did not include three tribes of hispines (Arescini, Hispolep- tini, and Sceloenoplini) so the results may be modified by future work. It is obvious from the preceding sum- mary that there is uncertainty over the rank and status of the Hispinae and Cassidinae. From a systematic point of view merely changing the rank or name of a taxon does not solve the problem of its relationship to other taxa. I agree with Schmitt (1996) that listing intermediate forms as incertae sedis would clearly signal unsolved problems while lumping two taxa (including the in- termediate forms) would only gloss over our lack of knowledge and not remedy it. Morphologically and biologically there is no valid reason for retaining Hispinae and Cassidinae as separate subfamilies. The presence of intermediate forms (the tribes Basiprionotini, Cephaloleiini, Delocranini, Epistictinini, Hemisphaerotini, Imatidiini, Notosacanthini, Oediopalpini, and Spilo- phorini) effectively bridge the subfamilies which were erected based on Palearctic, Nearctic, and a few tropical genera. The most obvious characters mentioned to sep- arate the subfamilies is that cassidines have the margins of the pronotum and elytra ex- panded, the head often covered by the pron- otum, and the free living larvae possessing paired caudal appendages on the last ab- dominal segment while hispines do not have the expanded elytral and pronotal mar- gins and the larvae are leaf miners and lack caudal appendages. However, these char- acters break down and are possessed by 723 genera placed in different subfamilies. The adult of Notosacantha looks like a typical cassidine but the larvae are leaf miners and have a caudal fork (Medvedev and Erosh- kina 1988, Rane et al. 2000). The adult of Oediopalpa looks like a typical hispine but the larvae are free living and have a caudal fork (Bruch 1905). The larvae of Delocran- ia and some Hemisphaerota (both cassidi- nes and not leaf miners) have the caudal fork reduced or absent (Bondar 1922). The larvae of /matidium (cassidine) is waterpen- ny-like and lacks caudal appendages (Bon- dar 1940). The larvae of Prosopodonta (hispine) are leaf miners and have the ninth abdominal segment modified into append- age-like structures (Maulik 1931) as do the larvae of Brontispa (Maulik 1938). Adults of Pseudocalaspidea and some Sceloenopla (hispines) have the elytral margins expand- ed; Adalurnus and Demotispa (hispines) have both the elytral and pronotal margins expanded. Workers who have proposed the merger of the two groups, with which I agree, usu- ally call the combined subfamily the His- pinae. However, since Gyllenhal (1813) proposed both Cassidinae and Hispinae in the same publication the valid name is Cas- sidinae since it appeared on p. 434 while Hispinae appeared on p. 448 [ICZN Article 23.1 (1999)]. The name would still be Cas- sidinae if one applied the determination of the first reviser [ICZN Article 24.2 (1999)] (Chen 1940). Chen (1940), Suzuki (1985, 1988, 1994, 1996), Reid (1995), and Lawrence and Newton (1995) combined hispines and cas- sidines as a subfamily but retained the di- vision between them at the tribal level. Since hispines and cassidines are not clearly distinct at the subfamily level, merely re- ducing the taxonomic rank does not solve the problem. Either a third tribe would need to be erected to accomodate the intermedi- ate forms which is not acceptable or the en- tire idea of hispines and cassidines as sep- arate taxonomic units needs to be rejected. I reject the traditional cassidine-hispine di- 724 vision as inadequate because of the inter- mediate forms linking the groups and am treating them as the subfamily Cassidinae and going directly to the 42 current tribes. TAXONOMIC TREATMENT The tribe Chalepini was proposed for genera with |1l-segmented antennae while the tribe Uroplatini was proposed for gen- era with 8- or less segments (Weise 1910). Since 1910 the genera Charistena, Deca- telia (in part), Sternostenoides, and Ster- noplispa with 10-segmented antennae have been placed in Chalepini. The remaining species of Decatelia have 9-segmented an- tennae. The genera Cnetispa and Nonispa have 9-segmented antennae and are placed in Uroplatini. These genera totally obscure Weise’s tribal divisions, therefore I am combining the two tribes under the name Chalepini. Bruch (1905) described all life stages of Oediopalpa negligens (Weise) and found the larva and pupa to be cassid-like. Weise (1910) suggested the tribe may belong in the Cassidinae based on the general biology and the larvae having a caudal fork. Mon- ros and Viana (1947) mentioned the differ- ent biology of the group but retained it in the Hispinae where it has remained. Borow- 1ec (1995) did not discuss the Oediopalpini. The cassidine tribes Delocraniini, Hem- isphaerotini, and Spilophorini are often confused with hispines. In fact, Spilophor- ini shares the same morphological charac- teristics as Oediopalpini (a seta present in each pronotal angle and the prosternum projecting forward to partly cover the mouth) and the two tribes are synonymized as Spiliphorini. Delocraniini and Hemis- phaerotini are included in the key for the convenience of workers. I agree with Borowiec (1995) that the cassidine tribe Imatidiini is a synonym of the hispine tribe Cephaloleiini. Reasons not mentioned by Borowiec is that Demotispa (Cephaloleiini) has the same type species as Pseudimatidium (Imatidiini) and the de- scribed larvae of the two groups share the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON same morphology (see Maulik 1931 and Bondar 1940). Hsiao and Windsor (1999) based on DNA analysis found that /mati- dium, Aslamidium, and Rhodimatidium formed a distinct group with Cephaloleia. The genera /matidium, Parimatidium, As- lamidium, Rhodimatidium, Calliaspis, Stilp- naspis, Pseudostilpnaspis, and Spaethaspis are included in this paper. The genus Cubispa is an anomaly. Some characters are hispine-like but an equal number are eumolpine-like. The genus needs extensive study to determine its prop- er placement but it is not a hispine due to the absence of setae in any of the pronotal angles, the structure of the mouthparts, the lack of bifid tarsal setae, and lack of a pars stridens (Staines 2000). R. W. Flowers also considers Cubispa to be an eumolpine (per- sonal. communication.). Seeno and Wilcox (1982) list the genus Cladophora Dejean 1837: 390 as incertae sedis. Dejean (1837) listed this name and included the species flabellata Dejean. Nei- ther name had a description associated with it, so they do not fulfill the requirements of Article 12.1 (CZN 1999) and are nomina nuda. Description.—Head: Opisthognathous, prominent, visible from above, at least to behind eyes; frons prominent, exposed or rarely retracted; antenna not retractable, closely inserted between eyes. Pronotum: Narrower than elytra; more or less quadran- gular or trapezoidal, with definite anterior angle which may have small tubercle. Scu- tellum: Always visible. Elytra: Without lat- eral expansions or with reduced and dis- continuous expansions; margins usually denticulate or with spines. Larvae: Either leafminers or free living; eight pairs of ab- dominal spiracles well developed and dor- sally placed; eighth abdominal segment ter- minal, with free hind margin. Ecologically, New World hispines fall into three feeding groups: external feeders; sheath, appressed or rolled-leaf feeders; and leaf miners. In the Old World some species have been reported as stem borers in her- VOLUME 104, NUMBER 3 Fig. 1: of head; 2 = anterior angle of pronotum; 3 = pronotal Dorsal morphology of Hispinae. | = vertex disc; 4 = lateral margin of pronotum; 5 = scutellum; 6 = humerus; 7 = scutellar puncture row; 8 = elytral costa; 9 = exterior apical angle of elytron; 10 = su- tural angle of elytron. baceous or semi-ligneous plants, but this has not been reported from the New World. The biology of few species has been stud- ied; most species are not associated with a host plant or plant family. ADULT MORPHOLOGY From the literature on hispines, there is no uniform nomenclature for the various body parts. In order to aid in the use of this key, the following terms are defined as they are used throughout the manuscript. They are shown on Figs. | and 2. Occiput: Region surrounding occipital oO Fig. 2. Ventral morphology of Hispinae. | = an- tenna; 2 = eye; 3 = oral cavity; 4 = prosternum; 5 = mesosternum; 6 = metasternum; 7 = femur; 8 = tibia; 9 = tarsus. and postoccipital sutures and in live adults mostly covered by pronotum. Vertex: Separated from occiput by oc- cipital suture, occupies post-ocular region. Eye: Elongate, following dorso-ventral axis, slightly prominent, multifaceted, slightly kidney-shaped. Frons: Upper anterior portion of head capsule between front and clypeus. Clypeus: Quadrangular, elongate, united to labrum. Labrum: Triangular, united to clypeus and oral orifice. Oral orifice: Subelliptical cavity which occupies middle part of head. Antenna: 3- to 11-segments, inserted in antennal pit in front of eyes. 726 Pronotum: Occupies all of dorsal part of prothorax, laterally margined. In anterior angles or posterior angles sometimes a small tubercle with a seta. Scutellum: Sclerotized portion of meson- otum, visible between elytral bases. More or less triangular or quadrate. Elytron: Strongly sclerotized mesotho- racic wings, remaining open during flight. When closed, they cover to last abdominal segment. Elongate and quadrangular form and flat surface, convex at sides. Sculptured with large punctures usually in regular rows; some species carinate. Most species with costae. Sutural margin somewhat raised; lateral and apical margins dentate or not. Prosternum: Contains two anterior coxal cavities. Mesosternum: Narrow between middle coxae, separated from metasternum by a transverse suture. Forms anterior part of middle coxal cavities. Metasternum: Large, with median lon- gitudinal suture. Forms posterior border of hind coxal cavities. Episternum: Anterior and larger lateral thoracic sclerite between sternum and no- tum. Parapleura: Lateral sclerite between dorsal and sternal parts of thorax. Coxa: Located in coxal cavity, spherical. Trochanter: Small, united to base of fe- mur. Femur: Largest part of leg; more or less cylindrical, united at base to trochanter and at apex to tibia. Tibia: Variable in length; joins femur and tarsus. Tarsus: Each with five tarsomeres, pen- ultimate degenerated. Last tarsomere with claws. Basal tarsomere small, triangular; second bilobed, larger than first; third larger than second, with two pronounced lobes; last tarsomere more or less conical, with one or two curved claws at distal end; seg- ments | to 3 flattened, with dense pubes- cence on lower surface. Abdomen: With five visible sternites; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sternites | and 2 mores strongly united that others. Key TO NEw WORLD TRIBES il Elytron without costae or costae are ves- tigial; rows of punctures little impressed; antenna 10- or 11-segmented = Elytron costate; rows of punctures strongly impressed; antenna 3- to 1l-segmented .. 9 Pronotum without seta in any angle (or along anterior margin) (Fig. 3) ........ 3 = Pronotum with seta present in 2 or 4 angles (onfalongvanterior margin) eee 4 Pronotum with prominent, anteriorly pro- jected lobes, rounded in front .. Delocraniini = Pronotum without prominent, anteriorly Projected sMObesis ieee Hybosispini Pronotum with a tuft of setae present in each angle; very large (more than 25 mm.) species; elytral punctures random or with onlyavestices)| of Ordena aioe Alurnini = Pronotum with a single seta present in 2 or 4 angles (or along anterior margin); elytral punctures more or less in rows; smaller Speciess@essithan) 20mm) eee 5) 5(4). Anterior margin of prosternum expanded forward, partially covering mouth (Fig. 5); pronotum with a seta present in all four Spilophorini = Anterior margin of prosternum not expand- ed forward; pronotum with a seta present angles in each anterior or posterior angle (or along anterior margin) Pronotum with a seta present in each pos- tenlonganelemeerea a+ cree Prosopodontini — Pronotum with a seta present in each an- terior angle (or along anterior margin) .. 7 Antennae not separated by a longitudinal keel; pro- and mesocoxae very large... . Se) SEEMS ais tot =e cite ns Gam SOs eee oes Arescini = Antennae separated by a longitudinal keel; coxae normal 8(7). Mesosternum deeply emarginate, emargi- nation deepened for reception of prosternal PFOCESS, 55 eieeets ye ne eee eek eee Ce id os eee Hemisphaerotini = Mesosternum not deeply emarginate ... Ba aS Ee 3 can ch onan eee Cephaloleiini Antenna thickened at apex ...... Chalepini = Antenna not thickened at apex ....... 10 10(9). Apical 4 antennal segments different from others (either longer or shorter, thinner or thicker, more pubescent, different in color); clypeus clearly visible ....... Sceloenoplini = Apical 4 antennal segments similar to oth- ErssiclypeusnObScure sm eer Hispoleptini VOLUME 104, NUMBER 3 TRIBE SPILOPHORINI CHAPUIS 1875: 364 Amplipalpini Weise 1910: 69. Oediopalpini Monros and Viana 1947: 140. New synonymy. Type genus.—Spilophora Boheman. Description.—Head: Second and_ third palpomeres of maxillary and labial palps enlarged, longer and wider than others; clypeus transverse, with upper margin curved, emarginate, overlapping antennal base; antenna |1l-segmented. Pronotum: Pit containing a seta present in each angle, pos- terior setae On cones, anterior Ones on disc or edge of lateral margin. E/ytron: With 10 regular rows of shallow punctures; scutellar row absent; intervals not costate. Venter: Prosternum with front margin elongate, transverse, raised, partially concealing mouth; abdomen with suture between sterna 1 and 2 obsolete in middle. Legs: Apical tarsomere triangular, flattened before claws; claws short. KEY TO GENERA |. Elytra (at base) much wider than base of pron- otumycounded (Bigs: 5, 7)! .2t.ce6 a2 2eun045 2 — Elytra (at base) not much wider than base of pronotum, parallel-sided (Fig. 6) Antennomere III longer than [TV and V com- bined, III—VI with internal angle with an acute SDIMC Arte eye OE ht epee hic ok cera Perens Spilophora — Antennomere II shorter than IV and V com- bined, II—VI internal angle without acute spine Oediopalpa i) Calyptocephala Calyptocephala Chevrolat (Fig. 5) Calyptocephala Chevrolat 1837: 391. Type species: Cassida nigricornis Germar (by monotypy). Description.—Head: Rounded, visible dorsally; front subconvex; labrum large, slightly wider than long, apical margin trun- cate, with medial longitudinal carina; max- illary palp with apical palpomere oblong, longer than preceding; eye oval, slightly convex. Antenna: Filiform; % length of body; 1l-segmented; antennomere I ob- long, obconic; II thinner and shorter than I; 27) Ill 2* as long as II; remainder gradually decreasing in length. Pronotum: 2 as wide as long, at base narrower than base of ely- tra, convex; anterior margin slightly emar- ginate, angles of emargination obtuse; an- terior angle rounded, with a seta in each angle; lateral margin curved, convergent anteriorly; posterior margin almost straight. Scutellum: Elliptical. Elytron: Subtriangu- lar or suboval, wide basally, narrowed and rounded apically, convex; humerus promi- nent, subacute. Venter: Prosternum with apex notched, narrowed between coxae, di- lated apically, truncate basally. Legs: Tibiae rounded, notched on apical 4%, exterior mar- gin subangulate; tarsi wide, tarsomere | narrower than rest, 4 little longer than lobes of 3; claws simple, divericate. Host plants.—Arecastrum, Astrocaryum, Chamaedorea, Elaeis (Arecaceae); Dios- corea (Dioscoreaceae); Chusquea (Po- aceae) (Jolivet and Hawkeswood 1995). Described species.—11. Key to species: Wagener (1881) covered 6 species. Range.—Mexico to Argentina. Oediopalpa Baly (Figs. 4, 6) Oediopalpa Baly 1858: 16. Type species: Hispa cyanipennis Fabricius (designated by Monros and Viana 1947). Amplipala Harold 1875: 185 [invalid re- placement name for Oediopalpa\. Charispa Baly 1875: 73 [invalid replace- ment name for Oediopalpal. Description.—Body more or less elon- gate, moderately convex. Head: Small; ver- tex flat or weakly depressed; eye oblong to oval, finely faceted; mouth small, exposed area covered by palps; mandible dentate. Antenna: 11-segmented; thick; antennom- ere III longest. Pronotum: Transverse, apex narrowed; regularly convex, may be im- pressed in each angle; sometimes with lat- eral impressions; disc with deep punctures. Scutellum: Quadrangular or pentagonal; rounded at apex. Elytron: Little wider than pronotum; slightly constricted behind pron- 728 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Lv aE Wea K 2 6 Sere ner er tease nat wes ee 7 —e . — 8 Figs. 3-11. 3, Setae in anterior angle of pronotum. 4, Oediopalpa sp., prosternum. 5—11, Dorsal habitus. 5, Calyptocephala nigricornis (Germar). 6, Oediopalpa cyanipennis (Fab.). 7, Spilophora trigemina (Guérin). 8, Aslamidium (s. str.) caprense (Herbst). 9, Aslamidium (Neoaslamidium) strandi (Uhmann). 10, Callasapis cin- nabarina Boheman. 11, Cephaloleia nitida Uhmann. VOLUME 104, NUMBER 3 otum, then subparallel; apex conjointly rounded; lateral and apical margins finely serrate or smooth. Legs: Short, thick, fem- ora widest, inner margin serrate or not. Ven- ter: Prosternum almost flat (Fig. 4). Some species orange or reddish-yellow brown; others may be metallic dark blue, green to black. Host plants.—Arundo, Bambusa, Oryza, Panicum, Poa, Saccharum, Zea (Poaceae) (Jolivet and Hawkeswood 1995). Described species.—37. Key to species: Weise (1910) covered twenty species; Mon- ros and Viana (1947), Argentina. Range.—Mexico to Argentina. Spilophora Boheman (ig 7) Spilophora Boheman 1850: 107. Type spe- cies: Calyptocephala trigemina Guérin- Méneville (designated by Monros and Viana 1947). Description.—Head: Rounded, visible dorsally; front concave; labrum large, quad- rate, apical margin triangularly emarginate, convex, subcarinate medially; maxillary palp with palpomere 4 oblong, subtruncate at apex; eye oval, convex. Antenna: % length of body, filiform, 11-segmented; an- tennomere I short, thick, obconic; II shorter and narrower than I; III very long, narrow, as long as IV to VI combined; III—VI with internal angle with an acute spine; VII—XI cylindrical, shorter and narrower than pre- ceding. Pronotum: Transverse, at widest % width of elytra, convex; anterior margin regularly rounded or slightly sinuate in middle; anterior angle obsolete; posterior angle rectangular; posterior margin sinuate. Scutellum: Triangular. Elytron: Convex; hu- merus rounded; lateral margin slightly di- lated. Venter: Prosternum sinuate in middle, slightly narrowed between coxae, flat, base dilated, subtruncate. Legs: Short, robust; femora fusiform; tibiae straight, sometimes subarcuate basally, obliquely truncate api- cally; tarsi robust, dilated, tarsomere | near- ly as wide as others, 4 dilated from base to 729 apex, slightly exceeding lobes of 3; claws simple, divaricate. Host plants.—Unknown. Described species.—16. Key to species: Wagener (1881) covered 7 species. Range.—Colombia to Ecuador and Peru. TRIBE CEPHALOLEIINI CHAPUIS 1875: 277 Callispites Chapuis 1875: 269. Himatidiites Chapuis 1875: 361 (in part). Himatidiitae Spaeth 1929: 113. Imatidiini Hincks 1952: 328. New syno- nym. Type genus.—Cephaloleia Chevrolat. Description.—Body shape approximately parallel or slightly narrowed behind or oval; explanate margin of pronotum and elytra narrow to moderately broad. Head: Suture present at union of oral cavity and antennal bases; without frontal plate; clypeus very short; labrum without carina; neck very short, ledge-shaped. Antenna: 10- or 11- segmented. Pronotum: Wide; a seta present in anterior angle or along anterior margin. Elytron: Noncostate; puncture rows. shal- low. Venter: Metepisternum fused with me- tepimeron. KEY TO GENERA Ie Antenna 10-segmented ....... Calliaspis - Antenna dil-sesmented” 22.22.24 4088 2 21): Mouthparts projecting forward and _ visi- ble from above; labrum large, joining head at an oblique angle; anterior margin of pronotum crenate in middle ....... 3 - Mouthparts not projecting forward; ante- rior margin of pronotum not crenate ... 4 3(2) Antennomere II longest; elytral margins smooth)? 2 2 =. S25 2 Homalispa (Xanthispa) — Antennomere III longest; elytral margins Serate- . 2a eae Homalispa (Homalispa) 4(2) Elytron rounded, broadly oval or widened apicalllynety sata ew we ete eet ee fe ass ice 5 - Elytronyparalllel=sidede. v2: ec ceke. tee 14 5(4). Antenna pectinate or with antennomeres Oded Sepak rene eee eee awa Se) 6 - Antennasyiilifonmpee en te kee Meee ek ee acu 7 6(5) Male antenna pectinate on antennomeres IV to X; female antenna with projections on antennomeres IV to X; pronotum with- out transverse sulcus Octocladiscus — Antennomeres I to X of both sexes with 730 98). 10(7). 11(10). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON projection; pronotum with transverse sul- CUS) agranee eons neeleeve ses oe) Seep Cladispa Interantennal carina distinct ......... 8 Interantennal carina not distinct ..... 10 Antennomere I longer and thicker than II; pronotum with notch external to setifer- ous tubercle on anterior emargination; tibiae obliquely and abruptly emarginate appearing pointed externally Antennomeres I to If subequal in length; anterior margin of pronotum without notches; tibiae not pointed externally Seas Oe CRer ter Or Demotispa Body elongate, parallel-sided:; pronotum rectangular; elytra slightly to moderately wider than pronotum;: prosternal process with deep groove along sides ...... EB Late a ante re Asladmidium (Neoaslamidium) Body broad, sides rounded; pronotum al- most semicircular; elytra much wider than pronotum; prosternal process with several shallow, short grooves at apex ..... SRR croetaoas as Aslamidium (s. str.) Posterior part of elytral margin serrate; posterior angle of pronotum sharp and projecting behind; elytra finely dentate externally toward apex; metepisternum separated from metasternum by a distinct suture Posterior part of elytral margin smooth; posterior angle of pronotum not as above: elytra not dentate; metepisternum more or less fused posteriorly with metasternum Antennomeres I and II globular Pseudostilpnaspis Antennomeres I and II elongate ..... 12 . Antennomere I subequal in length to II ee eect oo gad eee te aces eed Imatidium Antennomere I % the length of II ..... Be eects, © Ot OLE ee es as el pees 13 2). Pronotum with sides almost parallel a Nyse We smnciuiy?: off ries tetas Spaethaspis Pronotum with sides convergent from baSesLOkapexernea eet eee Stilpnaspis Elytra at base slightly wider than prono- CUED ye eeepc isres toyed sed clog caters ea os 15 Elytra at base much wider than pronotum dic) Py Rrwisi galas pe cet OR ee Se RIES 5 16 = Blytraswidenssapicalllya-ssies ieee Euxema Eby tray parallel-sided eis -aeeke Stenispa Pronotum longer than wide; base of ely- tron with carina to humerus; last three ab- dominal sterna hirsute ........ Solenispa Pronotum as wide or wider than long; base of elytron without carina; last three abdominal sterna not hirsute........ 7 17(16). Pygidium for most part visible from above; maxillary palp with palpomeres 2 and 3 smaller than 1 Cephaloleia — Pygidium not visible from above; maxil- lary palp with palpomere 3 longer than | and 2 combined Melanispa Aslamidium Borowiec (Fig. 8) Aslamidium Borowiec 1984: 412. Replace- ment name for /matidium: Aslam (1965: 689) a misapplication of Jmatidium Fa- bricius 1801. Type species: Cassida ca- pense Herbst (by original designation). Description.——Head: Frons concave pos- teriorly; eye projecting; interocular space equal to maximum width of an eye; antenna inserted equidistant between eyes and inter- antennal carina; interantennal carina sharp and united to a transverse carina which en- closes oral cavity behind and around la- brum; labrum very short and transverse; maxillary palp with palpomere 4 thicker and slightly longer than 3. Antenna: 11-seg- mented, narrowed apically; antennomere I thicker and longer than IJ; III at least 2X length of II; [IV to X decreasing in length; XI laterally compressed and longer than X. Pronotum: Small notch on external side of setiferous tubercle within apical emargina- tion; deeply impressed on either side at base; sparsely punctate; lateral margin con- vergent from base to apex, recurved; ante- rior angle rounded. Scutellum: Broadly tri- angular. Elytron: Lateral margin laminate, lamina reticulate; lateral and apical margin smooth; slightly indented behind humerus; with 10 rows of moderately impressed punctures plus scutellar row. Venter: Me- tepisternum fused posteriorly with metas- ternum; hypomera beside anterior coxal cavities with transverse impression; 4 visi- ble abdominal sterna. Legs: Tibia obliquely and abruptly emarginate, appearing pointed externally. Host plants.—Calathea (Marantaceae) (Jolivet and Hawkeswood 1995). VOLUME 104, NUMBER 3 Described species.—9. Key to species: Group 3 in Spaeth (1938). Range.—Guatemala to Ecuador. Aslamidium (Neoaslamidium Borowiec) (Fig. 9) Aslamidium (Neoaslamidium) Borowiec 1998: 371. Type species: Imatidium for- mosum Spaeth (by original designation). Description.—As in Aslamidium except: Body elongate, parallel-sided. Pronotum: Rectangular. Elytron: Slightly to moderate- ly wider than pronotum; lateral margin im- punctate; lateral and apical margins narrow, in widest part narrower than 2 width of ely- tral disc. Venter: Prosternal process with deep groove along margins. Host plants.—Unknown. Described species.—3. Range.—Costa Rica to Ecuador. Calliaspis Dejean (Fig. 10) Calliaspis Dejean 1837: 391. Type species: Cassida rubra Olivier (by monotypy). Calliapis Boheman 1850: 81 (lapsus cala- mi). Cyanaspis Weise 1904: 433. Type species: Cyanaspis testaceicornis Weise (by mon- otypy). Spaeth 1905 (synonymy). Description.—Convex; rounded. Head: Deeply inserted into pronotum; depressed between eyes; medial sulcus present. Anten- na: 10-segmented; antennomeres VII to X hirsute; I to VI glabrous. Pronotum: At base nearly as wide as base of elytra; sparsely punctate; lateral margin strongly curved and convergent toward apex. Scu- tellum: Broadly triangular. Elytron: Lateral margin laminate, recurved; indented behind humerus; apex conjointly rounded; lateral and apical margins smooth; with 10 rows of punctures plus scutellar row; interspaces wide. Venter: 4 visible abdominal sterna; sternite 4 with medial notch on apical mar- gin; metasternum punctate laterally, smooth in middle. Host plants —Unknown. 731 Described species.—18. Key to species: Wagener (1881) covered 10 species. Range.—French Guyana to Peru. Cephaloleia Chevrolat (Fig. 11) Cephaloleia Chevrolat 1837: 30. Type spe- cies: Hispa nigricornis Fabricius (desig- nated by Staines 1991). Cephalolia Blanchard 1845: 162 nec Weise 1910 (lapsus calami). Uhmannispa Monros and Viana 1947: 172. Type species: Uhmannispa maculata Monros and Viana (by monotypy); Uhmann 1957b (synonymy). Description.—Body elongate, rather par- allel (rarely oval); flat or moderately con- vex. Head: Small; eye finely faceted, slight- ly prominent. Antenna: 11-segmented, as long as head and pronotum combined, fili- form; antennomere I thicker and longer than II; slightly thickened at apex. Prono- tum: Quadrangular, usually wider than long, frequently widest behind anterior angle; weakly convex; lateral margin gradually converge toward base. Scutellum: Short; pentagonal or elongate triangle. Elytron: Varying in form and contour; lateral margin widened behind humerus or humerus insig- nificant; one segment of pygidium exposed; with 10 rows of punctures plus scutellar row. Host plants.—Alpinia, Renealmia (Zin- giberaceae); Heliconia (Heliconiaceae); Calathea, Ischnosiphon, Maranta (Maran- taceae); Cyclanthus (Cyclanthaceae); Cos- tus (Costaceae); Cocos, Diplothemium, Elaeis, Geonoma (Arecaceae); Cordyline (Agavaceae); Panicum, Pharus, Saccharum (Poaceae); Canna (Cannaceae); Musa (Mu- saceae); Ananas (Bromeliaceae); Cypera- ceae (Jolivet and Hawkeswood 1995). Described species.—202 +. Key to spe- cies: Uhmann (1936b) covered 29 species in his collection; Monros and Viana (1947), Argentina; Staines (1996a), Central Amer- ica and the West Indies. Range.—Mexico to Argentina. 732 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cladispa Baly Cladispa Baly 1858: 22. Type species: Cladispa quadrimaculata Baly (by mon- otypy). Description.—Body broadly oval; flat- tened. Head: Short; vertex punctate; frons vertical; eye oval; maxillary palp with pal- pomere 2 almost as long as other three combined; clypeus with margin evenly ar- cuate. Antenna: 11-segmented, % body length; antennomeres II to V wider than I; each antennomere with pointed, oblique an- gle on inner apical margin; antennomere I thicker than II or HI, cylindrical; Hf and Hl compressed laterally; apical 6 antennomeres slender and short, gradually decreasing in thickness. Pronotum: At midline wider than long, convex; disc impunctate; lateral mar- gin parallel for basal %4s, then sharply con- vergent, punctate, and margined laterally; basal margin bisinuate; basal impression present, defined by a transverse sulcus; an- terior margin slightly emarginate. Scutel- lum: Short, almost pentagonal; impunctate. Elytron: Slightly wider than pronotum; with 10 rows of shallow punctures; mar- gined laterally. Venter: Smooth; with 4 vis- ible abdominal sterna. Legs: Tibiae emar- ginate on outer surface, concave enough to cover tarsi; tarsomere 4 not projecting be- yond lobes of 3. Host plant.—Cyrtopodium (Orchidaceae) (Jolivet and Hawkeswood 1995). Described species.—1. Range.—Brazil. Demotispa Baly (Fig. 12) Demotispa Baly 1858: 65. Type species: Demotispa pallida Baly (designated by Monros and Viana 1947). Xenispa Baly 1858: 63. Type species: Xen- ispa pulchella Baly (by monotypy); Wei- se 1910 (synonmy). Demothispa Gemminger and Harold 1876: 3599 (lapsus calami). Pseudimatidium Aslam 1965: 691. Type species: Demotispa pallida Baly (by orig- inal designation); Borowiec 2000 (syn- onymy). Description.—Body elonagte to elon- gate-oval, flattened. Head: Small; frons ver- tical; antennal base with keel or may be broadly separated; interantennal carina dis- tinct; eye slightly protruding; maxillary palp with palpomeres 2 to 4 subequal in length. Antenna: 11-segmented, long, fili- form; antennomeres I—II short, subglobular; III longest, [IV to XI subequal in length. Pronotum: Slightly narrower than base of elytra, transverse; lateral margin curved, narrowly laminate. Scutellum: Pentagonal, cuspidate behind. Elytron: Oblong-oval, wider than pronotum, convex, indented be- hind humerus; lateral margin often expand- ed, smooth; with 10 rows of punctures plus scutellar row. Venter: Prosternum narrow; mesosternum transverse; 4 visible abdomi- nal sterna. Legs: Short; claws divaricate. Host plants.—Unknown. Described species.—48. Key to species: Uhmann (1937a, 1948) covered 23 species; Sanderson (1967), West Indies; group 4 in Spaeth (1938). Range.—Mexico to Argentina. Euxema Baly (192513) Euxema Baly 1885: 3. Type species: Eux- ema insignis Baly (by monotypy). Description.—Body attenuate, moderate- ly convex, gradually widening behind. Head: Medial sulcus absent; vertex im- punctate. Antenna: 11-segmented, filiform, barely % body length; antennomere I thick- er than others; II short; III longest, 3x length of IH; IV to XI uniform in width and length. Pronotum: Transverse, slightly nar- rowed behind, highly convex; apical margin prolonged into lobe which projects over head; margined laterally; basal impression present; lateral impression present in pos- terior angle; impunctate except for impres- sions which have large, dense punctures. Scutellum: Pentagonal. Elytron: Convex; margined laterally; lateral margin smooth; VOLUME 104, NUMBER ee) PAP OD es OF 6 Wf PGROLI LN RS, 1 (ay PRA SS RP PREETI BANA EES NPB ED Ee BINS Figs. 12-20. Dorsal habitus. 12, Demotispa pallida Baly. 13, Euxema insignis Baly. 14, Homalispa sp. 15, Imatidium thoracicum (EF). 16, Melanispa truncata Baly. 17, Octocladiscus fasciatus (Guerin). 18, Parimatidium rubrum (Boheman). 19, Solenispa leptomorpha (Baly). 20, Spaethaspis peruviana Borowiec. 734 apical margin finely dentate; with 10 rows of lightly impressed punctures. Host plants.—Unknown. Described species.—2. Range.—Panama to Venezuela. Homalispa Baly (Fig. 14) Homalispa Baly 1858: 33. Type species: Homalispa batesii Baly (by original des- ignation). Description.—Body oval or oblong-oval, rather flat. Head: Small; eye elongate; frons depressed, obliquely directed forward; la- brum large, visible from above; mandible long, bidentate; labium large, almost as long as wide; clypeus short, with one nar- row cross striation which is ledge-shaped and bent upwards; mouthparts projecting forward. Antenna: 11-segmented, thin, fili- form, inserted in a frontal depression, not separated by a keel; antennomere III not longest. Pronotum: Transverse; lateral mar- gin convergent apically; anterior margin an- gularly pointed; narrower than elytra. Scu- tellum: Pentagonal, cuspidate behind. Ely- tron: Wider than pronotum, indented be- hind humerus; humerus impunctate; lateral and apical margins smooth; with 10 rows of punctures plus scutellar row. Host plants.—Lasiacis, Olyra (Poaceae); Scleria (Cyperaceae) (Jolivet and Hawkes- wood 1995). Described species.—26. Range.—Nicaragua to Argentina. Homalispa (Xanthispa Baly) Homalispa (Xanthispa) Baly 1858: 31. Type species: Cephaloleia cimicoides Guérin-Méneville (by monotypy). Description.—As in Homalispa except: Antennomere III longest; lateral and apical margins of elytra serrate. Host plants—Unknown. Described species.—1. Range.—Brazil. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Imatidium Fabricius (Fig. 15) Imatidium Fabricius 1801: 345. Type spe- cies: Imatidium thoracicum Fabricius (designated by Latreille 1810). Himatidium Latreille 1804: 131 [unjustified emendation]. Himatidella Aslam 1965: 690. Type spe- cies: Imatidium thoracicum Fabricius (by original designation); Borowiec 1984 (synonymy). Description.—Head: Posterior half of frons flattened; eye not projecting; intero- cular width less than width of an eye; dis- tance between antenna not less than 2X dis- tance between antenna and an eye; inter- antennal carina not raised, transverse carina behind labrum absent; labrum distinct. An- tenna: 11-segmented, not thickened apical- ly; antennomere I equal in length to II; more than % length of III; [V to X decreas- ing in length; XI not laterally compressed and shorter than X; IV to XI setose. Pron- otum: Without notch external to anterior se- tae; sparsely punctate; no depressions pre- sent; lateral margin convergent from base to apex. Scutellum: Broadly triangular. El- ytron: Lateral and apical margins laminate; slightly indented behind humerus; with 10 rows of moderately impressed punctures plus scutellar row. Venter: Hypomera with transverse impression beside anterior coxal cavity; 4 visible abdominal sterna. Legs: Tibiae not pointed externally though exca- vated subapically. Host plants.—Areca, Astrocaryum, Are- castrum, Bactris, Cocos, Desmoncus, Elaeis, Geonoma, Licuala, Livistona, Phoe- nix, Pritchardia (Arecaceae); Heliconia (Heliconiaceae) (Jolivet and Hawkeswood EIS). Described species.—14. Key to species: Group 2 in Spaeth (1938). Range.—Mexico to Peru. Melanispa Baly (Fig. 16) Melanispa Baly 1858: 30. Type species: Melanispa truncata Baly (by monotypy). VOLUME 104, NUMBER 3 Description.—Head: Moderately slender, flat; triangular depression present on vertex; punctate between eyes; palpomere 3 of la- bial palp clavate, longer and wider than pre- ceding. Antenna: |1-segmented, slender, fi- liform, % length of body; antennomere I thickened; III longest; I to V shiny, others dull. Pronotum: Transverse, slightly nar- rower than elytra; disc depressed; sparsely punctate; lateral margin convergent from base to apex; margined laterally; basal mar- gin bisinuate. Scutellum: Large, triangular. Elytron: Lateral and apical margins smooth; humerus not produced; parallel-sided; trun- cate at apex; pygidium not exposed; with 10 rows of lightly impressed punctures plus scutellar row. Venter: Pro-, meso-, and me- tasterna smooth medially, punctate lateral- ly; with 5 visible abdominal sterna. Host plant.—Unknown. Described species.—1. Range.—Guadeloupe. Octocladiscus Thomson (Fig. 17) Octocladiscus Thomson 1856: 480. Re- placement name for Cladophora Guérin- Méneville 1844: 284 [name preoccupied by Cladophora Guérin-Méneville (Co- leoptea: Lycidae)]. Type species: Clado- Phora fasciatus Guérin-Méneville (by monotypy). Description.—Body broadly oval, flat- tened. Head: Vertex punctate; eye finely faceted. Antenna: 11-segmented, % length of body; antennomeres II and III short, fol- lowing antennomeres longer; male pecti- nate from antennomeres IV to X; female filiform, with projection on antennomeres IV to X. Pronotum: At midline 2 as wide as long; lateral marign strongly rounded, narrowing anteriorly; disc impunctate; large shallow punctures present laterally and ba- sally. Scutellum: Elongate triangle. Elytron: Oblong-oval, wider than pronotum; mar- gined laterally; pygidium visible from above; with 10 rows of punctures plus scu- 7135 tellar row. Venter: 5 visible abdominal ster- na. Host plant.—Unknown. Described species.—1. Range.—French Guyana. Parimatidium Spaeth (Fig. 18) Parimatidium Spaeth 1938: 307. Type spe- cies: Imatidium rubrum Boheman (by original designation). Description.—Head: Posterior frons flat- tened and anterior part depressed; intero- cular space less than the width of an eye; eye large; interantennal carina and trans- verse carina behind labrum absent; labrum distinct; maxillary palp with palpomere 4 longer than 3. Antenna: 11-segmented, not thickened apically; antennomere III longer than II; XI longer than X. Pronotum: Emar- gination on anterior margin shallow, with- out notch external to anterior seta; lateral margin strongly convergent from base to apex; lateral impressions absent; posterior angle sharp and acute. Scutellum: Broadly triangular. Elytron: Lateral margin lami- nate, dentate externally toward apex, slight- ly indented behind humerus; with 10 rows of punctures plus scutellar row. Venter: Metepisternum distinct posteriorly; hypom- era with transverse impression external to anterior coxal cavities. Legs: Tibiae not pointed externally, excavated subapically. Host plant.—Elaeis (Arecaceae) (Jolivet and Hawkeswood 1995). Described species.—5. Key to species: Spaeth (1938) treated as a subgenus of /m- atidium. Range.—Venezuela to Paraguay. Pseudostilpnaspis Boroweic Pseudostilpnaspis Boroweic 2000: 162. Type species: Stilpnaspis columbica Wei- se (by original designation). Description.—Body elongate, almost parallel-sided or slightly narrowed posteri- orly. Head: Interantennal area narrow (not wider than first antennal antennomere), flat 736 or slightly convex, not carinate. Antenna: 11-segmented; antennomeres I-—II short, subglobular; III distinctly longer that II; I- II sparsely pubescent and glabrous, III—XI densely pubescent, dull. Pronotum: Widest at base, slightly narrower than base of ely- tra; sides on basal % parallel or slightly con- verging anteriorly; explanate sides very narrow, not wider than width of antennom- eres. Elytron: With 10 rows of lightly im- pressed punctures plus short scutellar row; explanate margin narrow, at widest not wid- er than two elytral intervals combined. Host plant.—Unknown. Described species.—3. Range.—Costa Rica to Colombia. Solenispa Weise (Fig. 19) Solenispa Weise 1905a: 53. Type species: Solenispa impressicollis Weise (designat- ed by Uhmann 1957b). Description.—Body long, narrow. Head: Vertex deeply concave, descending to mar- gin of eye; eye protruding; clypeus with medial carina extending between antennal bases. Antenna: 11-segmented; antennom- eres I, III, and XI longer than others. Pron- otum: Longer than wide, completely mar- gined, convex; parallel-sided; basal impres- sion present; punctate flattened area present near midpoint on each side. Scutellum: Elongate, triangular. E/ytron: Lateral and apical margins smooth; apex conjointly rounded; basal margin carinate to humerus; with 10 rows of shallow punctures plus scu- tellar row. Venter: Epipleura visible from oblique angle; pro- and mesocoxae large, globular; prosternum narrow; 5 visible ab- dominal sterna, first two punctate laterally, last three hirsute. Host plants —Unknown. Described species.—9. Range.—Costa Rica to Ecuador. Spaethaspis Hincks (Fig. 20) Spaethaspis Hincks 1952: 343. Type spe- cies: Spaethaspis lloydi Hincks (by mon- otypy). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Description.—Body elongate; pronotum rectangular with sides almost parallel. Head: Vertex finely and sparsely punctate, deeply impressed; interocular space broad, flat; interantennal carina absent; clypeus very short, nearly 4X as wide as long; max- illary palp with palpomere 4 subequal in length to 3. Antenna: 11-segmented; basal two antennomeres glabrous; remaining an- tennomeres feebly pubescent; antennomere Il % length of I or HI; ['V subequal in length to III. Pronotum: Transverse; anterior mar- gin deeply emarginate in middle; lateral margin straight for basal %s, then slightly converging towards apex; anterior angle broadly rounded; basal margin prominent and more or less truncate adjacent to scu- tellum, slightly oblique on either side to rectangular posterior angle; posterior angle with shallow depression; sparsely, finely punctate on disc, strongly, distinctly punc- tate at sides. Scutellum: Broadly triangular. Elytron: Moderately wide, almost 2X as long as wide; transverse impression behind humerus; lateral margin expanded, some- what concave with polished margin, slight- ly widened behind, broadly rounded at apex; with 10 rows of fine punctures plus scutellar row; striae wide. Venter: Proster- num broad, strongly expanded behind with deep longitudinal striae, middle stria lon- gest and widest; abdominal sternites flat, very finely punctate, apical segment more or less truncate, without apical emargina- tion. Legs: Tarsal claws with large basal tooth. Host plants.—Unknown. Described species.—2. Range.—Ecuador and Peru. Stenispa Baly (Fig. 21) Stenispa Baly 1858: 13. Type species: His- pa metallica Fabricius (by original des- ignation). Description.—Body cylindrical or flat- tened; parallel-sided or narrowing posteri- orly. Head: Small, rounded; frons concave; VOLUME 104, NUMBER 3 maxillary palp with palpomere | small, 2 and 3 obconical, subequal in length and width, 4 oval, acuminate; antenna inserted close to eye in shallow pits, divided by lon- gitudinal keel; eye oval, finely faceted, slightly prominent. Antenna: | 1-segmented; antennomeres [| to II subglobose; HI to X cylindrical, II longer than IV; XI subacute, slightly dilated toward apex. Pronotum: Quadrate; slightly narrower than elytra; an- terior margin rounded; lateral margin par- allel. Scutellum: Pentagonal or subcylindr- ical. Elytron: Elongate; slightly convex; narrowing apically; lateral and apical mar- gins finely dentate; with 10 rows of punc- tures plus scutellar row. Legs: Short; ro- bust; femora stout; tibiae not arcuate, slight- ly flattened; tarsomeres dilated, densely pu- bescent beneath, tarsomere 4 with nearly half its length projecting beyond 3; claws stout, moderately arcuate, divaricate. Ven- ter: Metepisternum very narrow at middle; prosternum narrow, longitudinally fur- rowed; abdominal sternites | and 2 with su- ture obsolete in middle. Host plants.—Carex, Cyperus, Scirpus (Cyperaceae); Paspalum, Spartina (Po- aceae); Juncaceae (Jolivet and Hawkes- wood 1995). Described species.—21. Key to species: Horn (1883), North America; Monros and Viana (1947), Argentina. Range.—United States to Argentina. Stilpnaspis Weise (Fig. 22) Stilpnaspis Weise 1905b: 298. Type spe- cies: Stilpnaspis marginata Weise (by monotypy). Rhodimatidium Aslam 1965: 690. Type species: Himatidium coccinatum Bohe- man (by original designation); Borowiec 2000 (synonymy). Description.—Body oval to broadly oval; pronotum semicircular in outline, widest just before base, broadly explanate; explan- ate margin of elytra broad, at widest dis- tinctly wider than elytral intervals 9 and 10 qT3T combined. Head: Posterior % of frons flat- tened, eye level with it; interocular space less than width of eye; distance between an- tennal insertion more than 2 the distance between antenna and eye; interantennal ca- rina absent, area between antennal bases broad and flat; labrum long, somewhat nar- 2 rowed, thickened at sides, without posterior transverse carina; maxillary palp with pal- pomere 4 longer than 3. Antenna: 1|1-seg- mented, antennomeres narrow, elongate; antennomere I thickest, robust, longer than wide; II longer than I; III longer than II. Pronotum: Transverse, convex, narrower at base than elytra at humeri; with emargina- tion external to anterior seta; lateral margin convergent from base to apex, slightly re- curved; without basal depressions; disc im- punctate; sparsely punctate laterally. Scu- tellum: Broadly triangular. Elytron: Mod- erately convex; at base as wide as base of pronotum, slightly indented behind humer- us, slightly widened before middle; lateral margin laminate, smooth; with 1O rows of lightly impressed punctures plus scutellar row; striae wide. Venter: Prosternum with transverse impression beside anterior coxal cavity; 4 visible abdominal sterna. Legs: Tibiae excavated subapically, not pointed externally. Host plants.—Unknown. Described species.—17. Key to species: Group | in Spaeth (1938). Range.—Costa Rica to Argentina. TRIBE DELOCRANIINI SPAETH 1929: 113 Hoplhionotites Chapuis 1875: 357 (in part). Type genus.—Delocrania Guérin-Méne- ville. Description.—Antenna: Very long; api- cal antennomeres wider than long. Prono- tum: Without anterior angle. Venter: Pros- ternum moderately wide between procoxae; prosternal process wide, truncate apically; mesosternum truncate in front, touching prosternum, not margined behind. 738 Delocrania Guérin-Méneville (Fig. 23) Delocrania Guérin-Méneville 1844: 141. Type species: Delocrania cossyphoides Guérin-Méneville (by monotypy). Description.—Head: Barely visible dor- sally; labrum transverse, subemarginate on apical margin, small oblique carina in mid- dle; eye oval. Antenna: Placed on two tu- bercles; 11-segmented, filiform, extends just beyond base of pronotum; antennomere I thick; II cylindrical, shorter than I; re- mainder thinner, subequal in length. Pron- otum: Wider than long, convex; anterior margin deeply emarginate behind head, emargination quadrate; anterior angle di- late-foliate, rounded; lateral margin slightly curved, sinuate at base; depressed laterally; posterior angle rectangular. Scutellum: Quadrate, rounded apically. Elytron: At base as wide as base of pronotum, convex; lateral margin dilated, flat; exterior apical angle rounded; with 10 rows of regular punctures, plus scutellar row; one costa pre- sent. Legs: Short; tibiae obliquely notched apically on external margin; tarsi wide, first three tarsomeres equal in width, tarsomere 4 as long as lobes of 3; claws divaricate. Host plants.—Attalea, Cocos, Diploth- emium, Elaeis (Arecaceae) (Jolivet and Hawkeswood 1995). Described species.—3. Range.—Costa Rica to Ecuador. TRIBE HEMISPHAEROTINI MONROS AND VIANA 1951: 368 Himatidiites Chapuis 1875: 361 (in part). Porphyraspitae Spaeth 1929: 113. Type genus.—Hemisphaerota Chevrolat. Description.—Small species (less than 10 mm); more or less oblong or oval, convex. Head: Partially visible dorsally. Antenna: 11-segmented, short, not extending beyond base of pronotum; apical antennomeres shorter than wide, thickened. Pronotum: Anterior angle prominent, with seta. Ely- tron: With lateral margin expanded. Venter: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Prosternum short, wide between procoxae, not widened behind, truncate on anterior margin; mesosternum notched in front, de- pressed, posterior margin doubled; abdom- inal sternites | and 2 not fused. Legs: Short; robust; tibiae dilated; tarsi very wide. KEY TO GENERA 1. Apical tarsomere with two claws; epipleural keel present Spaethiella — Apical tarsomere with one claw (interior claw degenerate); epipleural keel absent Hemisphaerota Hemisphaerota Chevrolat (Fig. 24) Hemisphaerota Chevrolat 1837: 391. Type species: Cassida erythrocera Germar (= Imatidium cyanea Say) (by monotypy). Emperochela Spaeth 1901: 333. Type spe- cies: Porphyraspis palmarum Boheman (by original designation); Spaeth 1905 (synonymy). Porphyraspis Hope 1840: 152. Type spe- cies: Cassida erythrocera Germar (by original designation); Guérin-Méneville 1844 (synonymy). Description.—Body more or less hemi- spherical; elytra not strongly sculptured. Head: Visible dorsally; medial sulcus pre- sent; vertex punctate; clypeus transverse. Antenna: 11-segmented; antennomere I long, thick; II shorter than I, subequal in width; HI—VII short, transverse; VIII—XI wider than preceding. Pronotum: Trans- verse; convex; anterior margin emarginate behind head; lateral margin curved; poste- rior angle rounded; depression present in anterior angle. Scutellum: Trapezoidal; acute at apex. Elytron: Convex, at base lit- tle wider than base of pronotum; lateral margin curved; apex conjointly rounded; epipluron without transverse carina; exca- vation behind epipluron for apex of femur; with 10 rows of regular punctures plus scu- tellar row. Venter: Prosternum wide, ante- rior margin curved, posterior margin with deep metasternal notch; metasternum large, posterior margin almost straight, anterior VOLUME 104, NUMBER 3 739 4 ces S$ ‘_ A ROE el] | | -— Ss rw ~ 4 & Bo R 3 Us ce & S | § % & a Ed Ps = 24 = ra zg 2 e : a § Es iS 2 é 2 3 3 | § : 2 « 21 ~~ - mm ee RET OED OP) Miah Ss Z> — , TS DRA A Fa ae ER RS Figs. 21-29. Dorsal habitus. 21, Stenispa metallica (EF). 22, Stilnpaspis rubiginosus (Boheman). 23, Delo- crania panamensis Champion. 24, Hemisphaerota cyanea (Say). 25, Spaethiella circumdata (Boheman). 26, Hybosispa melanura Weise. 27, Arescus labiatus Perty prosternum. 28, Arescus labiatus. 29, Chelobasis bicolor Gray, prosternum. 740 margin with notch for prosternal process. Legs: Short, robust; femora thick, triangular in cross-section; tibiae shorter than femora, triangular in cross-section, excavation on outer surface for reception of tarsi; tarso- meres very wide, flat, tarsomere | triangu- lar, 2 larger than 1, transverse, bilobed, 3 deeply bilobed, larger than 2, 4 slightly ex- ceeds lobes of 3; internal claws degenerate. Host plants.—Acoelorrhaphe, Arecas- trum, Attalea, Chamaerops, Cocos, Sabal, Serenoa, Thrinax, Washingtonia (Areca- ceae) (Jolivet and Hawkeswood 1995). Described species.—9. Key to species: Wagener (1881). Range.—Southern United States to Bra- VAN Spaethiella Barber and Bridwell (Fig. 25) Spaethiella Barber and Bridwell 1940: 11. Replacement name for Hemisphaerota: Spaeth 1905 not Chevrolat 1837. Type species: Imatidium sanguineum Fabricius (by original designation). Description.—Body more or less hemi- spherical; elytra strongly sculptured. Head: Visible dorsally; medial sulcus present; ver- tex irregularly punctate; clypeus transverse, depressed. Antenna: 11-segmented; anten- nomere I long, thick; I] subequal in length and width to I; III % length of II; VII-XI thicker than preceding. Pronotum: Trans- verse, convex, circular depression present on each side of disc; anterior margin emar- ginate behind head, with a pair of setae lo- cated on small promience on either side of head; lateral margin curved, expanded; pos- terior angle rounded. Scutellum: Trapezoi- dal; subtruncate at apex. Elytron: Convex, at base little wider than base of pronotum; lateral margin more or less parallel or slightly curved; apex conjointly rounded; epiplurae with transverse carina which is obsolete before margin; excavation behind for apex of femur; with 10 rows of regular punctures plus scutellar row. Venter: Pros- ternum wide, anterior margin curved, pos- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON terior margin with pointed projection; me- tasternum large, posterior margin almost straight, anterior margin with notch for prosternal process; suture separating ab- dominal sterna | and 2 distinct. Legs: Short, robust; femora thick, triangular in cross- section; tibiae shorter than femora, trian- gular in cross-section, excavation on outer surface for reception of tarsi; tarsi very wide, flat, tarsomere | triangular, 2 larger than 1, transverse, bilobed, 3 deeply bi- lobed, larger than 2, 4 slightly exceeds lobes of 3; claws short, equal, with small rectangular lamina between. Host plants.—Arecastrum, Astrocaryum, Bactris, Cocos, Copernicia, Elaeis, Phoe- nix, Pritchardia, Washingtonia (Areca- ceae); Heliconia (Heliconiceae) (Jolivet and Hawkeswood 1995). Described species.—33. Key to species: Wagener (1881). Range.—Guatemala to Peru. TRIBE HYBOSISPINI WEISE 1910: 69 Type genus.—Hybosispa Weise. Description.—Head: Frons with excava- tion divided by longitudinal keel from an- tennal base to clypeal margin; clypeus tri- angular; maxillary palp with palpomeres 2— 3 short, 4 oval, as long as 2—3 combined; antenna inserted into pits at side of eye. An- tenna: 11-segmented; filiform. Pronotum: Without setae in any angle. Elytron: Trun- cate posteriorly, part of pygidium exposed; 10 rows of punctures plus scutellar row. Hybosispa Weise (Fig. 26) Hybosispa Weise 1910: 96. Type species: Hybosispa melanura Weise (by monoty- Py). Description.—Body subcylindrical, at- tenuate, slightly convex. Head: Moderately large; eye large, wider above than below, finely faceted; frons with triangular exca- vation divided by a longitudinal keel from antennal base to clypeal margin; clypeus triangular; maxillary palp with palpomeres VOLUME 104, NUMBER 3 2 and 3 short, 4 oval, as long as 2 and 3 combined; antenna inserted into pit at side of eye. Pronotum: Quadrate, %4 wider than long; parallel-sided; finely margined on an- terior margin; depression present on either side of middle near side margin; seta absent in all angles. Scutellum: As long as wide; pentagonal; cuspidate behind. Elytron: Wider than pronotum at humeri, gradually narrowing apically, truncate posteriorly, part of pygidium exposed; with 10 rows of punctures plus scutellar row. Venter: Me- sosternum more elevated than prosternum. Host plants.—Unknown. Described species.—S. Range.—Brazil to Bolivia. TRIBE ARESCINI CHAPUIS 1875: 298 Type genus.—Arescus Perty. Description.—Body attenuate, moderate- ly convex; color red or yellow with black or blackish-blue markings. Head: Short; frontal horn usually present. Antenna: In- serted near clypeus, longer than head and pronotum combined; | 1-segmented; anten- nomere I robust; II small; HI attenuate, nar- row; rest shorter and slightly thicker. Pron- otum: Quadrate, wider than long; almost parallel-sided, converging suddenly at apex, slightly sinuate behind middle. Scutellum: Triangular; longer than wide. Elytron: Wid- er than pronotum at humeri; lateral margin subparallel; apex conjointly rounded; with 10 rows of punctures plus scutellar row, punctures often confused at sides and be- yond middle. Venter: Pro- and mesosterna narrow; pro- and mesocoxae large, globu- lar; suture between abdominal sternites 1 and 2 obscure in middle. Sexual dimor- phism apparent in some genera. KEY TO GENERA 1. Prosternum not produced between mesocox- Est (Sa OAT) carats ay cine seein a ORO ee ed oes 2 = Prosternum produced between mesocoxae (R29) BPE areas AIA ae Se gg Ae: 3 2(1). Pronotum with seta present in anterior an- gle; first antennomere simple, without pro- longation; frontal horn long, cuspidate; sex- es different Xenarescus 741 Pronotum with seta present in all angles; first antennomere with spoon-shaped_pro- longation; frontal horn short, broadly trun- cate at apex-sexesisimilanerey et 0,2... Arescus 3(1). Pronotum with seta present in anterior an- gle; mesocoxae separated by diameter of a coxa (Fig. 30) Chelobasis ~ Pronotum with seta present in all four an- gles; mesocoxae separated by less than di- ameter of a coxa (Fig. 32) .... Nympharescus Arescus Perty (Figs. 27—28) Arescus Perty 1834: 100. Type species: Arescus labiatus Perty (by monotypy). Description.—Body elongate, parallel; color usually yellow or red with variable black or bluish-black markings, markings may be absent on some specimens. Head: Frontal horn variable in length, truncate in front, female process a short triangle in some species; vertex depressed between eyes. Antenna: Antennomere I lengthened into an ventral process as long as anten- nomeres II and III combined. Pronotum: Quadrangular; basal margin bisinuate, deeply lobed; seta present in all four angles. Scutellum: Elongate, triangular. Elytron: Similar in both sexes; male without apical appendage; three species with longitudinal carina behind middle. Venter: Mesocoxae almost touching; prosternal process not pro- jecting between mesocoxae. Host plants.—Heliconia (Heliconiaceae) (Jolivet and Hawkeswood 1995). Described species.—6. Range.—Colombia to Peru. Chelobasis Gray (Figs. 29-31) Chelobasis Gray 1832: 143. Type species: Chelobasis bicolor Gray (by monotypy). Description.—Head: Frontal horn trun- cate at apex; vertex impunctate, depressed between eyes. Antenna: Antennomere I lengthened into an ventral process as long as antennomeres II and HI combined. Pron- otum: Quadrate; anterior margin bisinuate, not deeply bilobed; seta present in anterior 742 angle; punctate laterally and basally. Scu- tellum: Elongate, triangular. Elytron: Mar- gined laterally; apex conjointly rounded; with 10 rows of punctures; non-costate; male without apical appendage. Venter: Mesocoxae separated by the diameter of a coxa; prosternal process projecting between mesocoxae; pro- and mesotibiae expanded at apex. Host plants.—Heliconia (Heliconiaceae); Calathea, Ischnosiphon (Marantaceae) (Jo- livet and Hawkeswood 1995). Described species.—4. Range.—Guatemala to Peru. Nympharescus Weise (Figs. 32-33) Nympharescus Weise 1905a: 320. Type species: Arescus separatus Baly (desig- nated by Uhmann 1957b). Description.—Head: Frontal horn trun- cate at apex; vertex convex between eyes, impunctate. Antenna: Antennomere I with ventral process as long as II; HI longest. Pronotum: Quadrate, flattened; parallel-sid- ed; seta present in all four angles; basal margin bisinuate; tooth present in anterior angle; posterior angle acute. Scutellum: Elongate, triangular. Elytron: Lateral mar- gin with smooth raised edge or rim; tooth present in sutural angle; with 10 rows of punctures; no costae. Venter: Mesocoxae almost touching; prosternal process not pro- jecting between mesocoxae. Legs: Exterior apical edges of tibiae expanded backwards. Host plants.—Heliconia (Heliconiaceae) (Jolivet and Hawkeswood 1995). Described species.—6. Range.—Colombia to Peru. Xenarescus Weise (Fig. 34) Xenarescus Weise 1905a: 320. Type spe- cies: Hispa monoceros Olivier (by mon- otypy). Description.—Body elongate, parallel- sided. Head: Vertex concave; frontal horn long, cuspidate, curved; area behind horn PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON depressed. Antenna: Antennomere I simple in female or with a pointed ventral process. Pronotum: Quadrangular, longer than wide, weakly convex; punctured pit before middle on each side (deepens in male); seta present in anterior angle; posterior margin with transverse sulcus; female with disc almost symmetrical. Scutellum: Triangular. Ely- tron: Costae united, first and second sepa- rate behind middle; male with tooth-like tu- bercle and carina at humerus; apex emar- ginate at suture, male with spoon-like ap- pendage on exterior apical angle. Venter: Prosternum forms small carina sloping back to mesocoxae. Host plants.—Heliconia (Heliconiaceae); Stromanthe (Marantaceae); Musaceae (Jo- livet and Hawkeswood 1995). Described species.—1, with 16 subspe- cies. Key to subspecies: Pic (1927a). Range.—Colombia and Venezuela. TRIBE ALURINI CHAPUIS 1875: 292 Sphaeropalpites Chapuis 1875: 359. Type genus.—A/urnus Fabricius. Description.—Body large, elongate, el- liptical, subparallel, flattened. Antenna: 11- segmented, not reaching middle of elytra. Pronotum: Tufts of setae present in each angle. Elytron: Punctation disordered, not forming rows; costae absent; apex rounded. Remarks.—This tribe contains the largest species of hispines. KEY TO GENERA Ne Pronotum nearly as wide as base of elytra (Big: 3S) Rec cs se a Ree 2 - Pronotum much narrower than base of elytra (Figs. 35-37, 39) Py Bodyralmostqoyalieeser sae > nae Adalurnus Body elongate Platyauchenia . Elytra subcordiform (Fig. 39); antennomere III as long as IV to VI combined ..... er ao 515. che Biden eee Pseudocalaspidea - Elytra subparallel (Figs. 35-37); antennom- ere III not as long as IV to VI combined 4 . Antenna filiform; antennomere III as long as IV to V combined; epipleura of elytra pos- teriorly setose; labrum anteriorly impressed SA Ea RAE Se noe ere, cieobeur: Alurnus - Antenna subserrate; antennomere III not as VOLUME 104, NUMBER 3 743 Figs. 30-38. Dorsal habitus. 30, Chelobasis bicolor mesocoxae. 31, Chelobasis bicolor. 32, Nympharescus separatus (Baly), mesocoxae. 33, Nympharescus separatus. 34, Xenarescus monoceros (Olivier). 35, Alurnus salvini Baly. 36, Coraliomela brunnea (Thunberg). 37, Mecistomela marginata (Thunberg). 38, Platyaucheria latreilleri (Castelnau). 744 long as IV to V combined; epipleura of el- ytra not setose; labrum not impressed .... 5 5(4). Antennomere IV distinctly shorter than III; elytra with fine punctures densely grouped Eis aie, te Eas EERE) Cee ee ee caer Coraliomela ~ Antennomere IV almost as long as III; punc- tures of elytra large, rugose .... Mecistomela Adalurnus Maulik Adalurnus Maulik 1936: 392. Type species: Adalurnus rotundatus Maulik (by mono- typy). Description.—Body broad, nearly as wide as long, convex, in side view, highest point just behind middle. Head: Concave between eyes; mandible large, convex. Pronotum: Almost as wide as base of ely- tra, more shiny than elytra; finely and sparsely punctate. Scutellum: Pentagonal; smooth. Elytron: Smooth, duller than pron- otum; confusedly and moderately punctate, punctures closely spaced; lateral and apical margins smooth, expanded. Venter: Almost glabrous; shining; prosternum with front convex, expanded to conceal lower mouth- parts; 5 visible abdominal sterna. Legs: Fairly robust; sparsely covered with fine hairs; generally not visible from above; femora widest in middle; tibiae wider at apex, setose at apex; tarsi large, broad, thick setae beneath; tarsomeres | small; 2 bi- lobed; 3 longer than 2; 4 long but not pro- jecting past 3; claws simple, strong. Host plant.—Unknown. Described species.—1. Range.—Brazil. Alurnus Fabricius (Fig. 35) Alurnus Fabricius 1775: 94. Type species: Alurnus grossus Fabricius (by monoty- py). Poecilalurnis Jacobson 1899: 247. Type species: Alurnus bipunctatus Olivier (by original designation); Fischer 1935 (syn- onymy). Description.—Head: Frons slightly con- cave. Antenna: Filiform, extending to hu- meri; antennomere I short, thick; IT smaller, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON wider than long; III as long as IV to V com- bined; VI to XI decreasing in length, thin- ner than preceding. Pronotum: Highly con- vex, more so anteriorly; wider than long, narrowing posteriorly; glabrous or finely pubescent; margined or not. Scutellum: Cordate or triangular. Elytron: Oblong-oval, convex; margined; sparsely pubescent. Ven- ter: Pubescent. Host plants.—Heliconia (Heliconiaceae); Cocos, Elaeis (Arecaceae) (Jolivet and Hawkeswood 1995). Described species.—21. Key to species: Fischer d935): Range.—Costa Rica to Ecuador. Coraliomela Jacobson (Fig. 36) Coraliomela Jacobson 1899: 253. Type spe- cies: Alurnus brunnea Thunberg (= AI- urnus corallina Vigors) (designated by Fischer 1935). Psilurnus Weise 1900: 218. Type species: Psilurnus consanguineus Weise (by orig- inal designation); Fischer 1935 (synony- my). Silurnus Weise 1900: 218 [lapsus calami]. Description.—Body large, elongate el- lipse; color generally red or black; similar in appearance to Mecistomela. Head: Flat or slightly convex between eyes; frons with small depressions on anterior margin; inter- antennal keel absent; sulci near eye faint; punctation dense and irregular. Antenna: Short, barely reaching base of pronotum; antennomere III not as long as IV to V combined; IV shorter than HI. Pronotum: Less transverse than in Mecistomela; oth- erwise similar. Scutellum: Triangular. Ely- tron: Wider than pronotum; punctation im- pressed; vestige of irregular costae visible; margined laterally. Venter: Rugose. Host plants.—Copernicia, Cocos, Di- plothemium, Elaeis, Orbignya, Phoenix (Arecaceae) (Jolivet and Hawkeswood 1995): Described species.—4. Key to species: Fischer (1935). VOLUME 104, NUMBER 3 Range.—Brazil to Argentina. Mecistomela Jacobson (Fig. 37) Mecistomela Jacobson 1899: 246. Type species: Alurnus marginata Thunberg (by monotypy). Description.—Body large, elongate el- lipse. Head: Flat or slightly convex be- tween eyes; frons with small depressions on anterior margin; interantennal keel absent; sulci near eye faint; punctation dense and irregular. Antenna: Antennomeres short, sparsely pubescent, each antennomere with ventral asymmetrical apical expansion which diminishes in size from base to apex; antennomere IV almost as long as III. Pron- otum: Transverse, slightly narrowing ante- riorly; lateral margins almost parallel: densely punctate; slight basal impression present on each side of middle. Scutellum: Triangular. Elytron: Wider than pronotum; finely, densely punctate; sides margined. Legs: Trochanters with tufts of setae. Pron- otum and elytra with orange markings. Host plants.—Alagoptera, Copernicia, Cocos, Diplothemium, Elaeis, Lantania, Livistona, Phoenix (Arecaceae) (Macedo et al. 1994). Described species.—1. Key to subspe- cies: Fischer (1935). Range.—Brazil to Argentina. Platyauchenia Sturm (Fig. 38) Platyauchenia Sturm 1843: 358. Type spe- cies: Cassida latreilleri Castelnau (by monotypy). Description.—Head: Clypeus transverse, deeply sulcate; maxillary palp with 4 pal- pomeres, palpomeres | to 3 short, equal in length and width, cylindrical, 4 large, rounded, truncate. Antenna: Filiform, apical antennomere widest. Pronotum: Nearly as wide as base of elytra, widest at base, nar- rowing anteriorly; lateral margin evenly ar- cuate from base to apex; disc highly con- vex; depressed laterally on each side of 745 disc; basal and anterior margins bisinuate. Scutellum: Triangular, rounded at apex. El- ytron: Convex; punctate; widest in middle, narrowing anteriorly and posteriorly; mar- gined laterally; basal and sutural margins yellowish. Host plants.—Cocos, Diplothemium (Arecaceae) (Jolivet and Hawkeswood 1995S). Described species.—2. Range.—Brazil. Pseudocalaspidea Jacobson (Fig:.39) Pseudocalaspidea Jacobson 1899: 245. Type species: Alurnus cassidea West- wood (by monotypy). Description.—Head: Pubescent; medial sulcus present; frons punctate. Antenna: Antennomere III as long as IV to VI com- bined. Pronotum: Much narrower than el- ytra, convex; widest at base, narrowing an- teriorly; pubescent, more dense on anterior and lateral margins; finely punctate; ante- rior angle rounded; basal margin bisinuate. Scutellum: Elongate, triangular. Elytron: Subcordiform, posteriorly straightened, highly convex; humeral angle produced; laminate laterally. Venter: Pro-, meso-, and metasterna with setae; abdominal sterna pu- bescent; sterna 2 to 4 with pale pustule lat- erally; sternum 5 with sides and apex pale and pustulate. Host plant.—Unknown. Described species.—1. Key to subspe- cies: Fischer @1935)- Range.—Brazil and Ecuador. TRIBE PROSOPODONTINI WEISE 1910: 69 Type genus.—Prosopodonta Baly. Description.—Head: Nearly attenuate, almost cylindrical, egg-shaped, small; clyp- eus prolonged to a ridge between the anten- nal bases. Antenna: 11-segmented, filiform, antennomere III longest; apical 5 anten- nomeres slightly thickened. Pronotum: Se- tae present in posterior angle. Scutellum: Longer than wide; rounded behind. Elytron: 746 Nearly “4 wider than pronotum; 10 rows of punctures plus scutellar row. Prosopodonta Baly (Fig. 40) Prosopodonta Baly 1858: 68. Type species: Prosopodonta limbata Baly (by original designation). Cheirispa Baly 1858: 71. Type species: Cheirispa suturalis Baly (by original des- ignation); Weise 1910 (synonymy). Chirispa Gemminger and Harold 3604 [lapsus calami\. 1876: Description.—Head: Nearly attenuate, almost cylindrical, egg-shaped, small; me- daal sulcus absent; clypeus prolonged into a carina between antennal bases; antenna separated by a sharp keel. Antenna: 1 1-seg- mented, filiform; antennomeres I and II subequal in length, thick; HI longest, cylin- drical, as long as I and II combined; IV to VI shorter, oblong, decreasing in length; VII to XI short, slightly dilated; 5 apical antennomeres slightly thickened. Prono- tum: Transverse; margined laterally and ba- sally; scattered coarse punctures posteriorly on lateral margin; seta present in posterior angle. Scutellum: Cuspidate; longer than wide; rounded at apex. Elytron: Oblong; nearly %4 wider than pronotum; margined laterally; apex rounded; punctures little im- pressed, no costae; with 10 rows of punc- tures plus scutellar row. Legs: Profemur toothed or not. Host plants.—Heliconia (Heliconiaceae); Arecaceae (Jolivet and Hawkeswood 1995). Described species.—26. Range.—Nicaragua to Ecuador. TRIBE SCELOENOPLINI UHMANN 1930a: 238 Cephalodontites Chapuis 1875: 313 (in part). Cephalodontini Weise 1910: 69. Type genus.—Sceloenopla Chevrolat. Description.—Antenna: Antennomeres Il to VI long, cylindrical or laterally com- pressed; apical 4 differing from the others (either longer or shorter, thinner or thicker, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cylindrical, very pubescent, or a different color). Pronotum: A seta present in anterior angle. Scutellum: Longer than wide; round- ed at apex. Elytron: With 10 rows of punc- tures plus scutellar row; humeral rows may have 1 or 2 secondary rows; secondary rows also appear between other rows. KEY TO GENERA Ite Body more or less flattened (Figs. 42—44): legs slender, the anterior pair longer than rest, frequently armed with a tooth in male — Body moderately convex (Figs. 41, 45); legs in both sexes simple, short and strong ... 2 2(1). Lateral margin of elytra rounded (Fig. 45) RS eeins: sls 8 chavs este op Pent Serratispa ~ Lateral margin of elytra parallel (Fig. 41) Ree eos. ios Zt. Ee ee Acentroptera 3(1). Apical 4 antennomeres shorter than preced- NDE Ab. oe ERENCE Co a Gl oie acre Ocnosispa = Apical 4 antennomeres longer than preced- UN OMe cho, Fact, 01/5. + 15. 8 dy egigs Sen eel 6 Ree 4 4(3). Sides of elytron not almost vertical from Costas: (Bie: 744). SAMS Sceloenopla ~ Sides of elytron almost vertical from costa 3) ((Ev 342) ee een ete neo. c c Pseudispa Acentroptera Baly (Fig. 41) Acentroptera Baly 1858: 121. Type species: Hispa_ pulchella Guérin-Méneville (by original designation). Description.—Body elongate, slightly convex. Head: Vertex depressed; medial sulcus present. Antenna: 11-segmented; an- tennomere I transverse; II subcylindrical, as long as I; II] to IV cylindrical, subequal in length, each longer than II; V subcylindri- cal; VI to X transverse; XI pointed at apex; VII to XI hirsute. Pronotum: Longer than wide, widest at base, convex; lateral margin bisinuate; basal impression present; tooth present in anterior angle. Scutellum: Quad- rate. Elytron: Humerus slightly produced; margined laterally; apex with slight lamina; with 10 rows of punctures plus scutellar row; 4 well produced costae, costa 4 may be obsolete in middle. Venter: With 5 vis- ible abdominal sterna. Legs: Short robust; anterior pair not prolonged. VOLUME 104, NUMBER eS) = TAT : : ; : 4 >» & x \E 3 ie \ : ~ & ; — ee. & “le 5 : a 45 a Figs. 39-47. Dorsal habitus. 39, Pseudocalaspidea cassidea (Westwood). 40, Prospodonata distincta Baly. 41, Acentroptera sp. 42, Ocnosispa pectoralis Uhmann. 43, Pseudispa sp. 44, Sceloenopla carinata (F.). 45, Serratispa quadricostata Staines. 46, Serratispa quadricostata head. 47, Acanthodes donckieri Weise. 748 Host plant.—Ananas (Bromeliaceae) (Jo- livet and Hawkeswood 1995). Described species.—13. Range.—Costa Rica to Argentina. Ocnosispa Weise, new status (Fig. 42) Ocnosispa Weise 1910: 113 (as a subgenus of Sceloenopla). Type species: Chalepus cardinalis Guérin-Méneville (designated by Uhmann 1957b). Description.—Varying in form. Head: Medial sulcus present. Antenna: 11-seg- mented, apical 4 antennomeres shorter than preceding 4; antennomere I globular, small; If subcylindrical, twice length of I; HI to VI cylindrical, IV longest. Pronotum: Longer than wide, flattened; margined laterally; basal impression absent; tooth present or absent in anterior angle. Scutellum: Cuspi- date. Elytron: Humerus produced; exterior apical angle toothed or not; with 10 rows of punctures plus scutellar row; 4 costae, costa 2 highly carinate. Legs: Elongate, es- pecially anterior pair. Host plants.—Unknown. Described species.—19. Range.—Mexico to Peru. Pseudispa Chapuis, new status (Fig. 43) Pseudispa Chapuis 1875: 328. Type spe- cies: Cephalodonta marginata Guérin- Méneville (by original designation). Description.—Body elongate, slightly convex. Head: Small; medial sulcus pre- sent. Antenna: 11-segmented; antennomere I short; Il elongate; HI to VII obconical, decreasing in length; VIII to XI shorter than preceding, cylindrical, thickened. Prono- tum: Subparallel; basal impression present; longitudinal medial sulcus present. Elytron: Oblong, parallel-sided, almost vertical from costa 3; lateral and apical margins smooth; humerus not pronounced; apex almost ver- tical, truncate; secondary rows of punctures present between intervals 2 and 4. Host plants —Unknown. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Described species.—12. Key to species: Monros and Viana (1947), Argentina. Range.—Mexico to Argentina. Sceloenopla Chevrolat (Fig. 44) Sceloenopla Chevrolat 1837: 388. Type species: Hispa maculata Olivier (= His- pa spinipes Fabricius) (by monotypy). Cephalodonta Chevrolat 1837: 388. Nomen nudum. Cephalodonta Chevrolat 1843: 350. Type species: Hispa goniapterus Perty (by monotypy); Uhmann 1930a (synonymy). Chalepotatus (Macrochalepus) Pic 1929b: 4. Type species: Chalepotatus antennalis Weise (by monotypy); Monros and Viana 1947 (synonmy). Microdonta Chevrolat 1837: 388 not Hope 1837 [misapplied name]. Type species: Hispa serraticornis Fabricius (by mono- typy); Uhmann 1930a (synonymy). Description.—Varying in form. Head: Small, slightly convex; medial sulcus pre- sent; eye slightly prominent. Antenna: 11- segmented; apical 4 antennomeres longer than preceding 4; antennomere III longest. Pronotum: Trapezoidal, slightly convex; basal impression present; densely punctate; anterior angle prominent, directed forward; posterior angle directed obliquely back- wards. Scutellum: Quadrate. Elytron: Lat- eral margin smooth; apical margin dentate or not; external apical angle may have lam- ina or tooth; humerus produced; secondary punctures between intervals 2 and 4 absent. Host plants.—Persea (Lauraceae); Chry- sophyllum (Sapotaceae); Anthurium, Philo- dendrum (Araceae); Cocos (Arecaceae): Lonchocarpus (Fabaceae); Rapanea (Myr- sinaceae); Esembeckia (Rutaceae); Cerco- pia, Pourouma (Cercopiaceae); Rubiaceae; Cyclanthaceae (Jolivet and Hawkeswood 1995); Cupania (Sapindaceae); Clusia (Clusiaceae); Davilla (Dilleniaceae); Ster- culia (Sterculiaceae) (Hespenheide and Dang 1999). Described species.—154. Key to species: VOLUME 104, NUMBER 3 Uhmann (1937a), species groups; Staines (1996b), Nicaragua. Range.—Mexico to Argentina. Serratispa Staines, new genus (Figs. 45—46) Type species.—Serratispa quadricosta Staines, new species. Description.—Head: Vertex depressed, coarsely punctate. Antenna: |1-segmented; filiform; reaching beyond humeri. Prono- tum: ‘Trapizodial; lateral margin serrate; disc convex; punctate; basal impression present. Elytron: Lateral margin sparsely dentate near humerus; exterior apical angle and apical margin sparsely dentate; with 10 rows of punctures plus scutellar row; 4 cos- tae; horizontal plicae present on disc. Serratispa quadricosta Staines, new species Holotype.—[Brazil] Sao Paulo d’Olivenga, M. de Mathen, Juin Juillet/E Monros Collection 1959/Holotype Serratis- pa quadricosta, des. C. L. Staines 2001 (red label). Deposited in the National Museum of Natural History, Smithsonian Institution. Description.—Ovid, castanaceous with paler yellowish markings; legs yellowish. Head: Vertex depressed, coarsely punctate with a seta arising from each puncture; me- dial carina present; interantennal carina pre- sent; frons not projecting. Antenna: Anten- nomere I cylindrical, longer than II; I transverse; III—V cylindrical, each shorter than I; VI—-X transverse; XI pointed at apex; I-III punctate; IV—V with few pale setae; VI-XI darkly setose. Pronotum: Anterior angle rounded; lateral margin straight and convergent for basal %, then rounded and convergent; posterior angle acute; pair of setae present on anterior margin behind head; pronotal length 0.9 mm; pronotal width 1.6 mm. Scutellum: Cordate with di- amond-shaped sulci on lateral margin; mi- cropunctate. Elytron: Narrowing to apex; exterior apical angle rounded, sparsely den- tate; humerus rounded; with 11 rows of punctures at base, 10 on rest; punctures in 749 rows 4—10 larger than rest; highly sculp- tured, with 4 horizontal plicae on disc not reaching suture; interspaces 2 and 3 carinate on basal 4, carinae unite just behind hu- merus; shallow depression present on inter- spaces 4—9 behind humerus; interspaces |-— 3 carinate on apical 4; interspace 4 carinate along edge of depression; elytral length 3.0 mm; elytral width 2.1 mm. Total length: 4.3 mm. Etymology.—From serratus (Latin) = saw plus ispa for the Hispinae; referring to the sawlike lateral margin of the pronotum. Quadra (Latin) = four plus costa = ridge; for the four lateral plicae on the elytra. The name is a noun in apposition. Comparative notes.—Serratispa differs from all other hispine genera by the follow- ing combination of characters: Antenna | 1- segmented, filiform; head with vertex de- pressed, coarsely punctate; pronotum with a pair of setae on anterior margin behind head, lateral margin serrate; elytra costate with lateral plicae, with 11 rows of punc- tures at base. Host plant.—Unknown. TRIBE CHALEPINI WEISE 1910: 69 Octotomites Chapuis 1875: 310. Cephalodontites Chapuis 1875: 313 (in part). Uroplatini Weise 1910: 69. Type genus.—Chalepus Thunberg. Description.—Species of medium to small size; generally elongate, slightly con- vex. Head: Clypeus more or less long; la- brum reduced. Antenna: 3- to 11-segment- ed. Pronotum: A seta present on a small tubercle on anterior margin near angle. E/- ytron: Denticulate laterally and posteriorly; with 7 to 10 rows of punctures at base, scu- tellar row present or absent; usually with punctures in double rows separated by a costa, some with irregular puncture rows and tuberculate costae. KEY TO GENERA Le Antenna 3- to 7-segmented = Antenna 8- to 1l-segmented........ 1] 9(8). 10(5). 11(1). 12(11). 14(13). IS 3): 16(15). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Elytral costae irregular, tuberculate (Fig. 88) Ely tralicostac re cular an ene g} Prosternum expands anteriorly to partly cover mouth (Fig. 95) Sternocthispa Prosternum does not expand anteriorly, Physocoryna NO ONSooyvoINO I soon aoaaccbocuc 4 Elytron with 8 or 10 rows of punctures at DAS Cis crs eM e 2 Clcbays ssa eee 5) Elytron with 9 rows of punctures at base (igs W2)i4 ALM AM. Bae Se tthe Heptachispa Elytron with 8 rows of punctures at base sich 5) oy See HEMMER a HSA Be sls SP EPPO CHET 6 Elytron with 10 rows of punctures at base erat Lieu. cuct ty ites pokes kh os ope cian ee 10 Elytron with 10 rows of punctures on api- Gall A (EWE. 7/3) o-4.0 5 6 8 oo oo Heptatomispa Elytron with 8 rows of punctures for en- tineslen pth syst oe rate ae see cae 7 Antenna short or very short, clavate (Fig. SD) ot Mey Sad ee oes tg Se OR lee bees Bruchia Antennaylonessilifonmiscies ies o eee 8 Antenna 3- to 7-segmented; last anten- nomere acutely pointed at apex (Fig. 47) AEST IEUL A SRS sgt eal Bae, io Whkeet suet Acanthodes Antenna 7-segmented; last antennomere not acutely pointed at apex ......... 9 Penultimate antennomere longer than ul- Wubmavel ten (EIR YDY 3g Oe eee © oo 0 Oe Heptispa Penultimate antennomere shorter than ul- timate (Fig. 78) Apical shorter than three preceding combined (Fig. 66) Apical four antennomeres combined lon- ger than three preceding combined (Fig. 58) Nanocthispa four antennomeres combined Corynispa Brachycoryna (in part) Antennass-Sepmented) a. reeset 12 Antenna 9- to 1l-segmented........ 34 Antenna short or very short (barely reach- ing anterior margin of pronotum), clavate MOET Sat see ARSE TPM EL AORN Eee Pe ra A 13 Antenna long (reaching beyond base of pronotum), filiform or clavate ....... 19 . Elytral costae irregular, tuberculate (Figs. CANCE Moll Namen are? Sed cure erohr, came meme cer 14 Elytralvcostacmae pullatae) serene eer 15 Antennomeres VII and VIII almost fused, antennomeres I to VI thick (Fig. 48) ... ak ets A) 2 ae Pere va od weiner tes Soke Acritispa Antennomeres VII and VIII distinct, an- tennomeres I to VI normal (Fig. 81) ... Octotoma (in part) Third tarsomere not bilobed (Figs. 93-94) Stenopodius babraciie is, osu 16 Elytron on apical % with 8 rows of punc- tures 17(16). 18(16). 19(12). 20(19). 22(21). 27(26). 28(27). 29(28). 30(29). ail(Z7). Elytron on apical % with 10 rows of punc- tures Antennomere VII as wide as VIII (Fig. 81) Octotoma (in part) Antennomere VII narrower than VIII Parvispa Elytron with 10 rows of punctures at base (Fig. 58) Brachycoryna (in part) Elytron with 8 rows of punctures at base (Fig. 70) Fossispa Elytron with 10 rows of punctures and four costae: SSA oa eee eee 20 Elytron with 8 rows of punctures and three.cOstaé: 4.03 eee 23 First and fourth elytral costae united at apex; profemur with two teeth (Fig. 90) Probaenia First and fourth elytral costae not united at apex; profemur with one tooth or un- ATMS SF, os Su ae: were a eee RRC Rae DI . First elytral costa strongly elevated; 2nd elevated on apical 4%; 3rd elevated on bas- al 4%; 4th complete, only finely elevated; puncture rows complete (Fig. 75) ... 5, See EAT cs Bis. cue o's Heterispa IK AMVINOE EMO a csoudogoasoc 22 Antennomeres laterally compressed, tri- angular; vertex of head without longitu- dinal’sulcus (Fiss"86)) Somer Oxyroplata Antennomeres not laterally compressed, cylindrical; vertex of head with longitu- dinalsuleus (fig. 104)i5 4 33s Uroplata . Vertex of head without sulcus ...... 24 Vertex of head with 1 to 5 sulci ..... DS . Frons not projecting (Fig. 82) . . Octuroplata Frons projecting Mimoethispa . Elytron with 10 rows of punctures (Fig. OD) Mees he, Jit eee Spaethispa Elytron with 8 rows of punctures 26 . Elytron with tubercles (Fig. 69) Euprinota Elytron without tubercles .......... DY Vertex of head with medial sulcus 28 Vertex of head with 3 to 5 sulci ..... 31 Sulcus present on outer margin of eye (E1285 0) Mayes) 2s Cs eee Octhispa Sulcus absent on outer margin of eye Pardo oO Slee NG Cie ROR ET eRe eS Gor 29 Sulcus present on inner margin of eye (Fig. 103) Temnocthispa Sulcus absent on inner margin of eye Frons projecting; lateral and apical mar- gins of elytra serrate (Fig. 57) Bothrispa Frons not projecting; lateral and apical margins of elytra smooth (Fig. 77) ee oe ee ne Microrhopala Pronotum with basal impression .... . 32 Pronotum without basal impression . . . VOLUME 104, NUMBER 3 32(31). 331). 40(39). 41(37). 42(41). 44(43). 45(44). Antennomeres III to V cylindrical (Fig. 60) Oe eee eae aee, shad Sleeves Arete Carinispa Antennomere III cylindrical, TV and V transverse (Fig. 87) Antennomere III with tooth-like projec- tion on outer apical margin (Fig. 89) Platocthispa Pentispa Antennomere III without tooth-like pro- jection on outer apical margin (Fig. 71) Glyphuroplata WAMtennano-Sepmentediqnis + .\icislsei.. « 35 Antenna 10- or 1l-segmented....... 37 . Elytron with 10 rows of punctures at base (Fig. 68) Elytron with 8 rows of punctures at base Decatelia (in part) FORA GUE IGec ROO Gp caciy ae oeer e 36 . Exterior apical angles of elytra expanded (Rig OS) so. ee a cee seater et een Cnetispa Exterior apical angles of elytra not ex- pandedt (Bic. 79). re ase, cide © = Nonispa 7 AntennagiO-seementeds i... .25 55 -- 38 Antennaplil=seomented es. css... 41 . Elytron with 10 complete rows of punc- tures (Fig. 68) Elytron with 8 complete rows of punc- tures Decatelia (in part) . Prosternum projecting forward, with two blunt teeth on anterior margin (Fig. 97) PE ih cto Golo ae Oney oan Cee one Sternoplispa Prosternum not projecting forward, with- out teeth on anterior margin Mesofemur with tooth (Fig. 99) Sternostenoides Mesofemur without tooth Charistena Lateral margin of pronotum with two teeth (Fig. 83) Lateral margin of pronotum without teeth Odontispa Pronotum conspicuously margined later- ally (Fig. 67) Crasepodonispa Pronotum finely or not margined laterally BES eee Cea A aes Ot eee ere 43 . Elytron at base with 3 costae and 8 rows of punctures (in some species additional rows present either behind middle or at base), scutellar row present or absent Elytron at base with 4 costae and 10 rows of punctures (sometimes reduced to 8 or 9 rows in middle), scutellar row present or absent Anterior margin of prosternum projecting toward mouth (Fig. 98) Sternostena Anterior margin of prosternum not pro- jectins-toward mouth) 2a. as). is on 45 Femora with a large, ventral tooth at mid- limen(Rics 50) ie AP. SS. Seer Agathispa 46(45). 47(46). 48(47). 49(43). 50(49). 51(50). 57(56). . Abdominal sterna Femora without a large, ventral tooth at midline (may have small denticules) .. 46 Body slender, almost cylindrical, eye not projecting (Fig. 52); mesotibia strongly CUEVEU MES eter ete re cee eet ec. 47 Body broader, especially at humeri (Fig. 101); eye slightly projecting; mesotibia not stronely curved... eee Sumitrosis Costae on elytral intervals 2 and 6 unite on apical fifth (Fig. 53) = Anisostena (s. str.) Costae on elytral intervals 2 and 6 do not unite on apical fifth Apical five antennomeres wider than pre- ceding (Fig. 54) Apical five antennomeres not wider than Anisostena (Apostena) preceding (Fig. 55) aody esd) SRE ap omen Anisostena (Neostena) An outward pointing spine produced at humeral angle; exterior apical angle of el- ytra forms a strong, sharp tooth (Fig. 100) 2 la dla Sous Bye pe th oR cutene cick bes Stethispa Humeral angle with incurved tooth or hu- meral angle angular, without a spine; ex- terior apical angle of elytra rounded or ends ina broad; flattooth 22... ..:- 50 Humerus with recurved tooth ...... nicd' Hie yn 2k. ENE ES Chalepispa Fhimenus angular. Seieeee ees . s .s - ai Anterior margin of prosternum expanded towardimouthiysS << peat Chalepotatus Anterior margin of prosternum not ex- pandeditoward miouthy <774713.. 2 sat os 52 . Form elongate; elytron with 10 rows of PUNCHITESFAtDASE, sevy cesta oto a3 Form oval or cuniform; elytron with 11 rows of punctures at base ......... 55) . Apical tarsomere with one claw Anisochalepus Apical tarsomere with two claws .... 54 . Antennomere III twice as long as I or IV UD Nes A ee ee ae Goyachalepus Antennomere III subequal in length to II or IV Chalepus 1 and 3 with medial sulcus (Fig. 76) Abdominal sterna | and 3 without medial Metaxycera SUICUS ee ieee: cic Me wes slensee 56 . Apices of elytra regularly rounded, sutur- alvanglesrectanculate see ae eset. = ee Sy) Apices of elytra conjointly rounded, su- tural anglevemaroimates s <2 2 cee. w ccs a9 Elytral costa 3 complete (Fig. 84) ..... BES CRONE Oe Oita eC Ria Treo NOES Odontota Elytral costa 3 interrupted in middle or ADSENSE Fa. Meee Ae rele eee bene 58 To? 58(57). Head constricted behind eyes (Fig. 102) hbesh ea atiye cee me ewes ate Sie Temnochalepus ~ Head not constricted behind eyes (Fig. 56) RO aesMates ie ee Gs ters Heaelis Baliosus 59(56). Apical antennomere pointed at apex (Fig. S'S) ete aod er cteicgss aston ayee Oxychalepus — Apical antennomere rounded at apex... 60 60(59). Elytron with costae 1 and 4 complete, costae 2 and 3 almost completely obso- lete; costa 1 carinate, 4 less elevated (Fig. 105) - Elytron with costae 1, 2, and 4 complete, 3 only visible at base and apex (Fig. 106) See Eateacieonaad S. Xenochalepus (Neochalepus) Mee ere og Xenochalepus (s. str.) Acanthodes Baly (Fig. 47) Acanthodes Baly 1864: 262. Type species: Acanthodes generosa Baly (by original designation). Acanthispa Chapuis 1875: 323 [lapsus cal- ami). Description.—Body wedged-shaped. Head: Long, flattened; projection present between eyes. Antenna: Rigid, 3- to 7-seg- mented; apical antennomere acutely point- ed. Pronotum: Slighty narrower than elytra, transverse; medially depressed; basal mar- gin with excavation present on each side. Scutellum: Transverse, apex obtuse. Ely- tron: Parallel-sided, slightly enlarged api- cally; spine-like tooth in exterior apical an- gle; with 8 rows of punctures; extra punc- tures present, especially between rows 5 and 7. Host plants.—Quetzalia (Celastraceae) (Jolivet and Hawkeswood 1995). Described species.—15. Key to species groups: Spaeth (1937). Range.—Colombia to Argentina. Acritispa Uhmann (Figs. 48—49) Acritispa Uhmann 1940b: 143. Type spe- cies: Acritispa dilatata Uhmann (by mon- otypy). Description.—Head: Medial sulcus pre- sent; vertex micropunctate. Antenna: 8-seg- mented; antennomeres I and II subequal in length; II widened apically; VII as wide as PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VIII, difficult to distinguish; VIII hirsute club. Pronotum: Wider than long; tooth present in anterior angle; lateral margin evenly convergent from base to apex; two tooth-like projections present on lateral margin; densely coarsely punctate. Elytron: Greatly expanded at apex; lateral and apical margins dentate; with lateral and transverse plicae; translucent at exterior apical angle. Legs: Profemur expanded apically. Host plants.—Unknown. Described species.—2. Staines (1988). Range.—Panama to Bolivia. Key to species: Agathispa Weise (Figs. 50-51) Agathispa Weise 1905a: 64. Type species: Hispa dimidiata Olivier (by monotypy). Description.—Moderately attenuate. Head: Clypeus triangular. Antenna: | 1-seg- mented; antennomeres I to VI short; VII to XI longer and wider. Pronotum: Transverse, narrower in front, obtusely angulate in mid- dle. Scutellum: Quadrate. Elytron: Rectan- gular; with 8 rows of punctures, rows I—5 regular at base, irregular after middle; scu- tellar row absent; first costa complete, curves outward at apex. Legs: Long; fem- ora with a tooth; tibiae slightly curved. Host plant.—Unknown. Described species.—1. Range.—Dominican Republic. Anisochalepus Uhmann Anisochalepus Spaeth 1937: 154 (unavail- able name, no type species designated). Anisochalepus Uhmann 1957a: 109. Type species: Anisochalepus Uhmann (by monotypy). reimoseri Description.—Narrow, elongate, cylin- drical. Head: Vertex twice as long as wide, flat, depressed between eyes; medial sulcus present; row of punctures present behind eye; clypeus very narrow; frons cristate, deeply punctate. Antenna: 11-segmented; antennomere III elongate, 2 as long as II, nearly quadrate; IV to VI combined as long VOLUME 104, NUMBER 3 3: oe, ws) Figs. 48-56. Dorsal habitus. 48, Acritispa dilatata Uhmann antenna. 49, Acritispa dilatata. 50, Agathispa dimidiata (Olivier), femur. 51, Agathispa dimidiata. 52, Anisostena (s. str.) sp. 53, Anisostena (s. str.) sp., apex of elytra. 54, Anisostena (Apostena) sp., antenna. 55, Anisostena (Neostena) missionensis Monr6és and Viana, antenna. 56, Baliosus nervosus (Panzer). 754 as III. Pronotum: Scarcely wider than long; lateral margin weakly convergent, margined behind middle; coarsely punctate; disc with transverse carina present; tooth present in anterior angle; basal impression present. Scutellum: Cordate. Elytron: Lateral margin finely dentate; apical margin strongly den- tate; with 10 rows of punctures plus scutel- lar row, rows 5 to 8 reduced to two rows behind middle; 4 costae; costa 1 strongly elevated, 2 and 4 weakly elevated, 3 nearly obsolete; sutural margin weakly carinate. Legs: Femora and tibiae unarmed; mesoti- bia weakly curved; femora punctate; apical tarsomeres with one claw. Host plant.—Unknown. Described species.—1. Range.—Paraguay. Anisostena Weise (Bigs 253) Anisostena Weise 1910: 120. Type species: Charistena elegantula Baly. Description.—Body elongate, coarsely punctate. Head: Frons medially impressed; vertex with medial sulcus present or absent; eye little swollen. Antenna: 11-segmented; antennomere I subglobose; II cylindrical, as wide as I; III to VI gradually widening; VII to XI forming a club. Elytron: With 8 rows of punctures plus scutellar row; tricostate. Legs: Mesotibia strongly curved. Host plants.—Bothrichloa, Panicum, Paspalum, Schizachrium, Tripsacum, Val- ota (Poaceae) (Staines 1994b, c, d). Described species.—23. Key to species: Monros and Viana (1947), Argentina; Staines (1994b, c, d). Range.—Canada to Argentina. Anisostena (Apostena Staines) (Fig. 54) Anisostena (Apostena) Staines 1993a: 186. Type species: Anisostena angustata Pic (by original designation). Description.—As in Anisostena (s. str.) expect: Head: Medial sulcus present; apical 5 antennomeres wider than preceding. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pronotum: Basal impression present; tooth present in anterior angle. Elytron: Costate elytral intervals 2, 4, and 6 do not unite near apex. Host plants.—Poaceae (Monr6os and Vi- ana 1947). Described species.—2. Key to species: Staines (1993a). Range.—Argentina. Anisostena (Neostena Monros and Viana) (Fig. 55) Anisostena (Neostena) Monros and Viana 1947: 201. Type species: Anisostena bi- colorceps Pic (by monotypy). Description.—As in Anisostena (s. str.) except: Head: Cylindrical; medial sulcus present; apical 5 antennomeres not wider than preceding. Pronotum: Basal margin bisinuate. Scutellum: Rounded at apex. El- ytron: Costate elytral intervals 2, 4, and 6 do not unite near apex. Host plants.—Olyra, Panicum (Poaceae) (Staines 1993a). Described species.—3. Key to species: Staines (1993a). Range.—Brazil to Argentina. Baliosus Weise (Fig. 56) Baliosus Weise 1905a: 64. Type species: Hispa nervosa Panzer (= Hispa rubra Weber) (by original designation). Parabaliosus Monros and Viana 1947: 254. Type species: Parabaliosus ogloblini Monros and Viana 1947 (by monotypy); Uhmann 1957b (synonymy). Description.—Wedge-shaped, somewhat flattened. Head: Vertex smooth or slightly punctate. Antenna: 11-segmented; apical 5 antennomeres somewhat dilated. Elytron: Much wider at apex; apex obtuse or ap- proaching truncate; with 10 rows of punc- tures plus scutellar row; 4 costae, costa 3 feebly interrupted in middle, oblique from umbone to 2nd or 3rd interval for a short distance forward. Legs: Short to moderately long; mesotibia not curved. VOLUME 104, NUMBER 3 Host plants.—Ceanothus (Rhamnaceae); Hippocratea (Celastraceae); Arrabidea (Bignoniaceae); Jatropha (Euphorbiaceae); Acer (Aceraceae); U/mus (Ulmaceae); Tilia (Tiliaceae); Desmodium, Dioclea, Meibom- ia, Robinia (Fabaceae); Bauhinia (Caesal- piniaceae); Veronia (Asteraceae); Cordia (Ehretiaceae); Urtica (Urticaceae); Olyra (Poaceae); Lippia (Verbenaceae); Castanea, Quercus (Fagaceae); Salix (Salicaceae); Al- nus, Betula (Betulaceae); Carpinus (Carpi- naceae); Corylus (Corylaceae); Amelan- chier, Malus, Prunus, Pyrus, Rubus (Ro- saceae); Banisteria (Malpighiaceae); Citrus (Rutaceae); Guazuma (Sterculiaceae); Guaiacum (Zygophyllaceae); Basanacan- tha (Rubiaceae) (Jolivet and Hawkeswood 1995); Urera (Urticaceae); Odontonema (Acanthaceae); Bignoniaceae (Hespenheide and Dang 1999). Described species.—44. Key to species: Monros and Viana (1947), Argentina; Staines (1996b), Nicaragua. Range.—Canada to Argentina. Bothrispa Uhmann (Cait: By) Bothrispa Uhmann 1940b: 143. Type spe- cies: Uroplata depressa Chapuis (by monotypy). Description.—Head: Vertex with depres- sion between eyes, surface alutaceous; me- dial sulcus absent; frons projecting. Anten- na: 8-segmented; antennomere I flattened laterally; II as wide as long, cylindrical; HI longer than I or II; IV to VII transverse; VIII hirsute, pointed club. Pronotum: Wid- er than long, convex; densely punctate, ex- cept for small portion of disc; margined lat- erally; tooth present in anterior angle; basal impression present. Scutellum: Cuspidate. Elytron: Lateral and apical margins strong- ly dentate; with 8 rows of punctures at base, expands to 10 beyond middle; sutural mar- gin carinate; tricostate, none united. Legs: Profemur expanded to apex. Host plant.—Unknown. Described species.—1. Range.—Brazil. Brachycoryna Guérin-Méneville (Fig. 58) Brachycoryna Guérin-Méneville 1844: 280. Type Brachycoryna pumila Gueérin-Meéneville (by monotypy). Physocoryna: Baly 1885: 90 not Guérin- Méneville 1844 [misapplied name]. species: Description.—Head: Small; vertex not prominent, with longitudinal sulcus; eye not prominent, separated by more than width of an eye; area around eye, except along pit, rugose; antenna inserted into pit on front of head; pit divided by shallow, central keel. Antenna: Short; 7- or 8-segmented; basal antennomeres more or less similar, penul- timate slightly expanded, ultimate forming a hirsute, oval club. Pronotum: Wider than long, narrower than elytra, convex; punc- tures distributed over entire surface except along apical margin; medial callus usually present on disc. Scutellum: Small; quadrate. Elytron: Oblong-oval, longer than wide, parallel-sided; apex broadly, evenly round- ed; lateral and apical margins serrate; hu- meral angle rounded; with 10 rows of punc- tures, punctures usually distinct, arranged in pairs, but sometimes becoming confused, plus scutellar row; even intervals elevated and often costate. Legs: Short; tibiae and femora subequal in length; trochanters slightly angular; tibiae gradually widening to apex with tuft of setae at apex; femora widest at middle. Venter: Pro- and meso- sternum punctate in middle, alutaceous. Host plants.—Abutilon, Sida, Sphaeral- cea (Malvaceae); Waltheria (Sterculiaceae); Artemisia (Asteraceae) (Staines 1986). Described species.—7. Key to species: Staines (1986). Range.—Southern United States to Ar- gentina. Bruchia Weise (Fig. 59) Bruchia Weise 1906a: 227. Type species: Bruchia sparsa Weise (by monotypy). 756 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 59 : BA Oe PATA as © Figs. 57-65. Dorsal habitus. 57, Bothrispa depressa (Chapuis). 58, Brachycoryna pumila Guérin. 59, Bruchia sparsa Weise. 60, Carnispa nevermanni Uhmann. 61, Chalepotatus coarctatus (Chapuis). 62, Chalepus san- guinicollis (L.). 63, Charistena ruficollis (F). 64, Clinocarispa fasciata Weise. 65, Cnetispa darwini Maulik. VOLUME 104, NUMBER 3 Bruchiella Weise 1916: 39 (invalid replace- ment name for Bruchia). Description.—Head: Medial sulcus pre- sent. Antenna: 7-segmented; antennomeres I to V very sharp, gradually widening after III, apical two antennomeres less separated; I to III short, transverse; IV to V very short, strongly transverse; VI as long and wider than preceding two combined, as wide as VII; VII subovate, pointed at apex. Prono- tum: Narrow, width at base and apex equal; lateral margin deflexed, nearly parallel-sid- ed; disc convex, regularly subseriate-punc- tate; medial line finely impressed. Scutel- lum: Small, quadrate. Elytron: 3X as long as pronotum, slightly convex; apex con- jointly rounded; with 8 rows of punctures; lightly bicostate, costae interrupted, trans- versely united before apex. Host plants.—Unknown. Described species.—2. Range.—Costa Rica to Argentina. Carinispa Uhmann (Fig. 60) Carinispa Uhmann 1930a: 255. Type spe- cles: Carinispa nevermanni Uhmann (by monotypy ). Description.—Head: Vertex trisulcate, impunctate; frons projecting. Antenna: 8- segmented; antennomere I subglobular; II transverse; III to V cylindrical; VI to VII transverse; VIII pointed at apex. Pronotum: Wider than long, widest at base, convex, densely punctate; basal impression present; tooth present in anterior angle. Scutellum: Quadrate. Elytron: Lateral margin sparsely dentate; apical margin dentate; with 8 reg- ular rows of punctures; tricostate, costae weakly carinate. Host plants.—Bunchosia, Malpighia (Malpighiaceae) (Jolivet and Hawkeswood 1995)» Described species.—1. Range.—Costa Rica. TT Chalepotatus Weise (Fig. 61) Chalepotatus Weise 1910: 133. Type spe- cies: Odontota coarctatus Chapuis (des- ignated by Monros and Viana 1947). Description.—Body elongate, moderate- ly flattened. Head: Clypeus smooth or se- tose, with medial carina ending in a tooth before antennal base. Antenna: 11-seg- mented. Pronotum: Lateral margin round- ed; disc smooth, shiny. E/ytron: At humeri little wider than pronotum, sides moderate- ly parallel; apex conjointly rounded; with 10 rows of punctures plus scutellar row, puncture rows 5 to 8 narrowed to two rows on basal 42; 4 costae, costa 3 united with 4 for “4th its length. Venter: Prosternum with mid-anterior projection. Legs: Short. Host plants.—Unknown. Described species.—3. Key to species: Monros and Viana (1947), Argentina. Range.—Mexico to Argentina. Chalepsipa Uhmann Chalepsipa ignorata Uhmann (by mon- otypy). Description.—Head: Vertex extremely smooth, alutaceous; medial sulcus present; frons cuspidate, raised in profile; constrict- ed at base. Antenna (only two basal anten- nomeres present in holotype): Antennomere I globose, punctate; II transverse, punctate. Pronotum: Widest at base, weakly trans- verse; lateral margin parallel to weakly con- vergent; disc densely punctate; basal im- pression present; tooth present in anterior angle. Scutellum: Ligulate. Elytron: Paral- lel-sided, lateral margin smooth; apical margin with 5 teeth; humerus raised into backward pointed tooth; with 10 rows of punctures plus scutellar row; three complete costae (1, 2, 4), 3 visible at humerus and apex; costae 1 and 2 strongly elevated. Legs: Femora densely punctate; apical tar- somere with two claws. Host plant.—Unknown. 758 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Described species.—1. Range.—French Guyana. Chalepus Thunberg (Fig. 62) Chalepus Thunberg 1805: 282. Type spe- cies: Hispa sanguinicollis Linnaeus (des- ignated by Weise 1905a). Anoplitis Kirby 1837: 227. Type species: Hispa bicolor Olivier (by monotypy); Butte 1968b (synonymy). Parachalepus Baly 1885: 47. Type species: Parachalepus brevicornis Baly (by mon- otypy); Staines 1993b (synonymy). Description.—Head: Wider than long; vertex sulcate or micropunctate; frontal ca- rina feebly developed, elongate or dia- mond-shaped, extending between antenna and joining clypeal base; clypeus large and usually longer than wide, triangularly ele- vated, apex feebly arcuate, surface finely punctate or coarsely granularly tuberculate with long white setae. Antenna: 1|1-seg- mented; basal antennomere incrassate. Pronotum: Transverse; convex; slight basal depression with ante-scutellar transverse ca- rina; lateral margin angulate in middle. E/- ytron: Subelongate, parallel, slightly in- dented at sides; apex regularly rounded, finely serrulate; with 10 rows of punctures at base and apex, may be reduced after mid- dle, scutellar row absent. Venter: Sparsely, irregularly micropunctate; abdominal ster- nite 5 apically truncate. Host plants.—Theobroma (Sterculi- aceae); Bambusa, Brachiaria, Chusquea, Elymus, Hystrix, Lasiacis, Olyra, Panicum, Paspalum, Valota, Zea (Poaceae); Apios, Benthamantha, Calopogonium, Canavalia, Crotalaria, Cymbosema, Desmodium, Dio- clea, Dolichos, Falcata, Glycine, Lathyrus, Meibomia, Mucuna, Phaseolus, Pueraria, Robinia, Vicia (Fabaceae); Bauhinia, Cas- sia (Caesalpiniaceae); Prosopis (Mimosa- ceae); Aloysia, Vitex (Verbenaceae); Paul- linia (Sapindaceae); Philodendron (Ara- ceae); Cyanus, Eupatorium, Verbesina, Ve- ronia, Wedelia (Asteraceae); Cordia (Ehretiaceae); Quercus (Fabaceae); Cera- sus, Crataegus, Malus, Pyrus (Rosaceae); Ipomoea (Convolvulaceae); Commelina (Commelinaceae); Acer (Aceraceae); Betu- la (Betulaceae); Terminalia (Combreta- ceae); Bromelia (Bromeliaceae); Aristolo- chia (Aristolochiaceae); Arrabidea (Big- noniaceae); Actinostomon (Euphorbiaceae); Celastrus (Celastraceae); Sapindaceae (Jo- livet and Howkeswood 1995); Tilaceae (Hespenheide and Dang 1999). Described species.—92+. Key to spe- cies: Uhmann (1936a), covered 48 species; Monros and Viana (1947), Argentina; Butte (1968b), North America; Staines (1996b), Nicaragua. Range.—Canada to Argentina. Charistena Baly (Fig. 63) Charistena Baly 1864: 251. Type species: Hispa_ ruficollis Fabricius (by original designation). Description.—Head: Small; frons not prominent; eye finely granulate. Antenna: 10-segmented; antennomere I stout, oval; II longer than I; [II longer than II; [V to VI gradually shorter; VII to X distinct, shorter than others. Pronotum: Subcylindrical, not clearly margined. Elytron: Elongate, paral- lel-sided; apex rounded; with 8 rows of coarse punctures; tricostate. Legs: Mesoti- bia strongly arcuate, inner apical angle acute, slightly prolonged; tarsomeres dilat- ed, densely pubescent beneath; tarsomere 1 triangular, rather small; 2 reniform; 3 deep- ly bilobed; 4 slightly longer than lobes of 3; claws feebly curved, divergent. Host plants—Panicum, Paspalum, Zea (Poaceae) (Jolivet and Hawkeswood 1995). Described species.—6. Range.—Colombia to Argentina. Clinocarispa Uhmann (Fig. 64) Clinocarispa Uhmann 1935b: 227 (unavail- able name, type species not designated). Clinocarispa Uhmann 1940b: 143. Type VOLUME 104, NUMBER 3 species: Odontota sauveuri Chapuis (by original designation). Description.—Head: Medial sulcus pre- sent; rounded depression present on vertex; clypeus smooth, sparsely setose. Antenna: 1l-segmented, fusiform, not thickened at apex, slightly compressed laterally, striate; antennomere I subglobular; II transverse; II to V cylindrical, III longest; VI to X transverse; XI pointed at apex; VII to XI pubescent. Pronotum: Longer than wide, convex, uniformly punctate; basal impres- sion present; longitudinal medial carina pre- sent. Scutellum: Subquadrate. Elytron: Sub- parallel, slightly divergent posteriorly; lat- eral margin smooth; apical margin dentate; with 8 rows of punctures plus scutellar row, with some additional puncture rows on hu- merus; tricostate. Legs: Tarsal claws with two teeth. Venter: Prosternum elevated be- tween front coxae, depressed toward mouth, not projecting forward. Host plants.—Bambusa, Olyra (Poaceae) (Ramos 1996). Described species.—7. Key to species: Ramos (1996). Range.—Colombia to Peru. Cnetispa Maulik (Fig. 65) Cnetispa Maulik 1930: 49. Type species: Cnetispa darwini Maulik (by original designation). Description.—Small. Antenna: 9-seg- mented; short, barely reaching to base of pronotum, not produced into acute point at apex; antennomeres I and II distinct; III to IX forming elongate, slightly flattened club, III longest; IX broadest at apex. Pronotum: Subcylindrical, narrowed apically; light prescutellar impression present. Scutellum: Rectangular. Elytron: Lateral margin ser- rate; external apical angle expanding but not spine-like; with 8 rows of punctures plus scutellar row. Host plants.—Centrosema, Cymbosema, Desmodium (Fabaceae) (Jolivet and Haw- keswood 1995). 759 Described species.—4. Range.—Costa Rica to Argentina. Corynispa Uhmann (Fig. 66) Corynispa Uhmann 1940b: 143. Type spe- cies: Uroplata clavicornis Uhmann (by monotypy ). Description.—Head: Medial sulcus ab- sent; vertex alutaceous, with depression be- tween eyes; carina present from depression to antennal base. Antenna: 7-segmented; antennomere I subglobose, II transverse, III cylindrical; IV to VI transverse, VII 2X as wide as the preceding. Pronotum: Longer than wide, parallel-sided; with scattered large punctures; two areas on either side of disc with dense punctures. Elytron: Lateral and apical margins dentate; dilated at apex; with 10 rows of punctures; costae carinate, especially on basal 1/5; costa 2 highly car- inate at humerus, continues on basal %, more elevated at apex. Legs: Thickened; tarsal claws with tooth at base. Host plant.—Stigmatophyllum (Malpigh- iaceae) (Jolivet and Hawkeswood 1995). Described species.—1. Range.—Brazil. Craspedonispa Weise (Fig. 67) Craspedonispa Weise 1910: 125. Type spe- cies: Craspedonispa modesta Weise (by monotypy). Description.—Body moderately elon- gate, almost parallel, slightly convex. Head: Clypeus triangular. Antenna: | 1-seg- mented, moderately short, filiform; apical 5 antennomeres thickened. Pronotum: 2 as wide as long, narrowing basally; lateral margin rounded, broadly and conspicuously margined. Scutellum: Wider than long; nar- rowing behind; apex rounded. Elytron: Wider than pronotum; parallel-sided; lateral margin with tooth; apex narrowed, con- jointly rounded; with 10 rows of punctures plus scutellar row, punctures close together; punctures of row 10 transverse and reach to 760 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON NO pte CLAS SEES OTF Cot ae CR RT Meee) ee Figs. 66-74. Dorsal habitus. 66, Corynispa clavicornis Uhmann. 67, Craspodonispa sp. 68, Dectalia lema Weise. 69, Euprinota aterrima (Guérin). 70, Fossispa lutena Staines. 71, Glyphuroplata uniformis (Smith). 72, Heptachispa texta Uhmann. 73, Heptatomispa kesseli Unmann. 74, Heptispa solarii Weise. VOLUME 104, NUMBER 3 lateral margin; unicostate. Venter: Proster- num narrowed between coxae. Legs: Mod- erately short. Host plant.—Saccharum (Poaceae) (Jo- livet and Hawkeswood 1995). Described species.—2. Range.—Trinidad to Brazil. Decatelia Weise (Fig. 68) Decatelia Weise 1904: 435. Type species: Decatelia lema Weise (by monotypy). Paradecatelia Uhmann 1940b: 144. Type species: Paradecatelia viridis Uhmann (by monotypy); Monros and Viana 1947 (synonmy). Description.—Body small; rounded in front; eye large. Head: Clypeus triangular, overlapping labrum. Antenna: Reaching hu- ‘meri, filiform, 9- or 10-segmented; anten- nomere II longer than I; II not significantly longer than IH; apical 5 antennomeres thick- er and wider. Pronotum: Slightly wider than long, cylindrical; transverse sulcus present before base; lateral margin constricted be- fore base. Scutellum: Transverse, quadrate. Elytron: Wider than pronotum at humeri, almost parallel-sided; apex widely conjoint- ly rounded; truncate at base; with 10 rows of punctures plus scutellar row; rows 5 to 8 in a longitudinal cavity with angular sides; rows before cavity confused or re- duced to 2 or 3 rows. Venter: Abdominal sterna 1 and 2 separated by deep suture; prosternum simple, truncate behind. Legs: Mesotibia curved; tarsal claws moderately divergent. Host plants.—Unknown. Described species.—7. Key to species: Monroés and Viana (1947), Argentina. Range.—Colombia to Argentina. Euprionota Guérin-Méneville (Fig. 69) Euprionota Guérin-Méneville 1844: 278. Type species: Euprionota aterrima Guér- in-Méneville (by monotypy). 761 Description.—Head: Vertex with 3 to 5 longitudinal sulci. Antenna: 8-segmented; reaching to middle of elytra; antennomeres I to VII short, slightly compressed laterally; VIII wider, compressed laterally. Scutellum: Quadrate. Pronotum: Transverse; lateral margin sinuate, narrowing anteriorly; basal margin as wide as base of elytra. Elytron: Expanding apically; lateral and apical mar- gins strongly denticulate, teeth alternately large and small on apex; exterior apical an- gle with strong, laminate tooth; tricostate, costa | strong, entire, more elevated at base. Host plant.—Vernonia (Asteraceae) (Jo- livet and Hawkeswood 1995). Described species.—S. Range.—Mexico to Colombia. Fossispa Staines ig. 70) Fossispa Staines 1989a: 343. Type species: Fossispa lutena Staines (by monotypy). Description.—Head-: Surface micropunc- tate; medial sulcus present; 3 short lateral sulci present near eye; antenna inserted into quadrate pit; pit divided by keel; carina pre- sent around eye. Antenna: 8-segmented, clavate. Pronotum: Wider than long; com- pletely margined laterally; raised areas be- tween punctures micropunctate. Scutellum: Quadrate; micropunctate. Elytron: With 4 discal costae, costa 3 short and weak, costae 1, 2, and 4 united apically; punctures in double rows, with 8 rows of punctures ba- sally, 10 apically, basal rows 5 and 6 in longitudinal medial depression, latter rows briefly expand to 4 rows behind middle; a short, weak costa between apical rows 6 and 7; costae and raised areas between punctures micropunctate. Venter: Rugose at sides. Host plant.—Unknown. Described species.—1. Range.—Guatemala, Jamaica, and Mex- ico. Glyphuroplata Uhmann (Fig. 71) Glyphuroplata Uhmann 1940b: 143. Type species: Hispa pluto Newman (= Hispa 762 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON porcata Melsheimer) (by original desig- nation). Glyphuroplata Uhmann 1937b: 453 (un- available name, type species not desig- nated). Description.—Small, coarsely punctate. Head: Vertex trisulcate. Antenna: 8-seg- mented, gradually broadening apically. El- ytron: With 8 or 9 rows of punctures at base, at apical % may be 10 rows; punctures between rows 4 and 6 may be confused or expanded to 3 or 4 short rows; tricostate. Legs: Mesotibia with blunt inward project- ing tooth at apex; no distinct tooth before each tarsal claw. Host plants.—Digitaria, Eriochloa, Pan- icum, Valota (Poaceae) (Riley 1985). Described species.—4. Key to species: Riley (1985), North America. Range.—United States to Costa Rica. Goyachalepus Pic Goyachalepus Pic 1929a: 26. Type species: Goyachalepus donckieri Pic (by mono- typy). Description.—Head: Elongate; vertex micropunctate with scattered large punc- tures; medial sulcus present, becomes keel before base of antenna; two lateral sulci present near top of eye; frons projecting. Antenna: 11-segmented, robust, laterally compressed; antennomere I globose; II transverse; III largest, 2.5 length of II or IV; IV to X transverse; XI appendiculate. Pronotum: Wider than long; with large, deep punctures; lateral margin sinuate; tooth present in anterior angle; basal im- pression present. Scutellum: Quadrate; alu- taceous. Elytron: Elongate; lateral margin serrate; apical margin dentate, laminate; with 8 rows of punctures at base plus scu- tellar row; on apical 1/5 puncture rows 5 and 6 expand to two additional rows divid- ed by a small costa; suture carinate; tricos- tate, none united; costae | and 3 straight, 2 sinuate on apical 1/5. Venter: 5 visible ab- dominal sterna. Legs: Apical tarsomeres with two claws. Host plant.—Unknown. Described species.—1. Range.—Brazil. Heptachispa Uhmann (Fig. 72) Heptachispa Uhmann 1953: 876. Replace- ment name for Heptatoma Weise 1921: 187 [preoccupied by Heptatoma Meigen (Diptera)]. Type species: Uroplata cras- sicornis Chapuis. Description.—Head: Vertex alutaceous; medial sulcus deep, wide; antennal base on a projection from head. Antenna: 7-seg- mented, thick; antennomeres I to VI trans- verse, I to IV strongly so; VII longest, a rounded, flattened club. Pronotum: Com- pletely margined; basal margin bisinuate; with deep punctures. Scutellum: Quadrate. Elytron: Exterior apical angle expanding into tooth-like lamina; with 9 rows of punc- tures at base, rows 5 to 8 expand to an ad- ditional row after middle; tricostate. Legs: Tarsal claws with a tooth near base. Host plants—Unknown. Described species.—S. Range.—Brazil to Paraguay. Heptatomispa Uhmann (Fig. 73) Heptatomispa Uhmann 1940b: 143. Type species: Heptatomispa kesseli Uhmann (by monotypy). Description.—Small, parallel-sided. Head: Medial sulcus absent; vertex with depression behind eye; surface micropunc- tate. Antenna: 7-segmented, large; anten- nomeres I to VI transverse, punctate; VII thick, oval; VI not as long as preceding two combined, transverse. Pronotum: Convex; alutaceous; with large punctures, along base arranged in two rows. Elytron: With 8 rows of punctures at base, expands to 10 on api- cal %; 4 costae, 1, 2, and 4 entire; aluta- ceous. Host plant.—Unknown. Described species.—1. Range.—Brazil. VOLUME 104, NUMBER 3 Heptispa Weise (Fig. 74) Heptispa Weise 1906b: 241. Type species: Heptispa solarii Weise (by monotypy). Description.—Body strongly flattened. Head: Medial sulcus present; sulcus present on inner margin of eye; vertex with few scattered punctures. Antenna: 7-segmented; antennomeres I and II transverse, subequal in length; HI longer, cylindrical; [TV to VI transverse, decreasing in length; VII lon- gest, hirsute, slightly pointed at apex. Pron- otum: Transverse, completely margined: with large punctures; tooth present in an- terior angle; prebasal impression present. Scutellum: Transverse, rounded at apex. E/- ytron: Parallel-sided; lateral and apical mar- gins dentate; with 7 entire rows of punc- tures plus a short row at the humerus, scu- tellar row absent; tricostate, costae 1 and 2 entire, 3 begins at humerus, all united at apex. Venter: Metasternum punctate at sides; last abdominal sternite punctate in middle. Host plants.—Cassia (Caesalpiniaceae): Inga (Mimosaceae); Machaerium (Faba- ceae); Serjania (Sapindaceae) (Jolivet and Hawkeswood 1995); Mimosa (Mimosa- ceae) (Hespenheide and Dang 1999). Described species.—6. Key to species: Staines (1996b), Nicaragua. Range.—Mexico to Brazil. Heterispa Chapuis (Fig: 75) Heterispa Chapuis 1875: 321. Type species: Heterispa infuscata Chapuis (by original designation). Description.—Head: Small, globose; vertex with medial sulcus; conical projec- tion forward to between eyes; antenna with conical projection at base; eye large, con- vex. Antenna: 8-segmented, extends be- yond base of pronotum; club with whorls of setae. Pronotum: Quadrangular or slight- ly transverse, narrower than elytra; narrow- ing from base to apex; light prescutellar im- 763 pression present. Scutellum: Longer than wide. Elytron: Oblong or elongate, subpar- allel; obtuse projection present in exterior apical angle; lateral margin finely serrate; apical margin with many strong, sharp, un- equal spines; with 10 rows of large, regular punctures plus scutellar row; elytral interval 2 strongly costate; 4 costate at humerus and apex; 6 visible only at base; 8 entire, less elevated. Venter: Mesosternum swollen an- teriorly. Legs: Long, slender. Host plants.—Althaea, Malvastrum, Sida, Sphaeralcea (Malvaceae); Triumfetta (Tiliaceae); Guazuma (Sterculiaceae); Pan- icum, Stenotaphrum (Poaceae) (Jolivet and Hawkeswood 1995); Apeiba (Tilaceae) (Hespenheide and Dang 1999). Described species.—5. Range.—Mexico to Argentina. Metaxycera Baly (Fig. 76) Metaxycera Baly 1864: 255. Type species: Cephalodonta purpurata Guérin-Méne- ville (by original designation). Description.—Body wedge-shaped, flat. Head: Clypeus_ transverse-triangular, ex- tending to base of antenna. Antenna: 11- segmented, strongly filiform, apical 5 an- tennomeres thickened. Pronotum: Almost 2x as wide as long; lateral margin weakly rounded, slightly margined. Scutellum: Wider than long, apex rounded. Elytron: At humeri wider than pronotum, gradually nar- rowing to apex; apex widely conjointly rounded; with 10 rows of punctures plus scutellar row; 4 costae. Venter: Abdominal sterna | and 3 with deep sulcus in middle. Host plants.—Cecropia (Cecropiaceae) (Jolivet and Hawkeswood 1995). Described species.—12. Range.—Colombia to Uruguay. Microrhopala Chevrolat (Fig. 77) Microrhopala Chevrolat 1837: 339. Type species: Hispa vittata Fabricius (desig- nated by Baly 1864). 764 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON RAST PAT 0 OA RCS ARE A ESP 4 wm i FS 8 GOEL LLP ALLAN EDL EEE LA RE IS Figs. 75-83. Dorsal habitus. 75, Heterispa vinula (Erichson). 76, Metaxycera sexpustulata Baly. 77, Mi- crorhopala vittata (FE). 78, Nanocthispa atra (Weise). 79, Nonispa carlosbruchi Maulik. 80, Octhispa sp. 81, Octotoma scabripennis Guérin. 82, Octuroplata octopustulata (Baly). 83, Odontispa bimaculata Uhmann. VOLUME 104, NUMBER 3 Description.—Head: Subglobular; vertex mesally impressed, impression bordered on each side by longitudinal row of punctures; longitudinal sulcus present between anten- na; area surrounding eye punctate. Antenna: Reaching beyond base of pronotum; anten- nomeres VII and VIII wider and more pu- bescent than preceding. Pronotum: Longer than wide, widest posteriorly, often nar- rowed anteriorly, narrower than elytra at humeri; convex; anterior margin straight, lateral margin straight, arcuate, sinuate or bisinuate, posterior margin bisinuate; punc- tation usually dense; a slender, usually slightly elevated strip lacking coarse punc- tures present in front of scutellum. Elytron: Longer than wide; usually slightly nar- rowed behind humerus; with 8 rows of punctures plus a scutellar row; rows | and 2 extending to apex, 8 and 9 separate or sometimes confused apically, extending to near suture where they unite with | and 2; intervals | and 9 and also 3 and 7 uniting apically; interval 9 strongly elevated. Ven- ter: Prosternum margined anteriorly by row of short setae; abdomen sparsely punctate and pubescent, apical sternite more coarsely punctate and often more pubescent than preceding sterna. Host plants.—Ambrosia, Aster, Boltonia, Brickellia, Chrysopsis, Encelia, Franseria, Helianthus, Silphium, Sericocarpus, Soli- dago (Asteraceae); Salvia (Lamiaceae) (Jo- livet and Hawkeswood 1995). Described species.—19. Key to species: Clark (1983), North America; Staines (1996b), Nicaragua. Range.—Canada to Colombia. Mimoethispa Pic Mimoethispa Pic 1927b: 10. Type species: Mimoethispa irregularis Pic (by mono- typy). Description.—Oblong-elongate, some- what dilated apically, shining. Head: Vertex micropunctate; medial sulcus absent; frons with projection at base of antenna; clypeal margin evenly arcuate. Antenna: 8-seg- 765 mented, antennomeres I to VI small; VII elongate, as long as IV to VI combined; VIII largest, 3 length of VII, apex some- What attenuate. Pronotum: Widest at base, narrowing anteriorly; disc impunctate; basal impression present, with scattered punc- tures; laterally with dense, large punctures. Scutellum: Triangular; micropunctate. Ely- tron: Lateral and apical margins dentate; flattened lamina present from exterior api- cal angle to suture; humerus pronounced; with 8 rows of punctures at base, expanded to 9 rows posteriorly, additional row pre- sent between costae 2 and 3; tricostate, cos- ta 3 as wide as a puncture. Host plant.—Unknown. Described species.—1. Range.—Brazil. Nanocthispa Monros and Viana (Fig. 78) Nanocthispa Monros and Viana 1947: 312. Type species: Octhispa atra Weise (by monotypy). Description.—Head: Finely punctate; opaque; with very fine pubescence; frons depressed, concave, with some sulci; clyp- eus large, flat, less densely punctate than rest of head, subquadrate; eye small, with carina behind. Antenna: Short, 7-segment- ed; antennomere II 1.5 the length of I. Pronotum: Subcylindrical; transversely convex. Scutellum: Small, quadrangular; angle rounded. Elytron: Parallel-sided; apex conjointly rounded, with 2 small teeth; with 8 rows of punctures plus scutellar row; tri- costate, all unite at apex. Host plant.—Unknown. Described species.—1. Range.—Argentina. Nonispa Maulik (Fig. 79) Nonispa Maulik 1933: 606. Type species: Nonispa carlosbruchi Maulik (by mono- typy). Enneachalepus Spaeth species: Enneachalepus 1934: 207. Type deruginosus 766 Spaeth (by monotypy); Uhmann 1957b (synonymy). Description.—Body oblong, slightly nar- rowed in front, widest at apex. Head: Wide, convex, smooth; eye small, not convex, ca- rina absent on inner margin; clypeus wider than long, punctate. Antenna: 9-segmented. Pronotum: Longer than wide, scarcely punctate; slight basal impression present. Scutellum: Quadrate, rounded at apex. El- ytron: Parallel-sided; lateral margin smooth; upper surface smooth, without el- evations, depressions or roughness; apex rounded, small emargination at apical su- tural angle; with 8 rows of punctures, scu- tellar row absent; tricostate, costae more el- evated toward apex. Venter: Smooth. Legs: Slender, unarmed. Host plant.—Unknown. Described species.—1. Range.—Bolivia to Argentina. Octhispa Chapuis (Fig. 80) Octhispa Chapuis 1877: 23. Type species: Octhispa fossulata Chapuis (designated by Monros and Viana 1947). Oethispa Pic 1927b: 12 (lapsus calami). Description.—Body moderately attenuate or very narrow. Head: Small; eye slightly prominent; vertex with medial sulcus; clyp- eus prominent, punctate. Antenna: Moder- ately long or short, 8-segmented, more or less clavate; apical antennomere rounded, middle antennomeres compressed laterally. Pronotum: Narrowed anteriorly; basal im- pression present; punctate. Scutellum: Qua- drangular, angle rounded. Elytron: Humerus may have large tooth; lateral and apical margins dentate; with 8 rows of punctures plus scutellar row; tricostate; apex rounded, emarginate or truncate. Host plants.—Dioclea, Machaerium (Fa- baceae); Inga (Mimosaceae); Cassia (Cae- salpiniaceae); Basanacantha (Rubiaceae); Adenocalymna, Paullinia, Serjania (Ehre- tiaceae); Coccoloba (Polygonaceae); Byr- sonima, Malpighia (Malpighiaceae) (Jolivet PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and Hawkeswood 1995); Stigmatophyllum (Malphigiaceae); Colubrina (Rhamnaceae); Ochroma (Bombaceae) (Hespenheide and Dang 1999). Described speices.—102. Key to species: Staines (1996b), Nicaragua. Range.—Mexico to Argentina. Octotoma Chevrolat (Fig. 81) Octotoma Chevrolat 1837: 390. Type spe- cies: Hispa plicatula Fabricius (by mon- otypy). Description.—Body wedged-shaped. Head: Medial sulcus present or absent. An- tenna: 8-segmented; reaching to base of pronotum; antennomere I subglobose; VII as wide as VIII; VIII oval, subacute at tip. Pronotum: Transverse; tooth present in an- terior angle; margined laterally; four de- pressions present, two before and two after midline. Elytron: With 8 or 10 rows of punctures plus scutellar row; costae may be developed; some species with transverse and lateral carinae, these species having asymmetrical sculpture. Legs: Femora dis- tinctly sinuate beneath near apex; tibiae curved at base, protibia more dilated than others and with outer edge sinuate; tarso- meres dilated, densely pubescent beneath, tarsomeres | small, twice as long as wide, 3 longer than 2 and deeply bilobed, 4 little longer than lobes of 3; claws divergent. Host plants.—Clerdendrum, Lantana, Tectona, Verbena (Verbebaceae); Mentha, Monarda, Origanum, Salvia (Lamiaceae); Campsis, Tecoma (Bignoniaceae); Cana- valia, Cymbosema, Dioclea, Lespedeza (Fa- baceae); Fraxinus (Oleaceae); Sesamum (Pedaliaceae); Eupatorium (Asteraceae); Quercus (Fagaceae); Xanthorrhoea (Xan- thorrhoeaceae); Stigmatophyllum (Malpigh- iaceae) (Jolivet and Hawkeswood 1995). Described species.—10. Key to species: Staines (1989b, 1994a). Range.—United States to Brazil. VOLUME 104, NUMBER 3 Octuroplata Uhmann (Fig. 82) Octuroplata Uhmann 1937b: 454 (unavail- able name, type species not designated). Octuroplata Uhmann 1940b: 143. Type species: Uroplata octopustulata Baly (by original designation). Description.—Head: Oblong, rounded anteriorly; vertex without sulci; interocular impression present which becomes keel an- teriorly; eye slightly prominent; clypeus smooth, flat. Antenna: 8-segmented; apical antennomere pointed. Pronotum: Trans- verse; laterally rounded; punctate. Scutel- lum: Oblong. Elytron: With 8 rows of punc- tures plus scutellar row; tricostate (1, 2, 4) in anterior center, becomes 2 lines of sunk- en punctures and vestige of costa 3 on api- cal %. Legs: Short; tarsal claws with tooth at base. Host plant.—Senna (Fabaceae) (Teixeira Etaale 999): Described species.—9. Range.—French Guyana to Argentina. Odontispa Uhmann (Fig. 83) Odontispa Uhmann 1940a: 120. Type spe- cies: Prospodonta latipennis Pic (by monotypy). Description.—Elytra much wider than pronotum. Head: Vertex smooth; frons elongate, projecting between antennal ba- ses; antennal insertion below midpoint of eye. Antenna: 11-segmented; antennomere III longest; apical 5 antennomeres thick- ened. Pronotum: Strongly transverse, 2X as wide as long, convex; lateral margin bisin- uate, tooth present in middle of lateral mar- gin; disc smooth, strongly punctate lateral- ly; basal impression present. Scutellum: Quadrate. Elytron: Much wider than pron- otum; short-oval; lateral margin indented behind humerus; lateral and apical margins flattened, finely, sparsely dentate; humerus prominent; punctures large, shallow; inter- val 8 costate on apical %; suture elevated. 767 Legs: Small tooth present before each tarsal claw. Host plants.—Unknown. Described species.—2. Range.—Brazil. Odontota Chevrolat (Fig. 84) Odontota Chevrolat 1837: 388. Type spe- cies: Hispa scapularis Olivier (designat- ed by Butte 1968c). Description.—Wedged-shaped or oblong. Head: Slightly wider than long; vertex tri- sulcate, area between ocular and medial sul- ci finely rugopunctate; frontal ridge cristu- late, either elongate or diamond-shaped; clypeus strongly transverse, prominently raised, base subangulate. Antenna: |1-seg- mented; basal antennomere incrassate. Pronotum: Transverse, generally widest medially; lateral margin angulate in middle, bisinuate; dorsum transversely convex, bas- al depression present or absent. Elytron: Oblong, gradually but slightly dilated from base to apex; apex regularly rounded; ex- terior apical angle rectangulate; lateral mar- gin serrulate or not; with 10 rows of punc- tures plus scutellar row. Venter: Abdominal sternite 5 generally truncate, female with small irregular patch of setae on center el- evation. Host plants.—Amorpha, Amphicarpaea, Apios, Glycine, Meibomia, Pueraria, Robi- nia, Sophora, Tephrosia (Fabaceae) (Jolivet and Hawkeswood 1995). Described species.—7. Key to species: Butte (1968c), North America. Range.—Canada to United States. Oxychalepus Uhmann (Fig. 85) Oxychalepus Uhmann 1938a: 438. Type species: Odontota proxima Guérin-Meéne- ville (by original designation). Description.—Head: Deep medial sulcus present; slightly constricted behind eyes; projection present at base of antenna. An- tenna: 11-segmented; antennomere XI 768 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON iS STAMPS THEREIN ee ‘ees | ; : | : | | » \ | 3 | x : P ; : 788 : | ix RETR AMET OL AO LS ARO ie Figs. 84—92. Dorsal habitus. 84, Odontota scapularis (Olivier). 85, Oxychalepus sp. 86, Oxyroplata sp. 87, Pentispa melanura Chapuis. 88, Physocoryna scabra Guérin. 89, Platocthispa emorsitans (Baly). 90, Probaenia sp., profemur. 91, Probaenia sp. 92, Spaethispa pulchella (Suffrian). VOLUME 104, NUMBER 3 sharply pointed at apex; III to XI laterally compressed; suture between antennomeres VII to XI very weak. Pronotum: Trapezoi- dal, widest at base; transverse basal im- pression present. Elytron: Lateral margin dentate; apical margin strongly dentate; with 10 rows of punctures plus scutellar row; 4 costae, 1, 2, and 4 complete, 3 vis- ible at humerus and apex. Host plants.—Canavalia, Cymbosema, Dioclea, Mucuna, Phaseolus (Fabaceae); Inga (Mimosaceae); Cassia (Caesalpini- aceae); Solanum (Solanaceae); Heliconia (Heliconiaceae); Flagellaria (Flagelari- aceae); Cocos (Arecaceae); Freycinetia, Pandanus (Pandanaceae); Pleomele (Aga- vaceae) (Jolivet and Hawkeswood 1995). Described species.—12. Key to species: Staines (1996b), Nicaragua; Ramos (1998), Brazil. Range.—Mexico to Argentina. Oxyroplata Uhmann (Fig. 86) Oxyroplata Uhmann 1937b: 454 (unavail- able name, type species not designated). Oxyroplata Uhmann 1940b: 143. Type spe- cies: Uroplata bellicosa Baly (by original designation). . Description.—Head: Medial sulcus pre- sent; frons strongly projecting. Antenna: 8- segmented, compressed, antennomeres sharply pointed; antennomere I subglobular, slightly compressed laterally; II transverse, shortest; HI large, laterally compressed, al- most as long as IV and V combined; IV to VII transverse, decreasing in width and length; VIII elongate, acutely pointed at apex. Pronotum: Longer than wide; widest at base; tooth present in anterior angle; bas- al impression present. Scutellum: Quadrate. Elytron: Lateral margin dentate, margined; apical margin laminate, strongly dentate, subcordate-emarginate; with 10 rows of punctures; 4 costae, 2 and 4 strongly ele- vated, | complete, 3 visible at base and apex. Host plants.—Unknown. 769 Described species.—4. Range.—Costa Rica to Peru. Parvispa Uhmann Parvispa Uhmann 1940b: 143. Type spe- cles: Parvispa marmorata Uhmann (by monotypy ). Description.—Head: Medial sulcus deep, long; longitudinal carina present on each side of medial sulcus; vertex alutaceous; row of deep punctures present around eye. Antenna: 7-segmented; antennomeres I and Il subglobular; HI transverse; [IV to VI as long as wide; VII hirsute club. Pronotum: As wide as long; margined on all sides; lat- eral margin bisinuate; longitudinal callus present; transverse basal depression present; large deep punctures present over entire surface. Scutellum: Quadrate, alutaceous. Elytron: Apex conjointly rounded; with 8 rows of punctures plus scutellar row; tri- costate; costae | and 2 united on apical fifth; costa 2 not reaching union of | and 3; costae carinate, uniform in height, no de- pressions or interruptions; sutural margin elevated. Legs: Tarsomeres 3 deeply bi- lobed; 4 barely projecting beyond lobes of 3. Host plant.—Unknown. Described species.—1. Range.—Bolivia. Pentispa Chapuis (Fig. 87) Pentispa Chapuis 1875: 322. Type species: Uroplata melanura Chapuis (designated by Uhmann 1957b). Penthispa Weise 1911: 52 [unjustified emendation]. Description.—Head: Small, globose; with 3 to 5 longitudinal sulci on vertex; eye slightly convex, large. Antenna: Short, ro- bust, 8-segmented, apical 2 antennomeres thickened; antennomere I transverse, large; II transverse, smaller than I; III cylindrical, pitted; IV transverse, pitted; V to VII trans- verse, with setae; VIII elongate, setose, pointed at apex. Pronotum: Wider than long, widest at base, slightly constricted in 770 middle; basal impression present; densely punctate. Scutellum: Subquadrate, trans- verse or square. Elytron: Oblong, slightly expanding apically; exterior apical angle rounded; lateral margin dentate; apical mar- gin strongly dentate; with 8 rows of punc- tures, scutellar row present or absent; tri- costate. Venter: Metasternum rounded an- teriorly, slightly convex. Legs: Short, ro- bust. Host plants.—Colea, Pithecoctenium (Bignoniaceae); Paullinia, Serjania (Sap- indaceae); Baccharis, Clibadium, Elephan- topus, Eupatorium, (Asteraceae); Malpighia (Malpighiaceae); Chusquea (Poaceae) (Jolivet and Hawkes- wood 1995). Described species.—26. Key to species: Staines (1996b), Nicaragua. Range.—Southern United States to Peru. Verbesina, Vernonia Physocoryna Guérin-Méneville (Fig. 88) Physocoryna Guérin-Méneville 1844: 279. Type species: Physocoryna scabra Guér- in-Méneville (by monotypy). Description.—Body elongate, attenuate. Head: Small, medial sulcus absent; vertex with triangular depression. Antenna: Very short, 7-segmented; antennomeres I to III moderate, similar in size, as wide as long or III slightly longer than I or II; ['V—VI transverse; VII much wider, oval, with whorls of bristles, hirsute. Pronotum: Transverse; tooth present in anterior angle; tooth present on lateral margin; with irreg- ular pits. Elytron: Parallel-sided, enlarged in exterior apical angle; lateral and apical margins serrate; apex almost truncate; sur- face with strong, confused punctures; punc- ture row | visible on basal %, interrupted by tubercles; elytral sculpturing symmetri- cal. Venter: Prosternum broad. Legs: Short; tarsi wide, tarsomeres | short, 2 twice as large as 1, 3 as long as 1 and 2 combined, 4 slightly exceeds lobes of 3. Host plants.—Canavalia, Cymbosema, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Dioclea, Phaseolus (Fabaceae); Stimato- phyllum (Malpighiaceae) (Staines 1999). Described species.—3. Key to species: Staines (1999). Range.—Nicaragua to Argentina. Platocthispa Uhmann (Fig. 89) Platocthispa Uhmann 1939: 332 (unavail- able name, type species not designated). Platocthispa Uhmann 1940b: 144. Type species: Odontota gregorii Chapuis (by original designation). Description.—Body flattened. Head: 3 parallel sulci between eyes; frons project- ing. Antenna: 8-segmented, antennomeres transverse, apical antennomeres thickened, reaches to middle of elytra; antennomere I slightly compressed laterally; Il shorter and narrower than I; III longer than IV and V, compressed laterally, tooth-like projection on outer apical margin; IV to VI com- pressed laterally, ['V with small tooth; VII compressed laterally; VIII elongate, as long as V to VII combined. Pronotum: Wider than long; lateral margin evenly convergent from base to apex, margined laterally; tooth present in anterior angle; basal impression present; basal margin bisinuate; longitudi- nal sulcus present on disc. Scutellum: Quadrate. Elytron: Lateral margin dentate; apical margin more dentate, laminate; apex subcordate-emarginate; exterior apical an- gle rounded; with 8 rows of punctures, scu- tellar row absent; tricostate, costa 1 most elevated, 3 extends from humerus to middle of elytra. Venter: With 5 visible abdominal sterna. Host plants.—Cassia (Caesalpiniaceae) (Jolivet and Hawkeswood 1995); Piper (Pi- peraceae); Costus (Zingiberaceae); Cala- thea (Marantaceae); Ochroma (Bombaceae) (Hespenheide and Dang 1999). Described species.—7. Range.—Southern United States to Peru. VOLUME 104, NUMBER 3 Probaenia Weise (Figs. 90-91) Probaenia Weise 1904: 447. Type species: Odontota crenata Blanchard (designated by Monros and Viana 1947). Description.—Body elongate, rectangu- lar. Head: Eye not prominent; clypeus tri- angular, smooth, edges margined, medial carina present. Antenna: 8-segmented. Pronotum: Trapezoidal, at base as wide as or slightly narrower than elytra, strongly narrowed behind anterior margin, conical. Elytron: Apical and lateral margins dentate; flat triangular lamina present in exterior apical angle; with 10 rows of punctures plus scutellar row; 4 costae; costae 3 and 4 unit- ed at apex. Legs: Profemur with | to 3 teeth on inner margin. Host plants.—Verbena (Verbenaceae); Igna (Mimosaceae); Rolandra, Verbesina, Vernonia (Asteraceae) (Jolivet and Haw- keswood 1995); Mikania, Piptocarpha (As- teraceae); Arrabidaea (Bignoniaceae) (Hes- penheide and Dang 1999). Described species.—45. Range.—Nicaragua to Argentina. Spaethispa Uhmann (Fig. 92) Spaethispa Uhmann 1939: 333. Type spe- cies: Uroplata pulchella Suffrian (by monotypy). Description.—Head: Vertex with 3 lon- gitudinal sulci; clypeus transverse, with longitudinal lines. Antenna: 8-segmented; antennomere I subglobose; II transverse; III cylindrical; IV to VII transverse, wider than preceding; VII elongate, truncate at apex. Pronotum: Wider than long; margined lat- erally; tooth present in anterior angle; densely, coarsely punctate. Scutellum: Tri- angular. Elytron: Lateral margin dentate; apical margin more dentate; with 10 rows of punctures, additional puncture row pre- sent between interspace 3 and 4 behind middle; tricostate, costae weakly produced, sutural margin carinate. Host plant.—Unknown. Described species.—1. Range.—Cuba. Stenopodius Horn (Figs. 93-94) Stenopodius Horn 1883: 301. Type species: Stenopodius flavidus Horn (by monoty- py). Description.—Head: Small; vertex not prominent; frons vertical; eye large, oval, facets numerous. Antenna: Clavate, short, extends a short way beyond apical margin of pronotum, 8-segmented; antennomere VIII oval, pointed at apex. Pronotum: Wid- er than long; in most species apex more or less prolonged (lobed) at middle and sinu- ate at each side; base wider than apex, broadly lobed, sinuate laterally, overlapping base of elytron, except in middle. Scutel- lum: Subquadrate; apex truncate, feebly ar- cuate or slightly emarginate. Elytron: Ob- long-oval; about 2 longer than wide; par- allel-sided and broadly rounded in apical 4; lateral and apical margins more or less serrulate; 8 or 10 rows of large, circular punctures. Legs: Glabrous; metafemur slightly longer than tibia; tarsomeres not di- lated, with pale setae beneath, first 3. tar- someres subequal, 3 emarginate above, 4 as long as | to 3 combined; claws feebly ar- cuate. Host plants.—Al/thaea, Malva, Sphaer- alcea (Malvaceae) (Jolivet and Hawkes- wood 1995). Described species.—7. Key to species: Blaisdell (1939), North America. Range.—Canada to Mexico. Sternocthispa Uhmann (Figs. 95—96) Sternocthispa Uhmann 1938b: 113. Type species: Sternocthispa — gracillima Uhmann (by monotypy). Description.—Head: Trisulcate; micro- punctate. Antenna: 7-segmented; anten- nomere I transverse; II and II cylindrical, subequal in length; IV to VI transverse; VII 772 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON guinitedos 90: 99 Sdaoores Figs. 93-101. Dorsal habitus. 93, Stenopodius flavidus Horn, tarsi. 94, Stenopodius flavidus. 95, Sternocth- ispa gracillima Uhmann, prosternum. 96, Sternocthispa gracillima. 97, Sternolispa nigrohumeralis (Pic). 98, Sternostena sp. 99, Sternostenoides daguerrei Monrés and Viana. 100, Stethispa sp. 101, Sumitrosus rosea (Weber). VOLUME 104, NUMBER 3 enlarged, hirsute club, nearly as long as | to VI combined, pointed at apex. Prono- tum: As wide as long, convex; lateral mar- gin sinuate; tooth present in anterior angle; medial carina present; with large, shallow punctures; surface distinctly micropunctate; basal impression present. Scutellum: Quad- rate; micropunctate. Elytron: At base much wider than pronotum, elongate, parallel-sid- ed; lateral margin serrate; apical margin dentate, laminate; exterior apical angle rounded; with 8 rows of punctures plus scu- tellar row; tricostate; costa | more elevated on apical %; 2 and 3 elevated for entire length. Legs: Apical tarsomeres with two claws; small tooth present at base of each tarsal claw; femora punctate. Venter: Pros- ternum produced, partially concealing mouth; 5 visible abdominal sterna. Host plant.—Unknown. Described species.—1. Range.—Brazil. Sternoplispa Uhmann (Fre. 97) Sternoplispa Uhmann 1935d: 237 (unavail- able name, type species not designated). Sternoplispa Uhmann 1940b: 144. Type species: Sternoplispa triformis Uhmann (by original designation). Description.—Head: Vertex with small triangular depression, impunctate; medial sulcus present. Antenna: 10-segmented; an- tennomere I subglobose; II and III cylin- drical; IV to VI transverse, narrower than preceding; VII to [X transverse, wider than preceding; X pointed at apex. Pronotum: Longer than wide, convex; basal impression present. Scutellum: Rounded at apex. Ely- tron: Lateral margin dentate; triangular tooth present on external apical angle; with 8 rows of punctures plus scutellar row; tri- costate. Venter: Prosternum projecting for- ward, with two blunt teeth present on an- terior margin. Host plants.—Unknown. Described species.—7. Key to species: Uhmann (1948). Range.—Brazil to Argentina. Sternostena Weise (Fig. 98) Sternostena Weise 1910: 120. Type species: Charistena basalis Baly (designated by Monros and Viana 1947). Description.—Body elongate, almost cy- lindrical. Head: Small, elongate; interocular area with some markings; antenna inserted into anterior of head. Antenna: 11-seg- mented; antennomere I robust, longer than wide; II longer, more slender than I; III lon- gest, more slender than II; [V to VI similar, shorter than Il; VII to XI somewhat thick- ened, more closely united than others. Pronotum: Slightly longer than wide; lat- eral margin curved. Scutellum: Quadrate, angle rounded. Elytron: Elongate, subpar- allel; apical denticulation irregular; with 8 rows of punctures plus scutellar row; tri- costate. Legs: Short, relatively robust; me- sotibia curved. Venter: Prosternum project- ing forward, covering part of mouth. Host plant.—Paspalum (Poaceae) (Joliv- et and Hawkeswood 1995). Described species.—7. Key to species: Monros and Viana (1947), Argentina. Range.—Costa Rica to Argentina. Sternostenoides Monros and Viana (Fig. 99) Sternostenoides Monros and Viana 1947: 267. Type species: Sternostenoides da- guerrei Monros and Viana (by monoty- Py). Description.—Body elongate, subcy- lindrical. Head: Medial sulcus faint; frons punctate; clypeus finely punctate, with lon- gitudinal carina. Antenna: 10-segmented; antennomeres I and II similar, wider than the next. three; Il to,.V lengthened; . VI shorter and wider than V; VII to X thicker than rest, forming a poorly defined club. Pronotum: Narrower than elytra, cylindri- cal; parallel-sided, slightly elongate; irreg- ularly punctate. Scutellum: Quadrangular. Elytron: Lateral and apical margins finely 774 dentate; apex conjointly rounded; with 8 rows of punctures plus scutellar row; tri- costate, united on apical 1/5. Legs: Meso- tibia strongly curved; mesofemur with short spine. Venter: Prosternum with short me- dian plate on anterior margin. Host plant.—Unknown. Described species.—1. Range.—Argentina. Stethispa Baly (Fig. 100) Stethispa Baly 1864: 265. Type species: Stethispa bonvouloirii Baly (by original designation). Description.—Body elongate. Head: Clypeus transverse, rarely triangular, fre- quently expanding to base of antenna. An- tenna: Short, 1 1-segmented, filiform; apical 5 antennomeres thickened, longer than pre- ceding. Pronotum: Conical or rounded lat- erally; narrowing anteriorly. Scutellum: Transverse, narrowing behind, rounded at apex. Elytron: Elongate, rectangular; hu- merus with outward directed spine; strongly pointed tooth present on exterior apical an- gle; with 10 rows of punctures, scutellar row present or absent; 4 costae. Venter: Su- ture between first 2 abdominal sterna ob- scure in middle. Legs: Tarsal claws diver- gent. Host plants.—Coccoloba, Ruprechtia (Polygonaceae); Aristolochia (Aristolochi- aceae) (Jolivet and Hawkeswood 1995). Described species.—18. Key to species: Monros and Viana (1947), Argentina. Range.—Costa Rica to Argentina. Sumitrosis Butte (Fig. 101) Sumitrosis Butte 1969: 13. Replacement name for Anoplitis Chapuis 1875: 316 (preoccupied by Anoplitis Kirby 1837). Type species: Hispa rosea Weber (by original designation). Description.—Head: Slightly wider than long; vertex finely granulose with deep me- dial sulcus; frontal carina feeble, joining PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON clypeal base; clypeus feebly transverse. An- tenna: 11-segmented. Pronotum: trans- verse; lateral margin obtusely subangulate at middle and feebly narrowing towards apex and obliquely more or less so towards base; transversely convex; basal impression present or absent. Elytron: Elongate-ovate; lateral margin serrulate or not; apex con- jointly, distinctly rounded; with 8 rows of punctures plus scutellar row; tricostate. Host plants.—Amorpha, Amphicarpaea, Cajanus, Canavalia, Desmodium, Dolichos, Glycine, Lespedeza, Meibomia, Phaseolus, Pueraria, Robinia, Strophostyles (Faba- ceae); Potentilla (Rosaceae); Cassia (Cae- salpiniaceae); Celastrus (Celastraceae); As- ter, Eupatorium, Helianthus, Rudbeckia, Solidago, Veronia, Wedelia (Asteraceae); Cyrilla (Cyrillaceae); Laportea (Urtica- ceae); Chenopodium (Chenopodiaceae); Bambusa, Chusquea, Lasiacis (Poaceae); Quercus (Fagaceae) (Jolivet and Hawkes- wood 1995): Heliconia (Musaceae) (Hes- penheide and Dang 1999). Described species.—58. Key to species: Butte (1969), North America; Staines (1990), Argentina; Staines (1996b), Nica- ragua. Range.—Canada to Argentina. Temnochalepus Uhmann (Fig. 102) Temnochalepus Uhmann 1935a: 56. Type species: Odontota lugubris Chapuis (by original designation). Description.—Head: Clypeus hardly granulose, with medial carina which is pro- longed into a deep interocular sulcus; sharp postocular constriction present; projection present at base of antenna. Antenna: 11- segmented; antennomeres I to II cylindrical, Il shorter and narrower than I; III longer than I or I; IV to VI transverse, decreasing in length; VII to X transverse, slightly com- pressed laterally; XI rounded at apex. Pron- otum: Wider than long, widest at base; small tooth present in anterior angle; trans- verse basal impression present; densely VOLUME 104, NUMBER 3 plEircenemees Figs. 102-107. F : y Dorsal habitus. 102, Temnochalepus imitans Uhmann. 103, Temnocthispa truncata (F). 104, Uroplata sp. 105, Xenochalepus (s. str.) sp. 106, Xenochalepus (Neochalepus) medius (Chapuis). 107, Hispoleptis diluta (Guérin). punctate. Elytron: Lateral margin smooth; apical margin scarcely denticulate; with 10 rows of punctures plus scutellar row; 4 cos- tae, 1, 2, and 4 well developed; costa 3 vis- ible at humerus and apex or totally absent. Host plants.—Panicum, Pharus (Po- aceae); Commelina (Commelinaceae) (Jo- livet and Hawkeswood 1995). Described species.—4. Key to species: Monros and Viana (1947), Argentina. Range.—Brazil to Argentina. Temnocthispa Uhmann (Fig. 103) Temnocthispa Uhmann 1939: 331 (unavail- able name, type species not designated). Temnocthispa Uhmann 1940b: 144. Type species: Hispa truncata Fabricius (by original designation). Description.—Head: Medial sulcus pre- sent; sulcus present on inner margin of eye. Antenna: Antennomere I subglobose, slightly compressed laterally; Il to VII transverse, III longer than II; VIII elongate club, pointed at apex. Pronotum: Much nar- rower than base of elytra; slightly constrict- ed behind head; tooth present in anterior an- gle; lateral margin curved. Scutellum: Quadrate. Elytron: Lateral and apical mar- gins dentate; with 8 regular puncture rows plus scutellar row; additional puncture row 7716 present on apical %; tricostate, costae 1 and 2 more elevated than 3. Legs: Femora or pro- and mesofemora toothed; tarsal claws without tooth. Host plants—Unknown. Described species.—3. Range.—French Guyana to Ecuador. Uroplata Chevrolat (Fig. 104) Uroplata Chevrolat 1837: 389. Type spe- cies: Uroplata mucronata Olivier (des- ignated by White 1988). Codiohispa Maulik 1930: 48. Type species: Codiohispa anonicola Maulik (by mon- otypy); Uhmann 1957b (synonymy). Mimuroplata Pic 1933: 389. Type species: Mimuroplata trregularis Pic (by mono- typy); Monr6és and Viana 1947 (synony- my). Plicatopalpa Pic 1932: 29. Type species: Plicatopalpa irregularis Pic (by mono- typy). New synonomy. Description—Head: Oblong, rounded; vertex smooth or with no sulci. Antenna: 8- segmented; antennomeres VII and VIII with whorls of golden setae. Pronotum: Varying in form, at base narrower than elytra; lateral margin more or less rounded; disc frequent- ly with 3 longitudinal vittae. Scutellum: Transverse. Elytron: With 10 rows of punc- tures, scutellar row present or absent; 4 cos- tae, 3 and 4 united on apex; 3 interrupted. Legs: Moderately long; protibia occasion- ally with a tooth; empodium present be- tween tarsal claws. Host plants.—Annona, Rollinia (Annon- aceae); Baccharis, Clibadium, Elephanto- pus, Eupatorium, Rolandra, Vernonia, Wedelia (Asteraceae); Banisteria, Malpigh- ia (Malpighiaceae); Lantana, Lippa, Ver- bena (Verbenaceae); Caesarea (Viviana- ceae); Arrabidaea, Bignonia, Colea, Pithe- coctenium (Bignoniaceae); Calopogonium (Fabaceae); Aristolochia (Aristolochi- aceae); Althaea, Sida (Malvaceae); Ocotea (Lauraceae); Panicum (Poaceae); Acantha- ceae (Jolivet and Hawkeswood 1995); Inga PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Mimosaceae); Gouania (Rhamnaceae) (Hespenheide and Dang 1999). Described species.—88. Key to species: Staines (1996b), Nicaragua. Range.—Mexico to Argentina. Xenochalepus Weise (Fig. 105) Xenochalepus Weise 1910: 136. Type spe- cies: Odontota omogera Crotch (desig- nated by Butte 1968a). Hemichalepus Spaeth 1937: 147 (unavail- able name, type species not designated). Hemichalepus Uhmann 1957b: 94. Type species: Odontota haroldi Chapuis (by original designation); Staines and Riley 1994 (synonymy). Description.—Head: Slightly wider than long; vertex sulcate; longitudinal carina be- tween antenna reaching base of clypeus; clypeus large; labrum prominent, wider than long; outer margin of mandible rather broadly and evenly arcuate; eye convex, elongate-oval. Antenna: 11-segmented; an- tennomere I incrassate. Pronotum: Wider than long; lateral margin bisinuate, angulate at middle; transversely convex; slight basal impression with transverse carina present. Elytron: Elongate-oval; apex conjointly rounded, deeply subquadrate-emarginate at suture; lateral and apical margins serrulate; with 10 rows of punctures plus scutellar row. Legs: Last tarsomeres produced _ be- tween bases of claws into a narrow trian- gular, truncate process. Venter: Abdomen sparsely micropunctate; apex of sternite 5 slightly emarginate; generally sternite 5 of female with small irregular patch of setae on both sides of central elevation, in male patch of setae scarcely perceptible. Host plants.—Theobroma (Sterculi- aceae); Bambusa, Panicum, Saccharum, Zea (Poaceae); Canavalia, Cymbosema, Dioclea, Dolichos, Faba, Glycine, Lathy- rus, Mucuna, Nissolia, Phaseolus, Vigna (Fabaceae); Inga (Mimosaceae); Urera (Ur- ticaceae); Schubertia (Malvaceae); Prunus (Rosaceae); Celtis (Ulmaceae) (Jolivet and VOLUME 104, NUMBER 3 Hawkeswood 1995); Cecropia, Coussapoa, Pourouma (Cecropiaceae); Anthurium (Ar- aceae) (Hespenheide and Dang 1999). Described species.—26. Key to species: Uhmann (1938a), key to species groups; Monros and Viana (1947), Argentina; Butte (1968a), North America; Staines (1996b), Nicaragua. Range.—Southern United States to Ar- gentina. Xenochalepus (Neochalepus Staines and Riley) (Fig. 106) Xenochalepus (Neochalepus) Staines and Riley 1994: 219. Type species: Odontota medius Chapuis (by original designation). Description.—As in Xenochalepus (s. str.) except differing in development of ely- tral intervals and shape of mesosternum as follows: Elytron with intervals 2, 4, and 8 costate, other intervals not developed or, in some species, interval 6 raised near base and again apically; mesosternum between mesocoxae Clearly transverse. Host plants.—Olyra, Oryza (Poaceae); Lathyrus, Mucuna, Nissolia, Phaseolus, Rox binia, Wisteria (Fabaceae); Inga (Mimosa- ceae); Bauhinia (Caesalpiniaceae); [pomoea (Convolvulaceae) (Uhmann 1938a, Monros and Viana 1947). Described species.—64. Key to species: Spaeth (1937). Range.—Southern United States to Ar- gentina. TRIBE HISPOLEPTINI CHAPUIS 1875: 283 Type genus.—Hispoleptis Baly. Description.—Head: Clypeus obscure. Antenna: 11-segmented; filiform, apical 4 antennomeres similar to preceding; at least apical portion compressed laterally. Pron- otum: Nearly quadrate, narrowing apically. Elytron: Elongate, subparallel; puncture rows strongly impressed. Tig Hispoleptis Baly (Fig. 107) Hispoleptis Baly 1864: 262. Type species: Promecotheca diluta Guérin-Méneville (by original designation). Description.—Moderately elongate, with patterns of black and yellow or reddish. Head: Vertex impunctate; interantennal ca- rina distinct, bluntly pointed; eye trans- verse, narrowed behind. Antenna: Filitorm, more or less compressed apically; anten- nomere I subglobular; I] very small; II lon- gest; IV to X progressively decreasing in length; XI longer than X; III to XI with fine longitudinal carinae; apical 4 antennomeres with deep longitudinal depression on outer side. Pronotum: Almost as long as wide, more or less narrowing apically; at base al- most straight, margined and _ transversely impressed; margined laterally. Scutellum: Impunctate. Elytron: At humeri wider than pronotum, elongate, subparallel; deeply punctate; dentate apically; no costae. Legs: Procoxae closer together than meso- or me- tacoxae; mesotrochanter with sharp projec- tion; femora progressively less incrassate from anterior to posterior; protibia with in- terior tuft of setae; mesotibia curved, with inner apical tooth; pro- and metatibiae straight, without teeth; first tarsomere of prolegs asymmetrical and much wider than on other tarsi; tarsomeres 2 and 3 with lon- gitudinal medial depression on all legs. Venter: Prosternum projecting in front in middle, transversely concave between cox- ae; abdominal sterna | and 2 connate, su- ture distinct; all sterna with impressions lat- erally. Host plants.—Cocos, Elaeis (Arecaceae) (Jolivet and Hawkeswood 1995). Described species.—4. Key to species: Aslam (1965). Range.—French Guyana to Brazil. ACKNOWLEDGMENTS The following institutions and individu- als have lent material for this study, the as- sistance of the curators responsible is grate- 778 fully acknowledged: American Museum of Natural History, L. H. Herman; Bishop Mu- seum, G. A. Samuelson; The Natural His- tory Museum, S. L. Shute; California Acad- emy of Sciences, D. H. Kavanaugh; Uni- versity of California, Berkeley, C. B. Barr; Carnegie Museum of Natural History, R. L. Davidson; Universidad de Costa Rica, H. Lezama; Deutsche Entomologisches Insti- tut, L. Zerche; Florida State Collection of Arthropods, M. C. Thomas; Manchester Museum, C. Johnson; Muséum National d’Histoire Naturelle, Paris, N. Berti; Insti- tuto Nacional de Biodiversidad, A. Solis; University of Kansas, R. W. Brooks; Mu- seum of Comparative Zoology, D. G. Furth; Museum fiir Naturkunde de Humboldt- Universitat, EF Hieke; Texas A&M Univer- sity, E. G. Riley; National Museum of Nat- ural History, Smithsonian Institution, R. E. White; Universidad Central de Venezuela, V. Savini. R. W. Flowers (Florida A&M Universi- ty), E. G. Riley (Texas A&M University), and A. S. Konstantinov (USDA, Systematic Entomology Laboratory) commented on earlier drafts of this manuscript. Funding for a trip to The Natural History Museum (London) was provided by the Thomas Lin- coln Casey Fund, and a trip to the Man- chester Museum was provided by the Ernst Mayr Fund. S. L. Staines provided technical and editorial assistance. S. W. Lingafelter, USDA, Systematic Entomology Laborato- ry, was of great assistance in the digital photography and setting up the plates. Costs for publishing this work were provided from a World Bank grant to the Instituto Nacional de Biodiversidad, Santa Domingo, Costa Rica. LITERATURE CITED Aslam, N. A. 1965. On Hispoleptis Baly (Coleoptera, Hispidae) and /matidium F. (Coleoptera, Cassidi- dae). 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E. 1988. The genus Uroplata, type-species and authorship (Coleoptera: Chrysomelidae). Pro- 783 ceedings of the Entomological Society of Wash- ington 83: 713-715. INDEX Acanthispa'Chapuis® oea-nisnteo. ce ariacranace a. 63s ae 752 AGGNthOdES TBAlY. sicjctettecte mts Tapstae orotate. (0 cinta et sexes TD2 AGCCNLTOPLETG Aly sateen sea senescence peeieneoeir 746 Acritispa Ubimanm e< sass. merece ea ace ais 752 Adalurnus Niailike so cectasec ce eee cath ee eee 744 Aivathispa WESC, Sandee sso eee eee da2 Alumni Chapuis;csicc.5: eos eeeeeeborente saath ee 742 AMT RUS TaD TCI S tase sect ise sees etic eerste ecacce 744 AMUDUID GIG Taro de eeaatee cc csiqe tet eee oeekeeiacaaee TEA Amplipalpimi: WeiSeu green ceteccreeese eyes aeseece 727 Antsochalepus: Ulmanm) 2225220 2-40senereee yee cies 752 Anisostena (Apostena) States. 's.55.-te.tis- 2.42200 754 Anisostena (Neostena) Monros and Viana ....... 754 AIUSOSTCNIG NV CISCi th trope see ere tenets same eee eee Oeane 754 ATLODIULLS KOTDY ) a2 aca. cence case se eeeneeee eee 758 Anoplitiss Chapuils tosses caese tees 2 oeeeae spaces 774 AvrescinisChapwis) geecs sees ceed eee eomeeeece eee eee 741 ATESCUS: PEILY T2a628 sat aaisnaiaeneie cetonereise a naeacetiee 741 Aslamidium (Neoaslamidium) Borowiec .......... 731 AISIGMNI din BOTO WICC w ce aaeeice sieertsecos cies elect: 730 BGLIOSUS! WEISG: seagate manetianc ieee eee e eee eee 754 Bothrispa Olbmanny ee aen-ast ors -ceereoee ease aces 755 Brachycoryna Guérin-Méneville ................... qa5 Bruchia WEISO* 3. Fhe Fossa Soomro apa ont ein sate ohte (BS Bruchiella WeISCs a os oeieceaseche sects oe aoee eee 756 GCalliaspissDejcamiyn-cresudis erect ae aeceaoe Aces 731 Callaspites(@haplis secs seg teense neelaaate eaeiter 729 GalyptocephalastChevrolat 2... csasc-cineoeeteccasle IZ Carinispa Uhmann ........ shiiciesessie cose astee eee sass 756 Cephalodonta Chevrolat\.2o. c.0seaeore ade eee een 748 CephalodontinteWeise 325405 tess ee ee tee eeeaee 746 Cephalodontites (Chapuisy <...a22 senses daccmer 746, 749 Gephaloleia Chevrolat Fixssecca coos tase eereasae 731 ECephalolenmnmeChapwis: ay.c-s>-eeucseeedtoosciaacscee: 729 G@halepini? Wises Conntena acim armen acstae sips ener 749 Chalepotatus (Macrochalepus) Pic .............+.. 748 Chalepotatus WISE \..255:286 aa. aceon dad. Chalepsipg Whmantn oa. ss aceuecneroccaseeetosaa awh Chalepus Thumbene: 25 stae5.24556 seca ne se sere 758 GHOTIS DE MB AWG ise siape cays fo Satin ce 6 ss Se EASA SeIISO RE WEA CharistenarBaly ss. cnces: ss isin. shes ho aeee eee eee oi 758 GChEeiriSPGMBalye .eae<% ahh 5 dah 8 Hee wAS EN neekitedae 746 ChelobasiscGray e256. tenes te Gh gece de densttae nee 741 Ghirispa;Gemminger and! Harold...) 3.242.0 746 (GUCTNK OCR) 11S MRR COnAD Cac IER eT ODO RD OREe 732 Glinocarispa WRMANM 2.c.s.26.c.0eccemeyanee access 758 GnetispagNiaulike - 32 s5ic0, ose arecisenee sed erat ee eee 759 Codiohispa Nua soe cree cove sroe mi sicieyeshetelere sieasis vers 776 GCoraliomela JaCOOSOM po 464 25 ascss are iotea tare oe mrosclcerys 744 Gonynispa Oman erocnccjscsiena sean csieleee see tocee (By) CV ASPEAOMISP A WESC tas eaoeie e vies avila siete esse eter ei si- 139 GYAN ASDEUSPNVCISE aoe cize aoe ain setters inseinle Sos otteinsateoistateyeiaistsls 731 DECATCTIANWEISE! (oie joc te Biateacialetieiciaeis tinea eens em e 761 Delocrania: Gucrimn=Mene Ville: si. 2a 8 ecm yo njteioseiciere 738 784 Delocranimigs pacthipere-ees-ceeee cee ee ease se 737 Demothispa Gemminger and Harold .............. 732 IDA MOWIN OBIS) sa sesooopoedadeansecessseasesseccsons WB2 JIU NARDG HAUG SVOREWN soucqsancasobassceessoonouscooNs 738 Enneachalepus Spaeth .........-.....2:.. Rocce ess 765 Buprionota Guern-Meneville ys... ere 761 IBVERA G2). SEIN. coaancaapangoodoncopodpaasonapsudeosasces 732 1EOSOUN Bah SNEWS) pacepoocanoosonsneAbnacubooaEpasnebos 761 GhyphuroplataWhmanni asec eee ee 761 Goyachalepus Rice ssse ate cls NTA ee re 762 Hemichalepus Spacthy aa4-- seco rece e eel: 776 HemichalepussUhmannt eeeaecee eee este eee 776 Hemisphaerotal@hevrolatwreseee- see eee eee 738 Hemisphaerotas Spacthys. ee. e-e0- eee 738 Hemisphaerotini Monros and Viana .............. 738 Heptachispa Whmanniepeeere seen eee eee 762 JEKG COHORT \MACS cocoonbasusacusnddnoancosnesoodcsNsc 762 ideptatomispa Whmanny aeae--een- eee ee 762 EV eptispar WEIS Cite. estes see tts na ie ee ae ola eG 763 EHeterispalChapuiS wre eco eee eee ene eee 763 ElurmatideliaeAslaniae eer eee eee ee cee 734 Hlimatiditacss pacthe teres -e-cieee eee ieee ce 729 limatidiites! Ghapuisyesee-eeeeeeeecee eee aaa eee ee 729 Eimatidiumbatteile ese nee eee ene See eee 734 uspoleptinit@hapuiser-ssaceeesee escent Wad FHispoleptistBalya ee otsecssscaeecose ee oes ocl ee eel, Homalispay(Xxanthispa)) Baliye. eae 734 LL OMOLUS PGES alyssa serene secret 734 Hoplronotites!Ehapuiss aes. eee tees sete elatciers qT laloonkyaya VSS. so4annooancosaasgaebcecdageestoobace 740 lahyoxewyjormait: \KENISe" 5 Snaooomaceshoncasecsense osaecesE se 740 Imati ciinisbtin ksi rae se ee eee TE) ImatidiumyRavnGiuss ee aseee en occ heron ce 734 MoO MTD INAEWIN, cocasooconaaabanadatooosbaSosseddHr 730 MecistomelavacobSOnweree eee e tee eee eee 745 MielanispatBaly rect nc.cee-ccscccse ene ceo eee 734 MetaxyceravBaly ea nedccssss tees foece os case ees es 763 MicrodontarGhevrolat reeset eeee eee ee eee 748 Microrhopalai@hevrolat e425 ee eee ee 763 Mimoethispar Pics on dice secs asase- Vane ce ese oe 7605 MimuroplatanPicren eetecec tee nee ee 776 Nanocthispa Monros and Viana .................-- 765 NonispaiMauilike 26 aio sscaece seule tits se ae eee oe 765 INymMphares GUSIW CISC Mee eee ee atee eee ere ene eae 742 OGNOSISPGWEISE? Possess es ease eden ne rere 748 Octhispai@hapuis weeaseee teen eee eee ae snag Uo OciocladiscusmMihomsonyee- ee eee eee ee 185 Octotomai@hevrolatys ss peesc eee ees ees 766 OctotomutesiChapuismeeepeee sees eee eee eee 749 OciuroplatatW@hmannieee ee eee eee eee eee 767 OdontispasUhmanniiy-n-4-o-- eee eee eee 767 OdontotaiGhevrolatieeseeos eee eee 767 OcdiopalpaB aly ese een seen reece VAY Oediopalpini Monrés and Viana ................... 2a OCTRISPAABICs: bc Resse ek Sa aS RAM eee 766 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Oxychalepus Uhmannih-aes-- eeeee eee eee ee eee 767 Oxyroplata Whmanny seeee ce eeee eee eee eee 769 Parabaliosus Monros and Viana ...............-.- 754 Parachalepus Balyaeesce eee eee eee 758 Paradecatelia Whmanntis--esssecn ee eee eee 761 Rarnimatidium: Spacthye esses eee eee eee 735 Rarvispas@hmann Ss as-seee eee eee ee eee 769 PemthispanNVeisSe. wc os ecseeeseeen ae eee eee 769 Pentispax@hapuis! 2.4 asgae chee cere eee ee 769 Physocoryna Guérin-Méneville ...................- 770 Physocornyna> Baly” 2...22.0s2cg eee eee ee ee 1S) PlatocthispaWhmann’ 2 e-preseeeeeeer eta 770 Riahvauchenitar Sturm =. --9s seecceeee eee eee 745 PliGatopalparPicy s...s2s seen eee ee eee 776 Poeeilalurnis Vacobsonl eaeeeenee eee eee eee eee 744 RonphyraspissElOpe’ s... eee eee erate eee 738 Porphiyraspitac; Spaeth) D 1S mMests HALT 59) Siti 2, nest #1198; 2 d, nest #1199; 5 3,8 2, nest tl 20030316 41 35 eemesh H202:027 2. purple martin nest #1215, July 23, 1971; 1 3, 1 &, tree swallow nest #1216, July 23, LOTAse tis 1298S Monte? Cristo; Richi €o:; purple martin nest #519, August 10, 1970; 5 6,4 &, mixed with P. sialia, Walton Can- yon, Rich Co., cliff swallow nest #418; | 3,3 2, mixed with P. halli Sabrosky, Ben- nett, and Whitworth. Logan, Cache Co., barn swallow nest #1290, August 14, 1971; 73,6 2, Box Elder Co., violet-green swal- low nest August 2, 1966, collector Ken Ca- pelle. WASHINGTON (19 6, 81 ¢, many puparia): Tacoma area, Pierce Co.; 3 d mixed with 65 P. braueri, house sparrow nest #1955, July 6, 1978; 3 ¢o, 4 &, tree swallow nest #1966 June 15, 1979; 8 males, 15 2, tree swallow nest #2025, June 23, 1990; 6 d, 12 2, purple martin nest #2164, July 30, 1990; 22 ¢, purple martin nest #2165, July 30, 1990; 15 2, purple martin nest #2229, August 5, 1991; 7 puparia, tree swallow nest #2384, August 15, 1993; 85 puparia, purple martin nest #2539, summer 1996; | puparium, tree swallow nest #2557, July 13, 1997; 2 ¢, 3 2, Snohomish, Sno- homish Co., tree swallow nest #2566, Au- gust 1, 1997; 5 2, Bellingham, Whatcom Co., violet-green swallow nest #3751, July 10, 2000; | 2, Bellingham, Whatcom Co., tree swallow nest #4166, August 2, 2000; 2 2, Bellingham, Whatcom Co., tree swallow PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nest #4171, July 30, 2000; 20 &, tree swal- low, nest #4172, July 30, 2000. WYO- MING (2 6,3 2): Gros Ventre River, Jack- son, violet-green swallow nest #448, 7/29/ AO: Specimens examined.—145 d, 228 9, numerous puparia. Distribution.—USA: Idaho, Montana, Oregon, Utah, Washington, Wyoming, Can- ada: British Columbia (Fig. 9). Basis for description of puparia.—Utah slides - 359-2 & 3; 447; 519-A & C; 1198- 1, 2, & 3; Washington slides - 2384, 1-2; 2539, 1—5, 7-10; 2557; 3751, 1-2; 4172, 1-2; Oregon slides - 3741, 1; 4194, 2; 4202, 1 & 3; 4204, 2; Idaho slides - 4044, 1—2. Collection data for these nest numbers can be found under type series information. Hosts.—Purple martin, house sparrow, and many swallow species including: bank, barn, cliff, tree and violet-green. Ecology and biology.—Protocalliphora rugosa was the primary species in the nests of purple martins in northwestern United States and also often found in tree and vi- olet-green swallow nests. It rarely occurred in barn swallow, bank swallow, or cliff swallow nests where P. halli, P. chrysor- rhoea (Meigen) and P. hirundo respectively were the primary parasites. Puparia were usually found wrapped in feathers like P. sialia and P. parorum. Adult specimens of P. rugosa reared from nests in Washington were disproportionately females. Three nests, lot numbers 2165, 2229, and 4172, produced 57 adults, all females, with no males. However, in Utah the sex of adults was about equally male and female (94 males, 107 females). Similar sex ratio dis- tortion has been observed in other insects due to the presence of son-killing or male- killing bacteria like Wolbachia, which kills male hosts of diploid insects early in de- velopment. I have not observed this sex ra- tio distortion in other species of Protocal- liphora, but Wolbachia has been found in all species of Protocalliphora examined to date (E. Baudry, personal communication) [information obtained in writing]. VOLUME 104, NUMBER 3 Etymology.—The specific name is de- rived from the Latin rugosus, which means wrinkled or folded and describes the heavy folds characteristic of the puparia of the species. KEY TO SPECIES The following key will serve to separate males and associated puparia of closely re- lated species with white calypteres and dig- itate surstyli. 1. Frons of male as wide or wider than breadth of 3rd antennal segment. Hyperstigmatal spines of puparia 25,1. or more, posterior ventral spine bandsinot reduced ‘to rear, Sais. $2.6 2 23 8 2 — Frons of male obviously narrower than breadth of 3rd antennal segment. Hyperstigmatal spines of puparia short, usually 20p or less, posterior ventral spine bands significantly reduced to rear. . Preocellar area of male with polished black tri- angle or streak (Figs. 4—6). Dorsal cuticle of puparia with ridges weak or absent, or if pro- nounced, prothoracic fringe 500 or more. .. 3 — Preocellar area of male with dull gray triangle or streak (Figs. 7, 8). Dorsal cuticle with pro- nounced ridges, diameter of prothoracic fringe Bee ce Ci oh ee ote Bete shannoni i) AV (}0 WUROTHESS ty teedeas Sq seeie. 6 BOA Lew Geos Soe 4 3. Puparium with prothoracic fringe diameter of 500 or more, dorsum heavily ridged, frons in male averaging about 1.4 times breadth of 3rd antepnalysesment (BissG)r iia 2. 22 sialia — Puparium with prothoracic fringe about 375 (Fig. 1b), dorsal ridges usually weak (Fig. Ic), frons in male about equal to the breadth of the 3rd antennal segment (Fig. 4). ... bennetti, n. sp. 4. Stigmatal region of puparium with a few broad, irregular folds (Fig. 3a), dorsum with broad, irregular folds (Fig. 3b). Upper portion of para- frontal in male not significantly narrowed (Fig. tS) etc lee Honey ete otdita tok ae? AL ec ee hirundo — Stigmatal region of puparium with narrower, concentric folds (Fig. 2a), dorsum with narrow, regular folds (Fig. 2c). Upper portion of para- frontal in male narrows significantly (Fig. 7). rugosa, N. Sp. ACKNOWLEDGMENTS This study was funded by my company, Whitworth Pest Solutions. I thank all my employees for understanding my preoccu- pation with bird nests and for only occa- sionally complaining about all the fleas, 811 mites, swallow bugs and spiders that visited us while I examined nests. I am especially indebted to Colleen DeLong and Tina Phil- lips of Cornell University, and several hun- dred cooperators of the Cornell Birdhouse Network who provided me with over 2000 bird nests to search for Protocalliphora. | also thank Russ Dawson of the University of Northern BC, Steve Matsuoka, U.S. Geological Survey, Anchorage, Alaska, and Christine Custer, U.S. Geological Survey, LaCrosse, Wisconsin, who provided nests and specimens of P. bennetti. Also thanks to entomologists Karen Needham of the University of British Columbia, Spenser Museum, Wilford Hanson, Utah State Uni- versity, Rich Zack, Washington State Uni- versity, and Norm Woodley, USDA Sys- tematic Entomology Laboratory, Washing- ton, DC, who loaned me specimens needed to complete this study. Finally thanks to Dawn Nelson of Seattle for the drawings and Patrick Craig of Monte Rio California for the slide photos. LITERATURE CITED Bennett, G.F, 1957. Studies on the genus Protocalli- phora (Diptera: Calliphoridae) University of To- ronto, Canada, unpublished Ph.D. dissertation, 194 pp, 33 plates. Dawson, R.D., T.-L. Whitworth and G.R. Bortolotti, 1999. Bird blow flies, Protocalliphora (Diptera: Calliphoridae), in cavity nests of birds in the bo- real forest of Saskatchewan. Canadian Field-Nat- uralist 113 (3): 503-506. Hall, D.G., 1948. The Blowflies of North America. Thomas Say Foundation, Entomological Society of America, Lafayette, Indiana, 477 pp, 51 plates. Sabrosky, C.W., G.E Bennett, and T.L. Whitworth, 1989. Bird blow flies (Protocalliphora) in North America (Diptera: Calliphoridae), with notes on Palearctic species. Smithsonian Institution Press, Wash. D.C. 312 pp. Whitworth, T.L., 1971. A study of the biology of the species of Protocalliphora in the northern Wa- satch Range. unpublished M.S. Thesis, 58 pp. Utah State University, Logan, Utah. ———.. 1977. Host and habitat preferences, life his- tory, pathogenicity and population regulation in species of Protocalliphora Hough (Diptera: Cal- liphoridae). Utah State University, Logan, Utah, Dissertation Abstracts International, B, 37 (10): 4933-B. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 812-813 NOTE First Records of the Sugarcane Pest, Blastobasis graminea Adamski (Lepidoptera: Coleophoridae: Blastobasinae), from Mexico and Central America Although described only recently (Adamski 1999), Blastobasis graminea Adamski has been known as a pest of sug- arcane (Saccharum officinarum Linnaeus; Poaceae) in Colombia and Venezuela for nearly 50 years (e.g., Box 1953, Guagliumi 1962). Cardenas and Hernandez (1988) de- scribed the biology of B. graminea in Co- lombia, and Adamski (1999) presented de- tails of its morphology, along with illustra- tions of the adult, genitalia, larval chaeto- taxy, and damage. Martorell (1976) reported two species of Blastobasinae feed- ing in flower heads of sorghum (Sorghum aethiopicum Hackel Ruprecht ex Stapf.; Po- aceae) in the Virgin Islands, and it is pos- sible that one of them represents B. gra- minea. Unfortunately, these specimens are presumably lost. During a survey of sugarcane fields in central Mexico in February 2001, speci- mens of B. graminea were collected in the states of Veracruz and Jalisco. In Jalisco (Autlan and Casimiro Castillo), larvae of B. graminea were collected from young stalks of sugarcane at ground level, while in Ve- racruz (La Gloria), adults were collected us- ing light-traps and blacklight. We assume that both of these collections represent res- ident pest populations in the sugarcane fields from which they were collected. Co- incidentally, specimens of B. graminea (lar- vae and adults) recently were sent to the Systematic Entomology Laboratory for identification from Costa Rica, where they had been reared from sugarcane. These new records (from Mexico and Costa Rica) sig- nificantly broaden the known geographic range of B. graminea (Fig. 1). At present, it is unknown whether the re- cords from Costa Rica and Mexico represent a recent invasion of this species into Central America, an old, previously undetected in- vasion, or a host switch by native B. gra- minea populations onto cultivated sugar- cane. The fact that B. graminea has been reared from corn (Zea mays Linnaeus; Po- aceae) and sorghum suggests that it has the potential to become a widespread pest throughout the lowlands of northern South America and Central America where these crops are cultivated. However, it seems that adequate time for this scenario to come to fruition has elapsed without the potential re- sult. The fact that B. graminea also has been recorded from Coix lacryma-jobi Linnaeus (Poaceae) and Setaria paniculifera Fournier (Poaceae), two widespread native grasses, suggests that it already may have a broad native distribution in the region and has switched over to cultivated plants opportu- nistically in certain parts of its range. Re- gardless, the potential of B. graminea to be- come a more widespread and economically important pest of sugarcane seems high in light of the information presented herein. Furthermore, we recommend that future sur- vey work be conducted for this insect in sug- arcane producing areas of northern Mexico and southeastern United States where B. graminea 1s not known to occur at present. We acknowledge the following for assis- tance with logistics, access to sugar planta- tions, and/or collecting specimens: Monica C. Vargas, Director, and Luis M. Rosado- Grajales, Campus Veracruz, Colegio de Postgraduados, Mexico; Alejandro Martinez and Ricardo Garcia, Plan de Ayala Sugar Mill, Cd. Valles, San Luis Potosi; Juan Fran- cisco Leal and José Reyes Hernandez, Man- te Sugar Mill, Cd. Mante, Tamaulipas; Joel Hernandez Sanchez and Hugo Valdez, Pan- uco Sugar Mill, Veracruz; Alberto Baez and Jorge Campos, Independencia Sugar Mill, VOLUME 104, NUMBER 3 @ previously recorded localities (Adamsk! 1999) © new locations (persented herein) & possible location (Martore!! 1976) Fig. 1. Known distribution of B. graminea. Closed circles are previously reported locations (Adamski 1999); open circles are new locations (presented herein); triangle is possible location (Martorell 1976). Independencia, Veracruz; and Leopoldo Montero, La Gloria Sugar Mill, La Gloria, Veracruz. We thank Pedro Esteban Diaz, In- stituto Nacional de Ecologia, SEMARNAP. Mexico, for providing Adamski with a col- lecting permit. César Guillén Sanchez and Kenji Nishida, Universidad de Costa Rica, San Pedro, and Eugenie Phillips Rodriguez, Instituto Nacional de Biodiversidad, Santo Domingo, Heredia, Costa Rica, forwarded to us specimens of B. graminea from Costa Rica. Vitor Becker, Planaltina, Brazil, David Smith, Systematic Entomology Laboratory, USDA, Washington, DC, and an anony- mous reviewer provided helpful comments on the manuscript. LITERATURE CITED Adamski, D. 1999. Blastobasis graminea, new species (Lepidoptera: Gelechioidea: Coleophoridae: Blas- tobasinae), a stem borer of sugar cane in Colom- bia and Venezuela. Proceedings of the Entomo- logical Society of Washington 101: 164—174. Box, H. E. 1953. List of sugar-cane insects: A syno- nymic catalogue of the sugar-cane insects and mites of the world, and their insect parasites and predators, arranged systematically. Common- wealth Institute of Entomology, London, 100 pp. Cardenas Duque, L. and Maria del Pilar Hernandez. 1985. Barrenador de la cana azucar en Colombia. Miscelanea, Sociedad Colombiana de Entomolo- gia 1: 12-17. Guagliumi, P. 1962. Las Plagas de cana de aztcar en Venezuela. Ministerio de Agricultura y Cria. Ma- racay, Venezuela. Monografia no. 2.2 partes, 789 Pp. Martorell, L. EK 1976. Annotated food plant catalogue of the insects of Puerto Rico. Agricultural Exper- iment Station, University of Puerto Rico, Depart- ment of Entomology, 303 pp. David Adamski, John W. Brown, System- atic Entomology Laboratory, Agricultural Research Service, U.S. Department of Ag- riculture, c/o National Museum of Natural History, Washington, DC, 20560-0168, USA (e-mail: dadamski.sel.barc.usda. gov); Juan A. Villanueva-Jiménez, Colegio de Post- graduados, Campus Veracruz, Mexico; Ma- nuel Méndez L6pez, Organismos Benéficos para la Agricultura, Autlan, Mexico. PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 814-816 NOTE Tapinoma sessile (Say) (Hymenoptera: Formicidae) Nest in Association with the Northern Pitcher Plant, Sarracenia purpurea L. (Sarraceniaceae) Tapinoma sessile (Say), the odorous house ant, is a native species found throughout the United Sates and Canada and is considered a house pest (Wang and Brook 1970). This species is extremely flexible in selecting nesting sites. Nests are excavated in the soil, with or without a cov- ering object such as stones. Nests are also constructed under bark and have been re- ported in a variety of preformed cavities (Creighton 1950). When colonies nest in the soil, the nests are shallow and tempo- rary. In studying nest relocation in four spe- cies of ants, Smallwood reported that 7. sessile moved its nest approximately every twenty-six days (Smallwood 1982). Nests of T. sessile have been reported in bogg areas as well as swamps (Smith 1928). While 7. sessile is a diminutive species, its colonies can consist of several thousand workers and are reported to be polygynous. New colonies can be formed by a queen and a group of workers leaving the parent colony (budding) or by a single founding queen. Workers of 7. sessile visit plants to col- lect honeydew from plant-feeding insects including mealybugs, aphids and plant hop- pers, as well as to collect floral and extra- floral nectars. They also are reported to feed on both living and dead insects. An elevated soil nest of 7. sessile was observed against a leaf of Sarracenia pur- purea L., the northern pitcher plant (Fig. 1), im. Suitland Bog. ¢ Prince Georges’ iCo:, Maryland. Suitland Bog is a hillside fen. The ant nest and associated pitcher plant were monitored for several weeks. During the observation period one to six worker ants were observed on the hood (B) and lip (A) of the leaf closest to the ant colony (Fig. 1). Ants were sporadically observed on other leaves of the same plant cluster. As one ant would leave the leaf closest to the nest, others would arrive. Yet, the num- bers of ants on the leaf at any one time remained between |—6 workers. An ant nest at that location was surprising because the ground was saturated with water. It was as- sumed that the nest did not extend below the surface. Worker ants were active for much of the day (period between 9:00 AM and 4:00 PM). At the end of the observation period, one side of the nest was partially opened. Ma- ture larvae and pupae were readily appar- ent. Since the nest was minimally disturbed, only a small fraction of the colony was vis- ible. By the time the site was revisited again in September, the ants had abandoned the nest. It is assumed that they moved to high- er and drier ground to prepare for over-win- tering. Sarracenia purpurea has dermal glands in the leaves that release extrafloral nectar (Joel 1986). The greatest density of these glands is on the lip of the leaf (Fig. 1A). The total carbohydrate content of the ex- trafloral nectar of S. purpurea has been studied by Cipollini (Cipollini et al. 1994). Nectar production is correlated with the age of the leaf. Older leaves do not produce de- tectable quantities of carbohydrates. Dress (Dress et al. 1997) reported measurable amounts of amino acids in S. purpurea ex- trafloral nectar samples. The discovery of extrafloral nectar sourc- es can prompt several different responses in ants. When foraging worker ants encounter a rich nectar source, the ants will consume the sugar source until they are replete. Oth- ers will feed and mark a trail back to the parent nest to recruit other workers. Koptur (1992) states that worker ants will treat ex- VOLUME 104, NUMBER 3 815 Fig, ll: leaf, B = hood of the leaf. trafloral nectaries in the same manner as they treat honeydew-producing insects. They will actually defend and protect the sugar source. We believe Tapinoma sessile has taken this strategy one step further. Upon locating and evaluating the extrafloral nectar source of S. purpurea, the ants built a nest next to the pitcher plant. Brood production for the ant appeared to coincide with extrafloral nectar flow from the new leaves. The northern pitcher plant is rather lo- calized in its distribution being restricted to bogs, fens and hillside seeps in the eastern United States and Canada. The observation reported here demonstrates the flexibility of T. sessile to adapt to a specific set of con- ditions. Since there was a second 7. sessile nest near a separate plant in a different part of Suitland Bog, this was not an isolated observation. The ants appeared to be taking advantage of a particular food source in the most efficient manner to maximize their brood production. Sarracenia purpurea plant with Tapinoma sessile nest to the right of the labeled leaf. A = lip of the LITERATURE CITED Cipollini, D. F Jr, S. J. Newell, and A. J. Nastase. 1994. Total carbohydrates in the nectar of Sarra- cenia purpurea L. (northern pitcher plant). Amer- ican Midland Naturalist 131: 374-377. Creighton, W. S. 1950. The ants of North America. Bulletin of the Museum of Comparative Zoology 104: 1-585, 57 plates. Dress, W. J., S. J. Newell, A. J. Nastase, and J. C. Ford. 1997. Analysis of amino acids in nectar from pitchers of Sarracenbia purpurea (Sarracen- laceae). American Journal of Botany 131: 374— Side Joel, D. M. 1986. Glandular structures in carnivorous plants: Their role in mutual expoitation of insects, pp. 219-234. Jn Juniper, B. E. and T. R. E. South- wood, eds. Insects and the Plant Surface. Edward Arnold, London. Koptur, S. 1992. Extrafloral nectary-mediated interac- tions between insects and plants, pp. 81-129. /n Bernays, E., ed. Insect-Plant Interactions TV. CRC Press, Boca Raton, Florida. Smallwood, J. 1982. Nest relocation in ants. Insectes Sociaux 29: 138-147. Smith, M. R. 1928. The biology of Tapinoma sessile Say, an important house infesting ant. Annals of the En- tomological Society of America 21: 307-329. Wang, J. S. and T. S. Brook. 1970. Toxicological and 816 biological studies of the odorous house ant, Tap- inoma sessile. Journal of Economic Entomology 63: 1971-1973. R. M. Duffield, Department of Biology, Howard University, Washington, DC PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 20059, U.S.A. (e-mail: rduffield@ howard. edu); R. R. Snelling, Entomology Section, Emeritus, Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles, CA 90007, U.S.A. ENTOMOLOGICAL SOCIETY OF WASHINGTON http://www.entomology.si.edu/ES W/ES W Menus. lasso Information and Officers Bylaws History First 100 Years Past Presidents History of the ESW Seal Information for Authors Available Publications Meetings Members Database Membership Application Subscriptions PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 817-819 NOTE Feeding Habits of Nyssodrysina lignaria (Bates) (Coleoptera: Cerambycidae: Lamiinae) Cerambycidae are essentially xylopha- gous, with few species utilizing other kinds of food (Duffy 1953; Duffy 1960; Linsley 1961; Martins 1997; Marinoni et al. 2001; Monné 2001a, b). A few species feed on seeds: Ataxia sulcata Fallén (Pteroplini), Leptostylus terraecolor Horn, Lophopoeum timbouvae Lameere, and Nyssodrysina lig- naria (Bates) (Acanthocinini) (Marinoni et al. 2001). Studies indicate that adults of Nys- sodrysina lignaria have been obtained from pods of legumes, such as /nga lushnathiana Benth. (Leguminosae) (Bosq 1943) and from the fruit of Theobroma cacao (Linnae- us) (Sterculiaceae) (Bondar 1939). The only reports describing seed-living larvae are Lima (1955) who cited species of /nga as host plants, and Biezanko and Bosq (1956) who noted the presence of larvae boring seeds of Inga uruguensis Hooker and Arnott and Inga virescens Benth. Three adults that developed in the pods of Inga sessilis (Vell.), collected in Curitiba, during the month of November (1998), al- lowed insight into the feeding habits of Nys- sodrysina lignaria. Lorenzi (1998) defined the fruit of /nga sessilis (inga-macaco; inga- ferradura; inga-carneiro) as being a ““woody legume, tomentous-velvety, flattened and curved like a horseshoe, containing few seeds covered by a thick layer of sweet aril.” The pod in which the insects were reared had a length of 14 cm on the convex side and a width of approximately 2.5 cm at the widest part. The seeds had a length of about 1.5 cm and a width a little less than | cm. Studies of pods and characteristics of the feeding behavior of N. lignaria were con- ducted after the emergence of adults, which occurred in September (two individuals) and November (one individual) of 1999. It was shown that in one of the pod valves, the adults made two emergence holes (Fig. | de- picts one of these openings). There was no hole on the internal wall (endocarp), indi- cating a passage of the larva to the internal cavity of the pod, which would make feed- ing on the seeds possible. These findings in- dicate that the two insects developed by feeding exclusively on mesocarp. Oviposi- tion must have been next to the suture of the pod valves. The larva, penetrating into the exocarp, produces a gallery parallel to the suture (Fig. 3), where the remains (excre- ment, particles and fibrous strips) are com- pacted inside. As the larva develops, it bores into the mesocarp toward the middle of the pod. The gallery takes the shape of a wide pupal chamber next to the exit hole (Fig. 3). The chamber appears as a simple cavity without any delimiting artifact. The third specimen reached the internal cavity of the pod, which was evident by the presence of an oval opening in the internal wall (endocarp) of the valve (Fig. 2). It fed on part of the seed (Fig. 4), and the amount consumed was small, indicating that a large part of larval alimentary needs had been sup- plied by the ingestion of cellulose from the valve. The larva still produced long strips from the woody tissue that constituted the en- docarp of the pod (Fig. 5). These strips were dispersed in the internal cavity, uncharacter- istic of the formation of a bird nest type pupal chamber, as occurs in various genera of Cer- ambycidae (Duffy 1953, 1960). The behavior of these three individuals indicates that the spermophagous nature of this species must be circumstantial. The in- gestion of pod walls should be the basic source of food. This behavior (carpophagy ) can be considered as transitory in the pro- cess of adaptation to a strictly spermopha- gous habit. Even though the spermopha- gous behavior of Nyssodrysina lignaria may be viewed as circumstantial, it should 818 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-5. be noted that two other species of the same tribe Acanthocinini (described above) have been reported to be seed consumers. Costa and Link (1988) and Marinoni and Ganho (in press) point out other findings on the behavior of these beetles. The first au- thors collected 1162 specimens of N. lig- naria, during a One-year period (August, 1986 to July, 1987), utilizing impact traps Pods of Inga sessilis with Nyssodrysina lignaria. 1, Adult emergence hole. 2, Internal view of one valve showing hole made by larvae. 3, Internal view of one valve showing gallery and pupal camera produced by larva (residues in gallery removed). 4, Seed partially consumed by larva. 5, Internal view of one valve showing fibrous stripes of endocarp produced by larva. with alcohol as an attractant in six tree planting fields (eucalyptus, pine, “‘acacia negra,” “ipé amarelo”’). This number cor- responds to about twice (591 individuals) that collected by the project ““‘Levantamento da Fauna Entomolégica no Estado do Pa- rand (PROFAUPAR)”’ in eight sites, utiliz- ing Malaise traps (384) and light traps (207) carried out during the same period (August, VOLUME 104, NUMBER 3 1986 to July, 1987) when this species was the most abundant among all the Ceram- bycidae from a total of 2000 specimens (Marinoni and Ganho, in press). These findings allow us to point out two important aspects. The first is the large dif- ference between collections using traps with and without alcohol as an attractant. The second is the abundance of the species (S91 specimens) in the collections by PRO- FAUPAR, which was greater than that of all other species. The second most abundant species was Compsa albopicta (Perty), with only 166 specimens. The reason for the large abundance of N. lignaria must be related to the utilization of fruit for food by larvae. The feeding habit of this species, although not essentially sper- mophagous, is associated with fruit and not wood (trunk, limbs and roots). The fruit of these host species, having an annual cycle, can provide a larger quantity of food com- pared to wood which the majority of Cer- ambycidae species utilize only after dying or falling (due to wind, lightning, senility, etc.). The greater abundance by collections us- ing traps containing alcohol as an attractant is likely connected with the search for fruit for reproduction. The seeds of /nga sessilis, such as those of /nga edulis Mart., are cov- ered by a thick layer of aril. The aril be- comes sweet when the fruit ripens. This fa- vors the development of a fermentation pro- cess resulting in the production of alcohol (Duke 1983), which must be the substance that attracts Nyssodrysina lignaria. We are grateful to George L. Venable for help in producing the plate, to Albino M. Sakakibara for the photos, and to Steven W. Lingafelter and Wayne N. Mathis for re- viewing a draft of this paper. We are also grateful to FUNPAR, CNPq and CAPES for the support. LITERATURE CITED Biezanko, C. M. and J. M. Bosq. 1956. Cerambycidae de Pelotas e seus arredores. Agros 9(3/4): 3-16. Bondar, G. 1939. Insetos daninhos e parasitas do cacau 819 da Bahia. Boletim Tecnologico do Instituto do Ca- cau, Bahia 5: 1-112. Bosq, J. M. 1943. Segunda lista de cole6pteros de la Republica Argentina, daninos a la agricultura. Buenos Aires, Ministerio de la Agricultura, Direc- cion de Sanidad Vegetal, 80 pp. Costa, E. C. and D. Link. 1988. Flutuagao populacion- al de Nyssodrysina lignaria (Bates, 1864) (Cole- optera, Cerambycidae). Anais do VI Congresso Florestal Estadual, Nova Prata, RS, pp. 541—550. Duffy, E. A. J. stages of British and imported timber beetles. London, British Museum (Natural History), 350 PP- . 1960. A monograph of the immature stages of Neotropical timber beetles (Cerambycidae). London, British Museum (Natural History), 327 Pp. Duke, J. A. 1983. Inga edulis Mart. (Mimosaceae). From Handbook of energy crops (unpublished), on 15 Oct 2001 (web site: ?hort.purdue.edu/new- crop?). Lima, A. M. C. 1955. Insetos do Brasil, Cole6pteros, vol. 9, Escola Nacional de Agronomia, Rio de Ja- neiro, 289 pp. Linsley, E. G. 1961. The Cerambycidae of North America. Part 1. Introduction. University of Cal- ifornia Publications in Entomology 18: 1—135. Lorenzi, H. 1998. Arvores brasileiras, Vol. 2, Ed. Plan- tarum Ltda, Nova Odessa, Sao Paulo, 352 pp. Marinoni, R. C. and N. G. Ganho. In press. Sazonali- dade de Nyssodrysina lignaria (Bates, 1864) (Cer- ambycidae, Lamiinae), no Estado do Parana, Bras- il. Marinoni, R. C., N. G. Ganho, M. L. Monné, and J. R. M. Mermudes. 2001. Habitos alimentares em Coleoptera (Insecta). Ribeirao Preto, Ed. Holos, 63 pp. Martins, U. R. 1997. Cerambycidae sul-americanos (Coleoptera). Taxonomia, Vol. 1, ed. Sociedade Brasileira de Entomologia, Sao Paulo, 217 pp. Monneé, M. A. 2001la. Catalogue of the Neotropical Cerambycidae (Coleoptera) with known host plant—Part I. Subfamily Cerambycinae, Tribes Achrysonini to Elaphidiini. Publicagoes Avulsas do Museu Nacional 88: 1-108. . 2001b. Catalogue of the Neotropical Ceram- bycidae (Coleoptera) with known host plant—Part Il. Subfamily Cerambycinae, Tribes Graciliini to Trachyderini. Publicagdes Avulsas do Museu Na- cional 90: 1-119. 1953. A monograph of the immature Renato C. Marinoni, Norma G. Ganho, and Cibele S. Ribeiro-Costa, Department of Zoology, Federal University of Parana, Caixa Postal 19020, 81531.990, Curitiba, Brazil (RCM e-mail: rcemari@ bio.ufpr.br; NGG e-mail: norma®@ bio.ufpr.br; CSRC e- mail: stra@bio.ufpr.br) PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 820-821 NOTE Replacement Names in the Eumaeini (Lepidoptera: Lycaenidae: Theclinae) In preparing the Eumaeini (Lepidoptera: Lycaenidae: Theclinae) part of the Atlas of Neotropical Lepidoptera Checklist (“Atlas Checklist” hereafter, Robbins, in press), I discovered two generic and two specific homonyms that need to be replaced. The purpose of this note is to publish them in advance of the Atlas Checklist. GENERIC NAMES Eucharia Boisduval 1870 (type species Papilio ganimedes Cramer) is a junior hom- onym of Eucharia Hiibner [1820] (Lepi- doptera: Arctiidae). D’ Abrera and Balint (in D’ Abrera 2001) proposed the generic name Annamaria with type species Thecla draud- ti Lathy, a congener of P. ganimedes, but Annamaria D’ Abrera and Balint is unavail- able under ICZN Article 13.1.1 (1999) be- cause this genus was not “‘accompanied by a description or definition that states in words characters that are purported to dif- ferentiate the taxon.’? Consequently, the new name Lamasina Robbins is proposed as a replacement name for Eucharia Bois- duval. Lamasina contains four species in the Atlas Checklist: Lamasina ganimedes (Cramer 1775) new combination, Lamasi- na draudti (Lathy 1926) new combination, Lamasina rhaptissima (K. Johnson 1991) new combination, and Lamasina saphon- ota (Constantino, Salazar, and K. Johnson 1993) new combination. Lamasina is mas- culine and named for Gerardo Lamas, with deep appreciation for his prodigious contri- butions to the Atlas Checklist and to our knowledge of Neotropical butterflies in general. D’ Abrera and Balint (in D’ Abrera 2001) proposed the generic name Gulliveria for type species Thecla gigantea Hewitson. However, this name is preoccupied by Gul- liveria Castelnau 1878 (Pisces). The new name Megathecla Robbins is proposed as a replacement name. Megathecla is femi- nine and is another way of drawing atten- tion to the size of the type species. This genus consists of two species in the Atlas Checklist: Megathecla gigantea (Hewitson 1867) new combination and Megathecla cupentus (Stoll 1781) new combination. SPECIFIC NAMES Thecla aurorina Draudt 1921 is trans- ferred to Theclopsis Godman & Salvin 1887 in the Atlas Checklist, but is preoc- cupied by the earlier name Thecla aurorina Oberthiir 1880. The new name Theclopsis aurina Robbins is proposed to replace The- cla aurorina Draudt 1921. Theclopsis au- rina is an indeclinable, non-latinized name that is intended to be euphonious and rem- iniscent of 7. aurorina. Serratofalca callilegua K. Johnson & Sourakov 1993 and Gigantofalca calilegua K. Johnson 1995 are transferred to Caly- copis Scudder 1876 in the Atlas Checklist. As a consequence, they are secondary hom- onyms under ICZN Article 58.7 (1999), de- spite different spellings, because they both refer to the name of the same geographical locality (Calilegua). The new name Caly- copis mirna Robbins & Duarte is pro- posed as a replacement for the junior hom- onym, Gigantofalca calilegua K. Johnson 1995. Calycopis mirna was described from Argentina, but we know it as a widespread species in southern Brazil and name it as a noun in apposition for Mirna Casagrande in recognition of her many contributions to our knowledge of Neotropical butterflies, including extensive collecting in southern Brazil. Acknowledgments.—I am most grateful to Gerardo Lamas (Lima, Peru) for superb advice on nomenclatural matters. Marcelo VOLUME 104, NUMBER 3 Duarte (Curitiba, Brazil) first realized that the species that we knew from southern Brazil was the same as Calycopis calilegua from Argentina, which is why he ts the co- author of the replacement name for this tax- on. LITERATURE CITED D’ Abrera, B. 2001. The Concise Atlas of Butterflies of the World. Melbourne/London, Hill House Pub- lishers, 353 pp. ICZN. 1999. International Code of Zoological Nomen- 821 clature, Fourth Edition. The International Trust for Zoological Nomenclature, 306 pp. Robbins, R. K. In press. Eumaeini. /n Lamas, G., ed. Checklist: Part 4A. Papilionoidea—Hesperioidea. In Heppner, J. B., ed. Atlas of Neotropical Lepi- doptera. Scientific Publishers, Gainesville. Robert K. Robbins, Department of Sys- tematic Biology, Entomology Section, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, DC 20560- O77. SCS-A: robbins.robert@ nmnh.st.edu) (e-mail: NEw PUBLICATION Revision of the Oriental Species of Aphthona Chevrolat (Coleoptera: Chrysomelidae) by Alexander S. Konstantinov and Steven W. Lingafelter This hardcover book of over 340 pages presents a revision of the large flea beetle Aphthona for the Oriental Region. This is the second monographic treatment of this genus genus (the senior author revised the Palearctic fauna in 1998). In the work, 93 species are treated, with 37 being described as new. Numerous new synonyms and combinations are recognized and are presented. Full descriptions and diagnoses are included along with dichotomous identification keys. Hundreds of illustrations, including four full color habitus drawings, are provided to facilitate identification. Of primary significance are the large, detailed aedeagal and spermathecal illustrations for every species. Distribution maps are also included. Available from the Entomological Society of Washington (see inside back cover). PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 822 NOTE Ornithodoros kohlsi Guglielmone and Keirans (Acari: [xodida: Argasidae), a new name for Ornithodoros boliviensis Kohls and Clifford 1964 Kohls and Clifford (1964) described all post-embryonic stages of Ornithodoros bo- liviensis from ticks found on bats and in houses in northeastern Bolivia. However, the name O. boliviensis is preoccupied by Ornithodoros boliviensis Bacherer Gutiér- rez 1931. He described the last nymphal stage of Otobius megnini (Duges 1883) as Ornithodoros boliviensis, from ticks fre- quently found in the ears of humans in the vicinity of Sucre, Bolivia (Bacherer Gutiér- rez 1931). Therefore, Ornithodoros boli- viensis Bacherer Gutiérrez constitutes a ju- nior subjective synonym of Ofobius meg- nini (Duges 1883) (new synonymy), and we propose the name Ornithodoros kohlsi Guglielmone and Keirans (new name) for Ornithodoros boliviensis Kohls and Clif- ford 1964. This research was supported by National Institute of Allergy and Infectious Diseases grant AI 40729 to JEK. Literature Cited Bacherer Gutierrez, R. A. 1931. Las garrapatas en el conducto auditivo externo. la Conferencia Sani- taria Boliviana 2: 1—47. Kohls, G. M. and C. M. Clifford. 1964. Ornithodoros (Alectorobius) boliviensis sp. n. (Acarina: Argas- idae) from bats and houses in Bolivia. Journal of Parasitology 50:792-—796. Alberto A. Guglielmone, /NTA, Rafaela, CC 22, CP 2300, Rafaela, Santa Fe, Argen- tina. (e-mail: aguglielmone @ inta.gov.ar); James E. Keirans, U. S. National Tick Col- lection, Institute of Arthropodology and Par- asitology, Georgia Southern University, Statesboro, GA 30460-8056, U.S.A. (e-mail: Jkeirans @ gasou.edu) PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 823-824 Book REVIEW Fauna of New Zealand No. 42. Aphodi- inae (Insecta: Coleoptera: Scarabaei- dae). By Z. T. Stebnicka. 2001. Manaaki Whenua Press, Lincoln, New Zealand. 64 pp. ISBN 0-478-09341-1. Price $29.50 (includes shipping). Order from Manaaki Whenua Press, PO. Box 40, Lincoln 8152, New Zealand or http://www. landcare.cri.nz/mwpress. The Scarabaeidae are a large family of beetles, numbering about 35,000 species. The Aphodiinae are one of the larger sub- families and account for about 3,500 spe- cies. This is the first comprehensive review of the Aphodiinae in New Zealand by one of the world’s leading authorities on apho- diines, Zdzislawa Stebnicka from Poland. While this fauna is small (only 20 species), this work presents them accurately and in detail. Although the Aphodiinae are a large, di- verse, and cosmopolitan group, little was known about them in New Zealand prior to this study. They are commonly referred to as “dung beetles,’ but not all species are associated with dung. Aphodiines feed and breed in the soil on various kinds of excre- ment and/or in vegetable debris. Two spe- cies in New Zealand are actually minor pests of cultivated plants. The monograph begins with a popular summary (in English and Maori) and a checklist of taxa. A brief section on “Systematics” includes an over- view of the classification of the subfamily. The next section on ‘‘Morphology”’ dis- cusses and illustrates the characters used to delineate aphodiines, including the diagnos- tic characters for tribes found in New Zea- land. A section on *‘Biology”’ includes in- formation on life history and feeding pref- erences. An important section on *‘Bioge- ography” follows where’ Stebnicka discusses the nine indigenous species and the 11 species that have been accidentally introduced (six from Australia, 2 from America, | from Africa, 2 from Europe). One species is found only on South Island While seven are found only on North Island (including one also found on the Kermadec Islands). One species is indigenous to the Three Kings Islands, one to the Chatham Islands, and one to the Kermadec Islands. The New Zealand fauna of Aphodiinae is similar to that of Australia (as might be ex- pected) but with a much smaller number of taxa. Stebnicka concludes that the distri- bution of the nine indigenous species and their systematic relationships suggests a Gondwonan origin. A key to the 20 species in four tribes and nine genera is presented, and this is fol- lowed by descriptions, geographic distri- butions (with 20 locality record maps), sea- sonality, and biological associations. Two new species are described and two new syn- onymies are proposed. Two habitus draw- ings are provided as well as 41 line draw- ings and 16 scanning electron micrographs. The work concludes with an extensive ref- erences section. The information included in this work is easily accessible and well-presented in a traditional systematic treatment. The tax- onomy is sound and the descriptions accu- rate. The key is easy to use. Although the focus is only on New Zealand, this work is nevertheless comprehensive and an excel- lent presentation. This seems to be a hall- mark of Stebnicka’s work. The primary users of this work will be systematists, ecologists, resource managers, and students in New Zealand wishing to know more about their entomofauna. It will also be useful to scarab taxonomists, cura- tors and collection managers organizing their holdings, biogeographers studying southern hemisphere distribution patterns, and a fairly large set of amateur beetle col- lectors. Lastly, the form of this treatment 824 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON could easily serve as a template for others Brett C. Ratcliffe, Systematics Research to follow when writing future faunistic Collections, W436 Nebraska Hall, Univer- studies because it is so clear, comprehen- sity of Nebraska, Lincoln, NE 68588-0514. sive, and functional. U.S.A. NEW PUBLICATION Revision of the Genus Anoplophora (Coleoptera: Cerambycidae) by Steven W. Lingafelter and E. Richard Hoebeke This hardcover book of 236 pages is the first comprehensive taxonomic treatment for Anoplophora woodboring beetles. This genus included the notorious Asian longhorned beetle that was introduced recently into New York City and Chicago. Using a thorough examination of primary types scattered throughout Asia and Europe as a foundation, this work proposed over 20 new synonyms and dozens of new combinations. Full color illustrations of adults of both sexes, detailed diagnoses and descriptions, distribution maps, an identification key, and hundreds of additional figures of characters used in Anoplophora identification are included. Over 50 pages of professionally prepared full color plates also are included. Thhis work is the only comprehensive resource that will enable identifications of this important group of beetles. Available from the Entomological Society of Washington (see inside back cover). PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael E. (SPSL Soa R12 RE Ge Pl Sa A JOR ce eS ee Aa ee eS seeeenetd A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric SaRisselWaniiviEnaen es, SGHalat: Oy DpitOO7 658 Ne ee Bs (te Revision of the Oriental Species of Aphthona Chevrolat (Coleoptera: Chrysomelidae), by Alexander S. Ronstinunoviangustevem Ws lineateltern, 349) ppi 2002 242s ee a Revision of the Genus Anoplophora (Coleoptera: Cerambycidae), by Steven W. Lingafelter and E. RIGHARGAELOCHEKE: 2S Opp a UU) 2s eerie kee Trace EN eS ard De co eae ieee ae a MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 __...- No. 4. A Manual of the Chiggers, by G. 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New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. crt Mel Opp yhOO Sire seco 2 Se cee OLE eee Ne Ts a, Se pean ae METER eee es Sve 2d Tee No. 22. Systematics of the North American Species of 7richogramma Westwood (Hymenoptera: Prichostammatdas) by JOnnDPintow28 7 ppl O99 voles ees eras eee ee) Des ee ae ae eh ee No. 23. Revision of the Net-Winged Midges of the Genus Blepharicera Macquart (Diptera: Blepha- riceridae) of Eastern North America, by Gregory W. Courtney. 99 pp. 2000 _.-- $10.00 15.00 40.00 30.00 $15.00 15.00 15.00 15.00 12.00 5.00 25.00 25.00 25.00 25.00 25.00 18.00 12.00 12.00 28.00 14.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume to non-members and $25.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10 percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Treasurer, Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. CONTENTS (Continued from front cover) HASTRITER, MICHAEL W. and MICHAEL F. WHITING—Macropsylla novaehollandiae (Siphonaptera: Hystrichopsyllidae), a new species of flea from Tasmania .................--. 663 HUSBAND, ROBERT W.—A new species of Eutarsopolipus Berlese (Acari: Podapolipidae) from the Galapagos Islands, a parasite of Agonum chathami Van Dyke (Coleoptera: Carabidae) .. 563 NORRBOM, ALLEN L.—A new species and key for the genus Zonosemata Benjamin (Diptera: Tephivitidac):) dcr: cho eee etree eee WO Se. Aehank le wa 3 eis n anata AAAI ieataae serial re ae a 614 PIKREAK: Se-P: STARY, G. GRAF, and D. ALLISON—FPauesia columbiana, n. sp. (Hymenoptera: Braconidae: Aphidiinae) on juniper aphids, and a key to related species ..............+--++-+- 646 PITTS, JAMES P. and DONALD G. MANLEY—Description of females of Stethophotopsis Pitts and Sphaeropthalma (Photopsioides Schuster) (Hymenoptera: Mutillidae) ..............---- 672 POLHEMUS, DAN A.—A new species of micropterous damsel bug (Heteroptera: Nabidae) from Nukuidiva;) Marquesas Islands cP acmenty sce itatinn Sei 0 spa1cicislnjels > +s eva ese eieual Saleen Miele antares 640 SCARBROUGH, A. G.—Redescription of two species of Ommatius Wiedemann, with lectotype and paralectotype designations for Ommatius tenellus van der Wulp and range extension, and a replacement name for Ommatius tibialis Ricardo (Diptera: Asilidae) ...................... 680 SCHEFFER, SONJA J.—New host record, new range information, and a new pattern of voltin- ism: Possible host races within the holly leafminer Phytomyza glabricola Kulp (Diptera: A STOMIY AIAG) iA. sok 5 sic Oe tethers Meapesin ea Se UN hate ntaleteic os «B's e's Sv do Ue 0 ale We Sea ea 571 SCHRODER, R. F. W. and M. M. ATHANAS— Biological observations of Centistes gasseni Shaw (Hymenoptera: Braconidae), a parasitoid of Diabrotica spp. (Coleoptera: Chrysomelidae) .. 554 SMITH, DAVID R. and AKIHIKO SHINOHARA—A new genus and new species of Cephidae (Hymenoptera) from Sulawesi Utara, Indonesia ................ eee eee ee eee eee teen eee 624 STAINES, C. L.—The New World tribes and genera of Hispines (Coleoptera: Chrysomelidae: (QE ISS ahha ete) PINE GAR bell nny Mine Mia Lice nel aia = Si Wet) (ek i eS COUGAR beatnik gaa acime 721 WHEELER, A. G., JR—Leptoypha elliptica McAtee and L. ilicis Drake (Hemiptera: Tingidae): New distribution records of seldom-collected lace bugs, with clarification of host-plant FELATIOMSHIDS: Of ahi uo dene da Sie on le siete hese EMER SIS cles, er oir eee NOTES ADAMSKI, DAVID, JOHN W. BROWN, JUAN A. VILLANUEVA-JIMENEZ, and MANUEL MENDEZ LOPEZ—First records of the sugarcane pest, Blastobasis graminea Adamski (Lepidoptera: Coleophoridae: Blastobasinae), from Mexico and Central America .......... 812 DUFFIELD, R. M. and R. R. SNELLING—Tapinoma sessile (Say) (Hymenoptera: Formicidae) nest in association with the northern pitcher plant, Sarracenia purpurea L. (Sarraceniaceae) 814 GUGLIELMONE, ALBERTO A. and JAMES E. KEIRANS—Ornithodoros kohlsi Guglielmone and Keirans (Acari: Ixodida: Argasidae), a new name for Ornithodoros boliviensis Kohls and G) IT Zax yo Bt Koy cy pata eae oy ane eat og Same mage Hes NEY MRE 2 Ln On eSpace aren ae 5b Aki 2 822 MARINONI, RENATO C., NORMA G. GANHO, and CIBELE S. RIBEIRO-COSTA—Feeding habits of Nyssodrysina lignaria (Bates) (Coleoptera: Cerambycidae: Lamiinae) ............ 817 ROBBINS, ROBERT K.—Replacement names in the Eumaeini (Lepidoptera: Lycaeinidae: Theclimaeye 733i se See a eee yeh re SEI ara Darcie USNS a Mae pea chattel stavei ae =a Sets eden 820 BOOK REVIEW RATCLIFFE, BRETT C.—Fauna of New Zealand No. 42. Aphodiinae (Insecta: Coleoptera: Scarabaeidae), by, ZT Stebnicka) o2 yeas geese he teres hte stars aioe cieleeet oe eet eicteteleistet tela. foloho lets 823 NEW BUBEICATIONS 7 hse. .icnke coe WL ang Motes siea hs Bd Maveh larch aterm ain cla) tl ataye pane stadia stay ete ete VOL. 104 OCTOBER 2002 NO. 4 He PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED QUARTERLY ADAMSKI, DAVID and JOHN W. BROWN—A new specieSNgf Fremey Gelechiidae: Gnorimoschemini) from México: A potential bidtealig PLAC OTI OUELITEA (SOLAMACEAG) Met Pee a ee bot sha) or Te es eile eels wont prance Oy eTesal ante ere a Peo catchers 1029 ARCE-PEREZ, ROBERTO—A new species of Psephenops Grouvelle (Coleoptera: Psephenidae) RRYETAW NAGS Xo CO BPMN raed MAY Png ema au clea Sale avate,s/5 sa craic tavaluietet te syahaneveteteiete Buspar ags metoio ere eet eeyoreis 964 BURGER, JOHN F.— Description of five new species of Tabanidae (Diptera) from Costa Rica and revised keys to species for the genera Fidena Walker, Scione Walker, and Chrysops laren tin (COSA) aol. aiak da aodho dss OSS BBBONpE Sete Cab ecaoooMDEMaUUMEmonoadcpias yao 928 CRYAN, JASON R. and LEWIS L. DEITZ—Enigmatic treehopper genera (Hemiptera: Membracidae): Deiroderes Ramos, Holdgatiella Evans, and Togotolania, new genus ........ 868 HALL, JASON P. W. and DONALD J. HARVEY—A revision of the Neotropical butterfly genus Scecomsalleandenanveya lsepidoptera:!RIOGINIGAS)) 55.2 - reacties ete ci ds isis eee rie -ae 941 HESPENHEIDE, HENRY A. and LOUIS M. LAPIERRE—A review of Ptous Champion (Coleoptera: (Cuirenilrormre bres (Ciayaliord mantel ci iXe)) Go enaeopanuna se beortaneebebdd eb ao oe ouneS oes auosepduedoae 856 JOHNSON, NORMAN F. and LUCIANA MUSETTI—Rediscovery of the genus Platyscelidris Szab6 (Hymenoptera: Scelionidae) and description of a new species ................0.--005- 948 MILLER, DOUGLASS R., GARY L. MILLER, and GILLIAN W. WATSON—Invasive species of mealybugs (Hemiptera: Pseudococcidae) and their threat to U.S. Agriculture ............ 825 MILLER, GARY L. and DOUGLASS R. MILLER—Dysmicoccus Ferris and similar genera (Hemiptera: Coccoidea: Pseudococcidae) of the Gulf State Region including a description of A inewy Soseewamelmeyy Uintieal Siatves meee hy So Sees aba sane dodo oases seca 57 sspenodnpoRscdn so 968 MORON, MIGUEL ANGEL and ROBERTO ARCE—Descriptions of the immature stages of five Mexican species of Gymnetini (Coleoptera: Scarabaeidae: Cetoniinae) ................ 1036 NEUNZIG, H. H. and M. A. SOLIS—The Ceracanthia complex (Lepidoptera: Pyralidae: Phycinmnae) ia) Gosta Rican |. ICeracarntnia RAGOMOL .o\..2.ce8 visie seicesn is so aataya ais sete De slergie ese es a's 837 NEUNZIG, H. H. and M. A. SOLIS—The Ceracanthia complex (Lepidoptera: Pyralidae: Phycitinae) in Costa Rica. Il. Megarthria Ragonot, Drescoma Dyar, and Lascelina Heinrich 980 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2002 GABRIELA CHAVARRIA, President MICHAEL G. PoGuE, Treasurer JONATHAN R. Mawps ey, President-Elect RONALD A. OcHoA, Program Chair Stuart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Ho..is B. WILLIAMS, Corresponding Secretary JOHN W. Brown, Past President Jon A. Lewis, Custodian Davip R. Smitn, Editor Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. MartuHIs Honorary Members KARL V. KROMBEIN RONALD W. HopGEs 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 7:00 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $25.00 (U.S. currency). PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub- lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send address changes to the Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomo- logical Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE PP. 247-248 OF THE JANUARY 2002 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, “% Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 27 September 2002 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 825-836 INVASIVE SPECIES OF MEALYBUGS (HEMIPTERA: PSEUDOCOCCIDAE) AND THEIR THREAT TO U.S. AGRICULTURE DOUGLASS R. MILLER, GARY L. MILLER, AND GILLIAN W. WATSON (DRM, GLM) Systematic Entomology Laboratory, Plant Science Institute, Agricultural Research Service, U.S. Department of Agriculture, Beltsville, MD 20705 U.S.A. (DRM e-mail: dmiller@sel.barc.usda.gov; GLM e-mail: gmiller@sel.barc.usda.gov); (GWW) En- tomology Department, The Natural History Museum, Cromwell Road, London, SW7 SBD, U.K. (GWW e-mail: g.watson@nhm.ac.uk) Abstract.—We provide a compilation of 158 species of mealybugs that are either a pest or threat to United States agriculture. Included for each species, where applicable, is reference to: the United States origin and date of introduction; whether it is established in the United States; its pest or threat status in the United States along with a validation citation; its principle hosts; and its biogeographical region of origin. Key Words: Mealybugs (Pseudococcidae) are major agricultural pests and pose serious problems When introduced into new areas of the world without natural enemies. In recent years, it has become increasingly obvious that introduced species have a major eco- nomic and esthetic impact on plants. Public concern stimulated Congress to develop a report on the subject (Office of Technology Assessment 1993) and a Presidential initia- tive (Executive Order 13112) to seek a so- lution to this problem is currently under- way. Scale insects are notorious invaders because they are small, often live in con- cealed habitats, and frequently are trans- ported on commodities that are common in international commerce. Mealybugs are a group of scale insects that are of special interest because of the recent introduction of four pest species in the United States. They are: the pink hibiscus mealybug, Ma- conellicoccus hirsutus (Green) (California 1999), the papaya mealybug, Paracoccus marginatus Williams and Granara de Wil- Pseudococcidae, mealybugs, invasive species, biological control, agriculture link (Florida 1998), the vine mealybug Planococcus ficus (Signoret) (California 1994), and the banana mealybug Pseudo- coccus elisae Borchsenius (Florida 1995). The origin of recent California infestations of P. ficus is not clear. This species was collected in several localities in the south- eastern United States beginning in 1924, but was always found on fig trees (Ficus carica L.). It has not been recollected since 1959 and was never collected east of Texas. The infestation in California is only on grapes and has not been found on figs. It is likely that the California infestation is from an Overseas source and not from the eastern United States. Since invasive species are a major issue in U.S. agriculture, and with several recent introductions of pest mealybug species, we decided to investigate several parameters concerning invasive mealybug species. Our objectives were: |) to develop a preliminary list of the pest mealybug species of the world; 2) to provide a list of introduced and 826 pest mealybugs of the United States; 3) to ascertain which of the species in the pre- vious two objectives are introduced or na- tive to the United States; 4) to examine data provided by the United States Department of Agriculture, Animal and Plant Health In- spection Service—Plant Protection and Quarantine (USDA, APHIS-PPQ) concern- ing the most commonly intercepted mealy- bug species at the United States ports-of- entry; and 5) using this information, to try to understand which mealybugs are most likely to invade the United States in the fu- ture. MATERIALS AND METHODS Executive Order 13112 established the National Invasive Species Council and pro- vided a definition of an invasive species as: a species that is 1) non-native (or alien) to the ecosystem under consideration and 2) whose introduction causes or is like- ly to cause economic or environmental harm or harm to human health.” This def- inition not only includes alien species to the United States but also encompasses native species. The definition also has an econom- ic or potential economic component. Using this definition, the grape mealybug, Pseu- dococcus maritimus (Ehrhorn) would be an example of an invasive species in the Unit- ed States even though it likely is native. The definition that we use is a bit more sim- plistic. We consider invasive species to be those that are not native to the United States (alien or adventive) regardless of economic harm. To make a table of pest mealybugs of the world, we have used a broad definition of the term “pest.” Essentially, if a species is described in the literature as a pest, or as causing damage, or being controlled, or of economic importance, we have included it in the list. Our perspective for this paper has focused on the impact or potential im- pact of a pest mealybug in the United States. For example, Antonina pretiosa Fer- ris is known only from bamboo, and there- fore, is considered to have relatively minor PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pest potential in the United States; it may be far more important in areas of the world where bamboo is a predominant component of the natural or ornamental vegetation. Conversely, Pseudococcus cryptus Hempel occurs on an array of agricultural crops that are important to the United States economy and is considered a major threat. The term “‘threat’’ is used for species that are consid- ered pests but do not occur in the United States. For adventive species, we have ap- proximated the United States date of intro- duction by searching the literature for the earliest collection record or have examined specimens in the National Entomological Collection of the National Museum of Nat- ural History, in Beltsville, Maryland for the earliest record. Obviously, these dates are only estimates of when the species first in- vaded the United States. It is difficult to determine the zoogeographic area of origin for some species. It is clear that they are from the old or new world, but it is not always evident from which area. In some cases, we have simply made an educated guess based on the current distribution of the species, the distribution of what appears to be its closest relatives, and the natural distribution of its primary host plants. We have used the same criteria to determine if a particular species is native to the United States. In several instances, it is not clear, e.g., Phenacoccus gossypii Townsend and Cockerell or P. dearnesii King. RESULTS Table | provides information on 158 mealybug species. Since the table includes six species that have been introduced into the United States but are not considered pests, we estimate that there are 152 mealy- bugs that represent either a pest or threat to the United States agriculture. In the United States, there are 66 pest mealybug species, 19 are considered to be native. Therefore, 47 mealybug pests in the country are in- vasive species. 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With the exception of the 1980’s, every decade since the 1880’s has seen the introduction of at least 2 species of mealybug pests (Fig. 1). Two periods, the 1910’s and the 1990’s, witnessed the greatest number of introduced mealybug pests (9 and 7 species, respectively). A summary of the area of origin of all invasive species in the United States is as follows: Palearctic Region—17 U.S. invad- ers, 4 of which are considered non-pest ad- ventives; Neotropical Region—12 U.S. in- vaders, none are considered non-pest ad- ventives; Oriental Region—7 U.S. invaders, one is considered a non-pest adventive; Australasian Region—4 U.S. invaders, one is considered a non-pest adventive; Afro- tropical Region—3 U.S. invaders, none are considered non-pest adventives; and Nearc- tic Region outside of the U.S.—4 U.S. vaders, none are considered non-pest ad- ventives. Examination of the origin of pest mealy- in- Introduction of mealybug pests into the United States from 1880—1999. bugs worldwide provides the following to- tals: from the Palearctic Region, 46; Nearc- tic Region, 27; Neotropical Region, 25; Oriental Region 23; Afrotropical Region, 19; and from the Australasian Region, 18. Host characteristics of these pests include: 22% polyphagous; 20% on grasses; 16% on citrus/tropical fruits; 6% on coffee; and the remainder are not polyphagous and occur on various other hosts. Based on the char- acteristics of the highest pest mealybug dis- tributions and greatest frequency of host plants, a list of 10 species most likely to invade the United States has been deter- mined. These species include: Cataenococ- cus hispidus (Morrison), Dysmicoccus neo- brevipes Beardsley, Heliococcus bohemicus Sule, Nipaecoccus viridis (Newstead), Phenacoccus avenae Borchsenius, Phena- coccus azaleae Kuwana, Planococcus lila- cinus (Cockerell), minor (Maskell), Pseuwdococcus cryptus Hempel, and Rastrococcus iceryoides (Green). Interception records from the past five years from the USDA, APHIS-PPQ also were searched. A list of the 10 species in- Planococcus VOLUME 104, NUMBER 4 tercepted most frequently at U.S. ports-of- entry are: Cataenococcus hispidus (Morri- son); Dysmicoccus bispinosus Beardsley; Dysmicoccus mackenziei Beardsley; Dys- micoccus neobrevipes Beardsley; Maconel- licoccus hirsutus (Green); Palmicultor pal- marum (Ehrhorn); Paracoccus marginatus Williams and Granara de Willink; Plano- coccus kraunhiae (Kuwana); Planococcus lilacinus (Cockerell); and Planococcus mi- nor (Maskell). Comparison of the two lists reveals four species that are common to both. They are: Cataenococcus hispidus; Dysmicoccus neobrevipes; Planococcus lil- acinus; and Planococcus minor. We suggest that these species are most likely to be the next invasive mealybugs into the United States. DISCUSSION Our data indicate that the decades start- ing in 1910 and 1990 had the largest num- ber of mealybug introductions. We specu- late that the first peak occurred because of implementation of the Plant Quarantine Act in 1912. At this time, new inspection pro- cedures were started causing the detection of many insect contaminants in import commodities at U.S. ports-of-entry. In- creased detection caused the development of strategies to reduce the risk of introduc- tion of the potential invasive species. We suspect that the recent increase in world trade and the difficulty of inspecting large volumes of containerized imports may ex- plain the larger number of mealybug estab- lishments in the 1990’s. Our findings provide predictions about the most likely mealybug species to be in- troduced into the United States in the future and give procedures that may help others make predictions about the next invasive mealybug in other countries. Another cri- terion for predicting invasive species prob- lems that was not considered here might in- clude examination of all polyphagous spe- cies that have several agricultural hosts. A final observation: although invasive species are not always economic before they invade 833 a new area, examination of the list of U.S. invaders reveals that most were considered to be at least minor pests before they were invasive in the U.S. ACKNOWLEDGMENTS We thank Paris Lambdin (The University of Tenessessee, Knoxville, USA), Douglas J. Williams (Department of Entomology, The Natural History Museum, London, UK), Wiliam E Gimpel, Jr. (Maryland De- partment of Agriculture, Annapolis, MD, USA), Steven W. Lingafelter (Systematic Entomology Laboratory, USDA, Washing- ton, DC, USA), and Ronald Ochoa (Sys- tematic Entomology Laboratory, USDA, Beltsville, MD, USA) for their comments and reviews of the manuscript. We are also grateful to Joe Cavey (APHIS-PPQ, Riv- erdale, MD) for providing mealybug inter- ception data at U.S. ports-of-entry and Pen- ny Gullan (University of California, Davis, CA, USA) for her help in determining the earliest collection date of Ferrisia malvas- tra in the U.S. Thanks is also due to Ray Gill (California Dept. of Food and Agricul- ture, Sacramento, CA, USA) for allowing us to cite his in press work concerning in- formation about Vryburgia trionymoides. LITERATURE CITED Ali, S. M. 1962. Coccids affecting sugarcane in Bihar (Coccidae: Hemiptera). Indian Journal of Sugar Cane Research and Development 6: 72-75. Azhar, I. 1983. Some preliminary observations on the ecology of selected cocoa pests in west Malaysia. 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The biology of Phenacoccus aza- leae (Pseudococcidae) in the forest of Bunge Prickly Ash, northern China. 40 In:, VIII[th Inter- national Symposium on Scale Insect Studies. 41 PP- PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 837-855 THE CERACANTHIA COMPLEX (LEPIDOPTERA: PYRALIDAE: PHYCITINAE) IN COSTA RICA. I. CERACANTHIA RAGONOT H. H. NEUNZIG AND M. A. SOLIS (HHN) Department of Entomology, North Carolina State University, Raleigh, NC 27695-7613, U.S.A.; (MAS) Systematic Entomology Laboratory, PSI, Agricultural Re- search Service, U.S. Department of Agriculture, % National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: asolis@sel.barc. usda.gov) Abstract.—The mostly Neotropical Ceracanthia complex (Ceracanthia Ragonot, Me- garthria Ragonot, Drescoma Dyar, Lascelina Heinrich) is defined. The genus Ceracanthia is redescribed and Drescomopsis Dyar is newly synonymized with Ceracanthia. Ten species of Ceracanthia are recognized in Costa Rica, five of which are described as new: Ceracanthia alturasiana, Ceracanthia cornuta, Ceracanthia eugenieae, Ceracanthia pseudopeterseni, and Ceracanthia squamimagna. Three of the remaining species are transferred from Megarthria and given the new combinations: Ceracanthia frustrator (Heinrich), Ceracanthia schausi (Heinrich), and Ceracanthia squamifera (Heinrich), and one species is referred from Drescomopsis to form the new combination Ceracanthia soraella (Druce). Keys are provided to separate the genera belonging to the Ceracanthia complex and for males of the species belonging to Ceracanthia. Habitus photographs of male adults, and line drawings of male antennae and male genitalia, of all species are included. Also figured are the female genitalia of six of the ten species and the costal scale cluster or scale ridge of the forewings of Ceracanthia soraella and Ceracanthia frustrator. Key Words: Phycitinae, taxonomy, Neotropical celina Heinrich. The characteristic fore- wing, of most species of the complex, is The subfamily Phycitinae is the largest of the five subfamilies currently in the Pyral- idae. Many included species are morpho- logically hyperdiverse, particularly with re- gard to the male antennae and male geni- talia. Recent extensive collecting of phyci- tines, particularly by personnel at the Instituto Nacional de Biodiversidad (IN- Bio), Santo Domingo, Costa Rica, has prompted an in-depth study of the Phyciti- nae in Costa Rica. One group belonging to the subfamily that has been frequently col- lected belongs to a complex consisting of the genera Ceracanthia Ragonot, Megar- thria Ragonot, Drescoma Dyar, and Las- mainly pale on its anterior (costal) half and chiefly dark on its posterior (inner) half (Figs. 2-9). Many have the male antenna with a basal sinus bearing an upper and lower spine or protuberance (Figs. 11—20). All have the eighth abdominal sternite of the male developed as a digitate pocket. The male genitalia have a strong fusion of the gnathos and scaphium, large, broad val- vae, and usually a long vinculum. Almost all have very obvious, long setal tufts ex- tending from the base of the sacculus (Figs. 223 25) 2a 9 olieSs 2949) heaie= 838 male genitalia are simple; the only feature of any significance is a signum in the cor- pus bursae that consists of a scobinate, part- ly-sclerotized patch with, or without, a small, to large spine (Figs. 43—48). Unaware that such a complex of many similar appearing species existed, early lep- idopterists, Zeller (1881), Ragonot (1893), and Druce (1899), who in their studies of South American and Mexican species relied chiefly on forewing habitus, grouped spe- cies together and recognized only three. Later, Dyar (1914, 1919) named a few ad- ditional species from Cuba and Central America. Heinrich, in 1956, was the first to include genitalia in his study and thereby provided a more accurate, expanded ac- count of the group, particularly in Costa Rica. Neunzig and Dow (1993) reviewed the species occurring in Belize, and Neun- zig described species from Mexico (1994) and the Dominican Republic (1996). This paper provides more information on the genus Ceracanthia in Costa Rica. A subsequent contribution (Ceracanthia com- plex. II) will cover species in Costa Rica that belong to the other three genera. Most of the specimens studied were borrowed from the collection of INBio. Additional Costa Rican material has been seen that has recently accumulated in the National Mu- seum of Natural History (USNM), Smith- sonian Institution, Washington, DC, U.S.A., the Essig Museum, University of California (UCB); Berkeley, (CA, U:S-A., and) tm. the collection of Vitor O. Becker (VOB) Bra- silia, Brazil. All types of previously described species belonging to the complex were examined, including those of Megarthria (Myelois) pe- terseni (Zeller) and Ceracanthia (Homoeo- soma) soraella (Druce) in The Natural His- tory Museum (BMNH) London, England. Holotypes of new species have been depos- ited in the insect collections of INBio and UCB. Paratypes can be found at INBio, USNM, UCB, in the collection of VOB and at North Carolina State University (NCSU) Raleigh, NC, U.S.A. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON It has not been possible to associate fe- males with males with all species because of striking similarities in color and macu- lation of most species in the complex. Therefore, caution needs to be exercised us- ing females in the key to genera, and in referring to the descriptions of genera. The appearance of the immature stages of all species belonging to the complex in Costa Rica remain unknown. Heinrich (1956) included Maytenus phyllanthoides Bentham (Celastraceae) as the host of a North American species in the genus Las- celina. KEY TO GENERA OF THE CERACANTHIA COMPLEX I. Scape of male antenna broad (about 3 as wide as width of base of shaft); shaft of male antenna with basal sinus, but distal part of si- nus simple, without mesially protruding spine or other element; male genitalia with gnathos having a very weakly developed, highly scler- otized, tubular apical element, a sacculus with more than two kinds of scale tufts attached to its base (most scales strongly coiled), and a distinctly broad vinculum (disproportionately so, particularly relative to size of uncus); fe- male genitalia with ductus bursae long and dis- tinctly twisted (about 2 as long as corpus bur- SEs) the ine Sik oie ae om Aas Megarthria Ragonot — Scape of male antenna more slender (cylindri- cal) (Figs. 11—20); shaft of male antenna with or without basal sinus (if sinus is present, a spine or protuberance usually occurs at the dis- tal end of the sinus) (Figs. 11—20); male gen- italia with apex of gnathos formed into well developed, sclerotized, slender element (Figs. DiS e S729. SSS 4356515) 39) rom with out rodlike grasping element, a sacculus with one or two kind(s) of scale tufts attached to its base (all scales straight to slightly curved), and vinculum not overly broad relative to rest of genitalia; female genitalia (Figs. 43-48) with ductus bursae shorter than, or about same lengthiasscorpussibursaey yeni ee 2 Apical part of gnathos developed into a hook or rod, only partially fused to scaphium (Figs. DN, 23), 23), 2. 2D), Bil, 33, 355 Dn SD)R ASAIO EA gus without basal supplemental appendage (Figs. 22, 24, 26, 28, 30, 32, 34, 36, 38, 40); scobinate patch of signum of corpus bursae simple (Figs. 44—47), or with small, short spine (Fig. 43) or with large, mostly laterally oriented, hooked spine (Fig. 48) .......... 3 i) VOLUME 104, NUMBER 4 — Apical part of gnathos not developed into a slender rodlike element or hook (gnathos com- pletely fused to wall of scaphium); aedoeagus with supplemental, basal appendage; scobinate patch of corpus bursae with large, mostly an- teriorly oriented; hooked spime’......... SE VAS ing RE rene eV See ne Lascelina Heinrich es) Base of shaft of male antenna simple; hind- wing with distinct notch in costa, just beyond base; transtilla present, in form of a ““U” (cen- tral part straplike); valva with uniformly ta- pered, pointed projection near apex of saccu- lus; scobinate patch of corpus bursae with large, hooked spine (length of spine equal to about % width of corpus bursae) ....... RT Resin foes, clint ra fatecatar aes Drescoma Dyar — Base of shaft of male antenna with shallow to deep sinus, and usually a pair of spines (in some species scale tuft also present) (Figs. 1 1— 20); hindwing without notched costa; transtilla usually absent; clasperlike projection near apex of sacculus usually distinctly swollen (Figs. 21, PB 29) 3 1.39, 355 50)s;scObinate; patch of signum of corpus bursae simple (Figs. 44—47), or with small short spine (Fig. 43), or with large, hooked spine (length of spine equal to about % width of corpus bursae) (Fig. 48)... MBA np pS arene SO eibe eases Ceracanthia Ragonot Ceracanthia Ragonot Ceracanthia Ragonot 1893:230. Type spe- cies: Ceracanthia vepreculella Ragonot 1893. Monotypy Procandiope Dyar 1919:50. Type species: Procandiope mamella Dyar 1919. Mon- otypy Drescomopsis Dyar 1919:61. New synon- ymy. Type species: Drescomopsis subel- isa Dyar 1919. Original designation. Drescomopsis subelisa is considered to be a junior synonym of Homoeosoma so- raella Druce 1899. Note.—We have synonymized Drescom- opsis with Ceracanthia because the type species of the two genera have similar male antennae and similar genitalia. Heinrich (1956) kept subelisa in Dyar’s Drescom- opsis because of a reduced number of veins in the hindwing of subelisa, a feature that we consider to be of minor taxonomic sig- nificance. Description.—Male antenna with scape cylindrical, slightly swollen apically; shaft 839 usually with distinct sinus at base (sinus shallow in pseudopeterseni and soraella),; basal and distal end of sinus usually with spine, or protuberance; inner surface usu- ally covered with appressed scales (mamel- la, pseudopeterseni and soraella with erect papillae); sinus with or without brush or tuft of scales attached to posterior base of sinus; sensilla trichodea (cilia) of antenna about % as long as width of shaft at mid-sinus. An- tenna of female simple. Labial palpus of male upturned, reaching above vertex; 3" segment about as long as 2™ segment (about 4% as long as 2™ segment on squam- imagna). Maxillary palpus of both sexes small, short scaled. Haustellum well devel- oped. Ocelli present. Forewing of male sim- ple, or with raised costal scale cluster or scale ridge (soraella, frustrator), or with slight costal concavity or fold (pseudope- terseni, squamimagna), underside simple or with subcostal streak of contrastingly-col- ored scales. Forewing of both sexes smooth, with 11 veins; R,; ,, and R; stalked for about %, or slightly over ¥%, their lengths; M, straight to shghtly bowed; M, and M, separate or connate at base; CuA, from lower angle of cell; CuA, arising from well before lower angle of cell. Hindwing with seven to eight veins (1A, 2A, and 3A treated as one vein); Sc + R, and Rs fused for less than % their lengths beyond cell; M, straight to shghtly curved; M, and M, usually fused for less than % their lengths (M, and M, completely fused in soraella); CuA, briefly fused with stalk of M, and M, (or fused to single vein M, + M,); CuA, from before, but near lower angle of cell or from lower angle of cell; cell slightly more, to slightly less, than % length of wing. Male abdominal segment with one, or more, pair(s) of scale tufts originating from within a pocket of 8 abdominal sternite (pocket present in mamella, but without scale tufts); scales forming tufts simple, sometimes slightly broadened distally, not overly long, curved somewhat medially. Male genitalia with uncus broadly rounded apically; gna- thos with apical part developed into hook 840 or rod (center part of hook or rod weakly formed, fused with scaphium); transtilla usually absent (present in soraella, cornuta, schausi, eugenicae, alturasiana); juxta a plate with short, setiferous, lateral arms; valva broad, covered on inner surface of distal half with broadened, spinelike setae (basal half with usual setae); outer surface of valva of most species with long, black, weakly attached scales that curl under usu- ally obscuring some features of inner part of valva; sacculus with apex usually dis- tinctly swollen, not pointed; base of sac- culus with large (primary) and, usually, small (secondary) tuft of scales (eugenieae without tufts); aedoeagus slender to some- what robust; vesica usually with two groups of microspines or with larger spines (ma- mella with just one group of microspines and alturasiana with group of microspines and group of medium-sized spines). Female genitalia with ductus bursae shorter than corpus bursae; corpus bursae with signum; signum usually consisting of patch of scob- inations with inner members forming patch more strongly developed and situated on an invaginated, teardrop-shaped, highly scler- otized plate (pseudopeterseni, soraella, squamifera, squamimagna) (signum in ma- mella similar, but one scobination devel- oped into a small short spine and plate rect- angular; alturasiana with large, curved, pointed hook associated with patch of scob- inations); ductus seminalis attached to cor- pus bursae near junction of ductus bursae and corpus bursae. KEY TO SPECIES OF MALE CERACANTHIA (A key including females is not possible because the sexes of some species cannot be associated with certainty at this time.) 1. Forewing mostly pale on anterior half and dark on posterior half, with dark bands extending diagonally from about % out on costa, and from apex of wing, to dark posterior half, dividing most of pale costal area into three mainly white patches (most distal white patch small and in- distinct in most species) (Figs. 2—9) — Forewing not as above (Figs. 1, 10) i.) N PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Forewing dull, yellowish ochre along costa; postmedial line of forewing indistinct (Fig. 1); antenna with inner surface of sinus covered with erect papillae, and without brush of scales attached to posterior base of sinus See aim i ioe erence cer cen mamella (Dyar) Forewing mostly dark along costa; postmedial line of forewing distinct (Fig. 10); antenna with inner surface of sinus covered with appressed scales and with small, short brush of scales at- tached to posterior base of sinus alturasiana Neunzig and Solis, n.sp. . Forewing with long group of protruding scales extending over % length of costa (Figs. 4, 42) eed). 2 3). See frustrator (Heinrich) Forewing without long group of protruding scales extending over 4% length of costa .... 4 . Forewing with small raised scale cluster at base of costa (Figs. 3, 41); hindwing with M, and M, completely fused ........ soraella (Druce) Forewing without small raised scale cluster at base of costa; hindwing with M, and M, only partially fused . Antenna with inner surface of sinus covered with erect papillae (Fig. 13) pseudopeterseni Neunzig and Solis, n.sp. Antenna with inner surface of sinus covered with appressed scales Antenna with distal end of sinus strongly pro- duced mesially into a smoothly-scaled, conical, hookedselement (Fic. 2.0) eee eee 5 a SMR Ph. S cornuta Neunzig and Solis, n.sp. Antenna with distal end of sinus not strongly produced into a smoothly-scaled, conical, hooked element Antenna with large tuft of scales attached to posterior base of sinus (tuft extends well be- yond sinus; some scales forming tuft obovate) (Fig. 15) Antenna with tuft of scales attached to pos- squamimagna Neunzig and Solis, n.sp. terior base of sinus smaller (tuft does not ex- tend beyond sinus, or only slightly beyond si- nus) (EVeswl6=18)"": SR ee eee 8 Genitalia with inner basal lobe of valva strong- ly projecting mesially (lobe from each valva touching, or almost touching, each other); and with well developed transtilla (Figs. 35, 37) 8) Genitalia with inner basal lobe of valva weakly developed, and without transtilla (Fig. 31) PEE Oia rete ae Ee ee squamifera (Heinrich) Genitalia without scale tuft on sacculus, and with vinculum narrow (Fig. 35) ........ Sree oats Ge seger eugenieae Neunzig and Solis, n.sp. Genitalia with well developed scale tuft on sac- culus and with vinculum broad (Fig. 37)... . schausi (Heinrich) VOLUME 104, NUMBER 4 841 Figs. 1-6. Male. 1, Ceracanthia mamella. 2, C. pseudopeterseni, holotype. 3, C. soraella. 4, C. frustrator. 5, C. squamifera. 6, C. squamimagna, holotype. Ceracanthia mamella (Dyar) of the forewing is mostly a mixture of pale (Figs/-1)- 11, 21-22. 43) brown, red and black scales (appearing pur- ple without magnification). The antemedial line is very weakly formed, and the post- Ceracanthia mamella can be recognized medial line only slightly more apparent. by the appearance of the forewing (Fig. 1). The male antenna (Fig. 11) has a well A broad, dull, yellowish ochre costal band developed sinus in the base of the shaft, extends the length of the forewing. The rest with a strong, mesially directed spine near Procandiopa mamella Dyar 1919:51. 842 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-10. Male. 7, Ceracanthia cornuta, holotype. 8, C. schausi. 9, C. eugenieae, holotype. 10, C. altu- rasiana, holotype. its base and another at its distal end. The inner surface of the sinus is covered with erect papillae (a feature found only in a few other Ceracanthia (pseudopeterseni, so- raella)). The male genitalia (Figs. 21—22) have the costa of the distal half of the valva with a slightly inwardly produced distension, and the primary tuft of the sacculus distinctly longer than the length of the valva. The fe- male genitalia (Fig. 43) possess a scobinate, sclerotized, rectangular patch, bearing a short spine, in the corpus bursae. The duc- tus bursae is shorter than the corpus bursae. Costa Rican material examined.—Tur- rialba, Provincia Cartago, 17-21, 22-28 February 1965, S. S. & W. D. Duckworth (1 3, 2 2); Rio San Lorenzo, Tierras Mor- enas, 1,050 m., Provincia Guanacaste, Sep- tember 1993-'Gy\ pRodriguezs) IINBio CRI002131584 (1 6); Parque Nacional Santa Rosa, Provincia Guanacaste, 12 De- cember 1978—10 January 1979, D. H. Jan- zen, INBio CRI00204312 (1 3). Other material examined.—Guatemala, Cayuga, May, (no year on label), W. Schaus & W. Barnes (1 ¢, 1 2); Panama, Rio Trin- idad, March, May, 1912, A. Busck (1 d, 1 2: syntypes of P. mamella in USNM). Ceracanthia pseudopeterseni Neunzig and Solis, new species (Figs. 2, 13, 23-24, 44) Diagnosis.—The combination of the fol- lowing features will separate male Cera- canthia pseudopeterseni from other male Ceracanthia: forewing length about 11.0 mm. and with its anterior or costal half mostly pale and its posterior half dark (a dark diagonal band at about % out on costa VOLUME 104, NUMBER 4 AI = ca SAN \ 13 c=) Ss, 22S =< —: cA —, SSS Ee: ‘= SS: _= S Wz 843 oS = = =—— = <2= = Ly SS NS aN WS : » Nt SONG 15 14 —— Wess PIS aS ANNs) at i af OnE e 20 19 Figs. 11-20. Male, basal part of right antenna, frontal view. 11, Ceracanthia mamella. 12, C. soraella. 13, C. pseudopeterseni. 14, C. frustrator. 15, C. squamimagna. 16, C. squamifera. 17, C. schausi. 18, C. eugenieae. 19, C. alturasiana. 20, C. cornuta. divides most of the pale costal half into three strongly contrasting, chiefly white patches) and antenna with a sinus whose inner surface is covered with erect papillae (Bigs): Description.—Forewing length 10.0— 12.0 mm. Head: frons white; vertex a mix- ture of black and brownish red scales; labial palpus outwardly mostly white, sprinkled with brownish red scales and with patch of 844 brown near eye and at distal 4% of last seg- ment; maxillary palpus simple in both sex- es, pale basally, a mixture of black and brownish red distally; antenna of male with basal and distal ends of sinus of shaft only slightly produced mesially (sinus shallow); distal and subbasal end of sinus with dis- tinct spines; inner surface of sinus with erect papillae; short tuft of ochre scales at- tached to posterior base of sinus (tuft ex- tends about % length of sinus). Forewing of male with long, costal fold and with under- side possessing subcostal streak of brown- ish red scales along most of outer half of wing; wing above mostly white on anterior half and chiefly brown on posterior half; base mostly dark brown to black; a pair of black, diagonally-converging bands extend from costa to fuse with brown of posterior half; bands divide costal half of wing into three contrasting, mostly white patches; middle white patch with small, isolated, elongate, black streak at about mid-costa, and with a single black discal spot; a sprin- kling of brownish red scales chiefly on white patches. Hindwing simple in both sexes, pale brown, darker along margins. Male genitalia (Figs. 23-24) with uncus subtriangular, rounded apically; apical pro- cess of gnathos a slender, tapered, hooked rod; transtilla absent; juxta a thin plate with short triangularly-shaped, setiferous, lateral lobes; valva broad with distal half evenly rounded along costa, and with broadened, spinelike setae on inner surface; inner basal lobe of valva weakly developed; sacculus with apex distinctly lobed and with large, and small, tuft of very thin scales at base (large tuft slightly longer than length of val- va); vesica of aedoeagus with group of very small spines and group of slightly larger, more strongly sclerotized spines; vinculum distinctly longer than greatest width, and medially constricted. Female genitalia (Fig. 44) with signum developed as scobinate patch (part of patch sclerotized, teardrop shaped and with pronounced scobinations); ductus bursae shorter than corpus bursae. Types.—Holotype: ¢. Monte Verde, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Punt. Prov., Costa Rica, 15—16 May, 1980, D. H. Janzen & W. Hallwachs, INBio CRIO002044009, genitalia slide 4542 HHN (INBio). Paratypes: same collection data as holotype, except 8-10 Dec. 1978, D. H. Janzen, INBio CRI002044008, genitalia slide 4543 HHN (INBio) (1 ¢); C. R. Pun- tarenas Pr., Monteverde, 1,400 m., III-29- 31-92, UL & MV lights, S. McCarty & J. Powell, genitalia slides 4877, 4878 HHN (WEB) d dc, I 2) Volcan Stay Maria: Guat., July, (no year on label), Schaus and Barnes Coll. (USNM) (1 od); Oconeque, Carbaya, S. E. Peru, 7,000 ft., G. Okenden, genitalia slide 3596 CH (102, 128 USNM) (USNM) (1 ¢). Etymology.—The name pseudopeterseni refers to the misidentification of this species by Heinrich (1956) and subsequently by others as Megarthria peterseni (Zeller). Ceracanthia soraella (Druce), new combination (Figs. 3, 12, 25-26, 41, 45) Homoeosoma soraella Druce 1899:565. Drescoma drucella Dyar 1914:328. Drescomopsis subelisa Dyar 1919:62. Note.—The rationale for transferring so- raella from Drescomopsis to Ceracanthia was given following the list of synonyms of the genus Ceracanthia. Ceracanthia soraella superficially re- sembles a small version of Ceracanthia pseudopeterseni. The color and maculation of the forewing of both species are similar, and the inner surface of the male antenna of both possess erect papillae. However, Ceracanthia soraella has many diagnostic features that separate it from Ceracanthia pseudopeterseni, as well as from all other Ceracanthia. For example, the male and fe- male of the former has M, and M, com- pletely fused in the hindwing, and the fore- wing of the male with a small, flared cluster of scales at the base of costa (Figs. 3, 41), and on the underside, between the discal cell and vein 1A, an elongate pocket en- closing a scale tuft and a mass of modified VOLUME 104, NUMBER 4 845 Figs. 21-26. Male genitalia, ventral view. 21, Ceracanthia mamella (aedoeagus omitted). 22, C. mamella, aedoeagus. 23, C. pseudopeterseni (aedoeagus omitted). 24, C. pseudopeterseni, aedoeagus. 25, C. soraella (aedoeagus omitted). 26, C. soraella, aedoeagus. scales. Other Ceracanthia have M, and M, The male antenna has a very shallow, stalked and have males that lack the small easily overlooked, sinus. Heinrich (1956) costal cluster and the pocket with its tuft wrote “shaft of male slightly swollen at and scales. base, otherwise simple.’? However, a close 846 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Se =e as — > Ss LSS Figs. 27-32. Male genitalia, ventral view. 27, Ceracanthia frustrator (aedoeagus omitted). 28, C. frustrator, aedoeagus. 29, C. squamimagna (aedoeagus omitted). 30, C. squamimagna, aedoeagus. 31, C. squamifera (ae- doeagus omitted). 32, C. squamifera aedoeagus. look shows a slight depression enclosing a canthia soraella lacks an associated brush group of papillae, and a small spine at the — or tuft of scales. distal end of the sinus (Fig. 12). Unlike The most diagnostic feature of the male most of its congeners, the sinus of Cera- genitalia (Fig. 25) is the straplike transtilla VOLUME 104, NUMBER 4 with a pair of anteriorly directed arms. The corpus bursae of the female genitalia has a signum with a teardrop-shaped patch of scobinations (Fig. 45). Costa Rican material examined.—Juan Vinas, June, November, (no year on label), W. Schaus & W. Barnes (1 6); Sitio, June, (no year on label), (1 2); Quepos, Parque Nacional Manuel Antonio, 120 m., Prov- incia Puntarenas, November, 1990, G. Var- ela & R. Zuniga, INBio CRIOO0O180596 (1 6); Cerro Tortuguero, 0-120 m., Parque Nacional Tortuguero, Provincia Limon, March, 1992, May 1991, R. Delgado, IN- Bio CRI000853321, J. Solano, INBio CRIO01398889, CR INBio 1001398587, INBio 1000358990 (4 3). Other material examined.—Mexico, Ja- lapa, (no date on label), M. Trujillo (3 @; syntypes of H. soraella in BMNH), Cor- doba, May, 1908, FE K. Knab (1 @); Belize, San Ignacio, April, 1989, L. C. Dow (1 &), Mountain Pine Ridge, May, 1990, L. C. Dow (1 6, 1 2); Guatemala, Cayuga, April, June (2 @; syntypes of D. subelisa in USNM); Panama, Paraiso, May, 1923, R. C. Shannon (1 ¢), Porto Bello, March, Oc- tober (1 3, 1 2; syntypes of D. drucella in USNM), Rio Trinidad, March, 1912, A. Busck (1 2); Ecuador, Zaruma, 1916, F W. Rohwer (1 d¢); French Guiana, St. Jean de Maroni, no date (1 d); Brazil, Isla Sant. Catarina, June, 1935, EF Hoppman (1 9°), Linhares, January, 1998, V. O. Becker (1 3), Cacaulandia, November, 1991, V. O. Becker (1 6d), Ibateguara, March, 1994, V. O. Becker (1 3). Amsel (1956, 1957), prob- ably correctly, reported this species from Maracay, Venezuela, but we have been un- able to verify his identification. Ceracanthia frustrator (Heinrich), new combination (Figs. 4, 14, 27—28, 42) Megarthria frustrator Heinrich 1956:87. The color and maculation of the forewing of Ceracanthia frustrator greatly resembles the appearance of the forewing of most oth- 847 er Ceracanthia. The male of Ceracanthia frustrator, however, has a feature not seen on males of other species belonging to the genus (or in the subfamily!): a group of scales protrudes beyond the costa for over ¥y the length of the forewing (Figs. 4, 42). The male antenna (Fig. 14) has a well developed basal sinus with a_ triangular cluster of scales (possibly enclosing a spine) protruding from its base and a low spine associated with its distal end. The in- ner surface of the sinus is covered with ap- pressed scales, and a brush of scales is pre- sent, attached to the posterior base of the sinus (the brush, composed of narrow scales extends slightly beyond the length of the sinus). The male genitalia (Figs. 27-28) lack a transtilla, and have a large scale tuft, and a small secondary scale tuft, attached to the base of the sacculus (the large tuft is slight- ly shorter than the length of the valva). The vinculum is broad and not constricted me- dially. We have not been able to associate with certainty females with males, and, there- fore, the appearance of the female genitalia is unknown. Costa Rican material examined.—Juan Vinas, February, (no year on label), W. Schaus & W. Barnes (1 d; holotype of M. frustrator in USNM); 8 km. N. Vara Blan- ca, 1,400 m., Provincia Alejuela, 25 March, 1992, McCarty & Powell (1 ¢, 2 @). Ceracanthia squamifera (Heinrich), new combination (Figs. 5, 16, 31-32, 46) Megarthria squamifera Heinrich 1956:87. Ceracanthia squamifera is best recog- nized by examining the antenna of the male (Fig. 16). A strong sinus is present at the base of the shaft, and there is a well devel- oped brush of dark scales attached to the posterior base of the sinus and lying within the sinus. The brush extends almost the en- tire length of the sinus. 848 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 33-38. Male genitalia, ventral view. 33, Ceracanthia cornuta (aedoeagus omitted). 34, C. cornuta, aedoeagus. 35, C. eugenieae (aedoeagus omitted). 36, C. eugenieae, aedoeagus. 37, C. schausi (aedoeagus omitted). 38, C. schausi, aedoeagus. The male genitalia (Fig. 31) have a par- ticularly broad valva with the costa of its distal half noticeably distended. The large brush of the sacculus is longer than the length of the valva. The corpus bursae of the female genitalia has a signum with a teardrop-shaped patch of scobinations (Fig. 46). Costa Rican material examined.—Mount Pods, May, (no year on label), W. Schaus VOLUME 104, NUMBER 4 849 p> fa eZ gig 2 Za) [ips vy LEDGER ZZ LOS NPIL ILD A a" WIA igs y ye: pcolgeeiee Sah ee oe =a EBL, lg; ere eee BIOS ga Ro See SS a en ee as SAF oY extn aes ~ = S262 ee —S EY IOS FP VOD a Se OQ ILO ES SSN, 42 Figs. 39-42. Male genitalia, ventral view, and scale cluster or scale ridge on costa of male right wing, dorsal view. 39, Ceracanthia alturasiana (aedoeagus omitted). 40, C. alturasiana, aedoeagus. 41, C. soraella. 42, C. frustrator. Ceracanthia squamimagna Neunzig and Solis, new species (igs 26715, 29-3047) & W. Barnes (1 6; holotype of M. squam- ifera in USNM); 9 km. S. Sta. Cecilia, Estacion Pitilla, 7OO m., Provincia Guanacaste, July, 1991, 4-18 December, 1991, January 1995, C. Moraga, INBio CRI1000527303, INBio CRI000599235, ER INBio 1002001987, INBio CRIO01627819 (4 ¢) ; Monteverde, Prov- Diagnosis.—The male of Ceracanthia squamimagna is among the more easily rec- ognized members of the genus. A large, pale tuft of scales is associated with the si- incia Puntarenas, 10—11 December, 1979, D. H. Janzen, INBio CRIO02044007, INBio CRI002044011 (2 oC). nus at the base of the shaft of the antenna (Fig. 15). Some of the scales forming the tuft are very broad (obovate), and the tuft (o/) Nn i=) extends about as far beyond the end of the sinus as the length of the sinus. The distal end of the sinus is also peculiar, in that it is strongly produced and pointed mesially, and is roughly covered with many slender setae. Description.—Forewing length 7.5—8.0 mm. Head: frons white washed with brown- ish red; vertex mostly brownish red, suf- fused with black anteriorly and white pos- teriorly; labial palpus outwardly white with patches of brownish red; maxillary palpus simple in both sexes, same color as labial palpus; antenna of male (Fig. 15) with well developed sinus at base of shaft; distal end of sinus strongly produced and pointed me- sially, and roughly covered with many slen- der setae; inner surface of sinus with ap- pressed scales; large, pale, scale tuft at- tached to posterior base of sinus (tuft ex- tends distally about as far beyond end of sinus as length of sinus; some scales form- ing tuft obovate). Forewing: with concavity in basal % of costa; underside with basal subcostal patch of reddish brown; wing above mostly white on anterior half and chiefly brown on posterior half; base mostly dark brown to black; a pair of diagonally converging black bands extend from costa to fuse with brown of posterior half; bands divide costal half of wing into three con- trasting, mostly white patches (most distal patch small, obscure in some specimens); middle white patch with small isolated, elongate, mostly red patch at about mid- costa, and with pair of black discal spots; a sprinkling of red to brownish red _ scales chiefly on white patches. Hindwing: under- side with basal, ochre, subcostal patch; wing above pale brown, darker along mar- gins. Male genitalia (Figs. 29-30) with un- cus broadly rounded at apex; apical process of gnathos a slender, tapered, slightly hooked rod; transtilla absent; juxta a thin, triangular plate with short, setiferous, lat- eral lobes; valva broad, with distal % evenly rounded along costa, and with broadened, spinelike setae on inner surface; inner basal lobe of valva moderately well developed; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sacculus with apex distinctly lobed and with large, dark tuft of thin scales at base; no small secondary tuft at base of sacculus; vesica of aedoeagus with group of very small spines and group of slightly larger, more strongly sclerotized spines; vinculum longer than greatest width, and medially constricted. Female genitalia (Fig. 47) with signum developed as scobinate patch (part of patch sclerotized, teardrop-shaped, and with pronounced scobinations); ductus bur- sae shorter than corpus bursae. Types.—Holotype: ¢. Quepos, 120 m., P. N. Manuel Antonio, Prov. Punt., Costa Rica, G. Varela & R. Zuniga, Nov. 1990, L- S-370900, 449800, INBio CRIO00228018, genitalia slide 4540 HHN (INBio). Para- types: Cerro Tortuguero, P. N. Tortuguero, 0-100 m., Prov. Limon, Costa Rica, J. So- lano, Abr. 1991, L-S-285000, 588000, IN- Bio CRI0004443 14, INBio CRI000444053, genitalia 444314 MC, 444053 MC (INBio) (1 6,1 2); Cerro Tortuguero, 0-120 m., P. N. Tortuguero, Prov. Limon, Costa Rica, Abr., May 1991, J. Solano, L-S-285000, 588000, INBio CRI001375039, INBio CRIO01398885, INBio CRI001398904, genitalia slides 4525 HHN, 4526 HHN, 4538 HEIN (USNM, NCSU)"@ 6252): Etymology.—The name squamimagna is a combination of the Latin squama (scale) and the Latin magna (large) in reference to the large tuft of scales associated with the sinus of the male antenna. Ceracanthia cornuta Neunzig and Solis, new species (Figs. 7, 20, 33-34) Diagnosis.—As with many Ceracanthia, the best diagnostic feature of Ceracanthia cornuta is found on the male antenna. The distal end of the well developed sinus, at the base of the shaft, is produced mesially into a large, smoothly-scaled, conical, slightly-hooked element (Fig. 20). Description.—Forewing length 11.0— 11.5 mm. Head: frons ochreous white; ver- VOLUME 104, NUMBER 4 Figs. 43-45. 45, C. soraella. tex a mixture of pale brown, brown and dark brown scales; pale brown between an- tennae; labial palpus outwardly with seg- ments mostly white at base and chiefly pale brown and pale reddish brown distally: maxillary palpus simple in male, white and pale brown; antenna of male with sinus in base of shaft, and with basal and distal ends of sinus distinctly produced mesially (distal end enlarged into a conical slightly-hooked process); inner surface of sinus of antenna with appressed scales; brown tuft of scales attached to posterior base of sinus (tuft ex- tends almost to tip of hooked, conical pro- cess); part of outer margin of antenna ser- rate, particularly just above sinus. Forewing of male simple, without costal fold, costal concavity or sex-scaling; wing above most- } 44 Female genitalia, anterior part, ventral view. 43, Ceracanthia mamella. 44, C. pseudopeterseni. ly white on anterior half and mainly brown and black on posterior half; base mostly black; pair of black diagonally converging bands extend from costa to fuse with brown of posterior half; bands divide costal half of wing into three contrasting, mostly white patches; middle white patch with small, iso- lated, elongate, black and brownish red patch at about mid-costa, and with pair of black and brownish red discal spots; a sprinkling of brownish red scales, chiefly on white patches. Hindwing of male simple, pale brown, darker along margins. Male genitalia (Figs. 33—34) with uncus broad at base; apical process of gnathos a slender, tapered rod; transtilla present, consisting of a transverse, arched, spined band; juxta a thin plate with short, rounded lateral, seti- Figs. 46-48. 48, C. alturasiana. ferous lobes; valva broad, with distal half of costa enlarged; inner basal lobe of valva with low, adjunct, eliptical protuberance; sacculus with apex distinctly lobed, and with large and small, pale tufts of very thin scales at base (large tuft slightly longer than length of valva); vesica of aedoeagus with elongate cluster of microspines, and a sec- ond group of slightly larger, more sclero- tized spines; vinculum longer than greatest width, with medial constriction. Female un- known. Types.—Holotype: d. Fila Orosilito, Est. Pitilla, 9 km. S. Santa Cecilia, Prov. Guana., Costa Rica, 800—1,000 m., Abr. 1995, C. Moraga, LN 328650, 378600 #4828, INBio CRI002204210 genitalia PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Female genitalia, anterior part, ventral view, 46, Ceracanthia squamifera. 47, C. squamimagna. slide 4516 HHN (INBio). Paratype: Est. La Casona, 1,520 m., Res. Biol. Monteverde, Prov. Puntarenas, Costa Rica, Jul. 1991, N. Obando, L-N 253250, 449700, INBio CRI000948482, genitalia slide 948482 MC (INBio) (1 3). Etymology.—tThe large, slightly-hooked, conelike protuberance at the distal end of the sinus of the male antenna is responsible for the specific epithet cornuta. Ceracanthia schausi (Heinrich), new combination (Figs. 8, 17, 37—38) Megarthria schausi Heinrich 1956:87. Heinrich (1956) separated Ceracanthia schausi (as Megarthria schausi) from other VOLUME 104, NUMBER 4 similar species by its unusual transtilla con- sisting of a sclerotized plate with thin, short arms projecting from its lower (anterior) corners. Our study has discovered another Ceracanthia with a similar transtilla (see next species), so a combination of features now needs to be used to identify Ceracan- thia schausit. In addition to the transtilla, Ceracanthia schausi has a broad, rounded vinculum and a strong tuft of scales at the base of the sacculus. The male antenna (Fig. 17) has a well developed sinus at the base of its shaft. The sinus has a basal spine, and its inner surface is covered with pale, appressed scales. A tuft of mostly dark brown, to black scales, is present, extending the length of the sinus. We have been unable to associate with certainty females of Ceracanthia schausi with males, and, therefore, the appearance of the female genitalia is unknown. Costa Rican material examined.—Juan Vinas, January, (no year on label), W. Schaus (1 6; holotype of M. schausi in WISIN): 9 kimi. S= Stay, Cecilia, 700; m.; Estacion Pitilla, Provincia Guanacaste, 2-15 May, 1992, C. Moraga, INBio CRIO00405603, INBio CRI000599224 (1 3, 1 2); Estacion La Casona, 1,520 m., Re- serva Biologica Monteverde, Provincia Puntarenas, October, 1992, N. Obando, CR INBio 1000818108 (1 3); Monteverde, 4,400’, Provincia Puntarenas, 2 September LOSS: Cs Va Covelly In, so): Other material examined.—Belize, 1,000 Foot Falls, Mountain Pine Ridge, May, 1990, L. C. Dow (1 do, 1 2), Butler Line, Mountain Pine Ridge, May, 1990, L. C. Dow (1 6), San Ignacio, June, 1990, L. C. Dow (1 o). Ceracanthia eugenieae Neunzig and Solis, new species (Figs. 9, 18, 35-36) Diagnosis.—Based on the male genitalia, Ceracanthia eugenieae is closely related to Ceracanthia schausi. Both have peculiar lobes that protrude inwardly from the base of the valva, and both have a transtilla con- 853 sisting of a plate with a pair of curved low- er (anterior) arms. But, Ceracanthia eugen- ieae, most noticeably, differs from Cera- canthia schausi, and all other Ceracanthia, in lacking a tuft of scales at the base of the sacculus (Fig. 35). Description.—Forewing length 8.0—8.5 mm. Head: frons brownish white; vertex reddish brown, dark brown near eye; labial palpus outwardly with basal 2/3 of each segment mostly white, or brownish white, and with distal part white, or brownish white, red, or brownish red, and brown, or black; maxillary palpus simple, white, or brownish white; antenna of male (Fig. 18) with strongly developed sinus; basal and distal ends of sinus without spines (basal end with small, black mound of very short? sensilla); inner surface of sinus covered with appressed setae; tuft or brush of scales attached to posterior base of sinus (tuft or brush extends length of sinus). Forewing of male with long costal fold; underside with subcostal streak (dark with pale center ba- sally becoming brownish red near distal part of costal fold); wing above mostly white on anterior half and chiefly brown on posterior half; base mostly dark reddish brown and black; a pair of black (or dark brown), diagonally-converging bands ex- tend from costa to fuse with brown of pos- terior half; bands divide costal half into three contrasting, mostly white patches; dis- tal patch small; middle patch with small, isolated, elongate brownish red and black patch at about mid-costa, and with two brownish red and black discal spots; a sprinkling of red or brownish red scales chiefly on white patches. Hindwing simple, pale brown, darker along margins. Male genitalia (Figs. 35—36) with uncus subtrian- gular, rounded apically; apical process of gnathos a short, slender, tapered, slightly- hooked rod; transtilla a square plate with pair of curved arms extending from its low- er (anterior) corners; juxta a thin plate with short, setiferous lateral arms; valva broad with distal half slightly produced inwardly, and with broadened, spinelike setae on in- 854 ner surface; inner basal lobe of valva strongly developed (as in Ceracanthia schausi, but with tips overlapping); vesica of aedoeagus with elongate cluster of mi- crospines and a second cluster of slightly larger, more heavily sclerotized spines; sac- culus with apex lobed; base of sacculus without large tuft of setae or scales (also without small secondary tuft present on some Ceracanthia); vinculum longer than greatest width, without medial constriction, with base laterally lobed. Female unknown. Types.—Holotype: d. Estacion Pitilla, 9 km. S. de Santa Cecilia, Prov. Guana., Cos- ta Rica, 700 m., AGO 1995, C. Moraga, Luz, L-N-329950 380450, #6207, CR IN- Bio 1002309873, genitalia slide 4539 HHN (INBio). Paratype: Turrialba, Costa Rica, 600 m., 10.1.1972, V. O. Becker, genitalia slide 4922 HHN (VOB) (1 ¢). Etymology.—We are pleased to name this species in honor of Eugenie Phillips- Rodriguez at INBio. Her enthusiasm in pro- viding specimens, and assisting in many other ways, greatly added to the success of our study. Ceracanthia alturasiana Neunzig and Solis, new species (Figs. 10, 19, 39—40, 48) Diagnosis.—Like Ceracanthia mamella, the forewing of Ceracanthia alturasiana does not resemble the forewings of other species in the genus. However, the latter species cannot be confused with the former because the color and maculation of the forewings of the two species are very dif- ferent. An obvious feature is a strong post- medial line on the wing of Ceracanthia al- turasiana (Fig. 10); this distinct line is ab- sent from Ceracanthia mamella, as well as from all other Costa Rican Ceracanthia. Description.—Forewing length 8.0—8.5 mm. Head: frons ochre; vertex pale reddish ochre; labial palpus outwardly with basal segment white and pale ochre, other two segments ochre with brown distally; max- illary palpus simple in both sexes, ochre; antenna of male with basal sinus with small PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON spine at basal end of sinus and small spine at distal end of sinus (two additional spines beyond distal end of sinus); inner surface of sinus with appressed scales; small, short weakly-formed tuft of scales attached to base of sinus (tuft extends about % length of sinus). Forewing: simple, without costal concavity, costal fold, or other modifica- tions or embellishments; wing above basal- ly and subbasally dark brown to black, sprinkled with brownish red, and with dust- ing of ochre, particularly along costa; an- temedial line fragmented, bur rather dis- tinct, ochre; postmedial line well devel- oped, white and ochre; medial area with ground color dark brown to black, with dusting of brownish red scales, and with distinct patch of ochre surrounding discal spots; discal spots dark brown to black and brownish red. Hindwing simple in both sex- es, mostly pale brown, darker along mar- gins. Male genitalia (Figs. 39—40): with un- cus subtriangular, rounded apically; apical process of gnathos a slender, long, slightly- tapered, hooked rod; transtilla present, rep- resented by a thin, sclerotized, medial band; juxta a thin plate with short, setiferous, lat- eral lobes; valva broad with distal half slightly produced along costa, and with broadened, spinelike setae on inner surface; inner basal lobe of valva weakly developed: sacculus with apex narrowly lobed, and with yellowish, basal tuft of thin scales (tuft slightly longer than length of valva); vesica of aedoeagus with cluster of microspines and a well developed group of medium- sized spines; vinculum longer than greatest width, and not medially constricted. Female genitalia (Fig. 48) with ductus bursae short- er than corpus bursae; corpus bursae with patch of many, small spines posteriorly that extends a short distance into ductus bursae, and with a more anterior sclerotized, scob- inate patch on its right side (when viewed ventrally) bearing an imbricate, hooked spine that extends mostly laterally; ductus seminalis attached to corpus bursae near junction of ductus bursae and corpus bur- sae. VOLUME 104, NUMBER 4 Types.—Holotype: 6. C. R.: Estac. Biol. Las Alturas, 1,550 m., 12 air km. NE San Vito, Punt. Prov., I-22/24—1993, J. Powell, at blacklight, genitalia slide 4857 HHN (UCB). Paratypes: same collection data as holotype, genitalia slide 4858 HHN (UCB, NCSU) (3 2); C. R.: Puntarenas Pr., Mon- teverde, 1,400 m., III-30-1992, Powell (WEB) C:2): Etymology.—The specific epithet is de- rived from the type locality (Las Alturas), and the Latin suffix -ana (meaning “‘be- longing to’’). ACKNOWLEDGMENTS We thank the many parataxonomists at INBio for collecting, spreading and labeling most of the specimens we studied. Also, at the same institution, Mario Comacho pre- pared slides of genitalia, and Eugenie Phil- lips arranged for the loan of material. Other study specimens were provided by Jerry A. Powell, UCB, and Vitor O. Becker, Brasilia, Brazil. Robert L. Blinn, NCSU, made the habitus photographs. LITERATURE CITED Amsel, H. G. 1956. Microlepidoptera Venezolana I. Boletin de Entomologia Venezolana 10: 1—336. 855 . 1957. Microlepidoptera Venezolana II. Bole- tin de Entomologia Venezolana 10: Tables I-CX. Druce, H. 1899. Lepidoptera—Heterocera. /n Godman, FD. and O. Salvin, Biologia Centrali- Americana; or Contributions to the Knowledge of the Fauna and Flora of Mexico and Central America. Zool- ogy: Insecta, Vol. 2, Taylor and Francis, London, 622 pp. Dyar, H. G. 1914. Report on The Lepidoptera of the Smithsonian Biological Survey of the Panama Ca- nal Zone. Proceedings of The United States Na- tional Museum 47: 139-350. 1919. Some Tropical American Phycitinae. Insecutor Inscitiae Menstruus 7: 40—63. Heinrich, C. 1956. American Moths of the subfamily Phycitinae. United States National Museum Bul- letin 207: 1-581. Neunzig, H. H. 1994. New genera and species of Mex- ican Phycitinae (Lepidoptera: Pyralidae). Proceed- ings of the Entomological Society of Washington 96: 357-366. . 1996. New Species of Phycitinae (Lepidop- tera: Pyralidae) from the Dominican Republic. Proceedings of the Entomological Society of Washington 98: 774-801. Neunzig, H. H. and L. C. Dow. 1993. The Phycitinae of Belize (Lepidoptera: Pyralidae). North Carolina Agricultural Research Service Technical Bulletin 304: 1-133. Ragonot, E. L. 1893. Monographie des Phycitinae et des Galleriinae. Jn Romanoff, N. M., ed. Mémo- ires sur les Lépidopteres, Vol. 7, Imprimerie Gé- nérale Lahure, Paris, 658 pp. Zeller, P. C. 1881. Columbische, Chilonden, Crambi- den und Phycideen. Horae Societatis Entomolo- gicae Rossicae 16: 154—256. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 856-862 A REVIEW OF PTOUS CHAMPION (COLEOPTERA: CURCULIONIDAE: CRYPTORHYNCHINAE) HENRY A. HESPENHEIDE AND Louis M. LAPIERRE Department of Organismic Biology, Ecology and Evolution, University of California, Los Angeles, CA 90095-1606, U.S.A. (e-mail: henryh@biology.ucla.edu) Abstract.—Three new species are described in the genus Prous, P. cecropiae, n. sp., P. woldai, n. sp., and P. liebermanorum, n. sp., and P. otidocephalinus Champion is rede- scribed. These species and at least three other undescribed forms can be separated into two groups. The species are illustrated and a key is provided. Two species have been reared from members of the plant genus Cecropia (Cecropiaceae), and a third has been collected on Cecropia as adults. Members of the genus Prous are hypothesized to be part of a mimicry complex possibly based on the weevil genus Myrmex as Mullerian models. Key Words: Cecropia, mimicry, Myrmex The genus Ptous was described by Champion in 1906 for a single specimen from Nicaragua. In recent years collecting, ecological sampling, and rearing have yielded a number of additional specimens of at least seven different species. The pres- ent paper describes four of these as part of a more general study of insects associated with the plant genus Cecropia by one of us (L. M. LaPierre, in preparation). The following collection acronyms are used throughout the text: BMNH: The Nat- ural History Museum, London, England: CHAH: Henry A. Hespenheide, University of California, Los Angeles, CA, U.S.A.; CMNC: Canadian Museum of Nature, Ot- tawa, Canada; CWOB: Charles W. O’Brien, Tallahassee, FL, U.S.A.; INBC: Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica; STRI: Smithsonian Tropical Research Institute, Ancon, Pana- ma; USNM: National Museum of Natural History, Smithsonian Institution, Washing- ton DGS UES 2A: Ptous Champion 1906: 632. Type species.—Ptous otidocephalinus Champion, by monotypy. Champion separated the genus Prous on a combination of characters that make it very distinctive in the Cryptorhynchinae but which produce a strong superficial re- semblance to members of the genus Myr- mex (= Otidocephalus) in the Myrmecinae, a resemblance that is probably mimetic (see below). Species are relatively slender, nar- rowly obovate or ovate, and more or less strongly shining black and glabrous above with a relatively inconspicuous pattern of stripes and fasciae of white setae on the el- ytra. The third elytral interval is usually raised and denticulate near the base of the elytra, and the first (sutural), fifth and sev- enth intervals also may be raised or faintly carinate. The eyes are relatively large and narrowly separated along the middle, as in the Zygopinae, and the ocular lobes on the pronotum are poorly developed. The legs are relatively long and slender. Champion VOLUME 104, NUMBER 4 placed Prous between the genera Trachalus and Cryptorrhynchus but did not discuss its relationships with either. It is interesting that there are similarities of Prous with Co- phes gibbus Champion, which also has been reared from Cecropia. Characters.—Species in the genus fall into two groups based on elytral pattern and form of the posterior tibiae. One group of species possesses an elytral pattern of a more or less complete lateral stripe on the sixth interval and a more or less well de- veloped transverse subapical fascia, has the apical half of the first (sutural) interval raised and denticulate, and has the posterior tibiae terete and straight. The other group of species has an elytral pattern of broad oblique fasciae from the base of fifth elytral interval to the suture of the elytra, without a subapical fascia and with or without a submarginal stripe, and posterior tibiae that are flattened and weakly arcuate on their posterior margins. Species within these two groups differ in details of the setal patterns, the nature of the surface sculpture, and male genitalia. KEY TO SPECIES OF PTOUS 1. Elytral pattern of oblique fasciae from base of fifth interval to suture at middle of elytron and then continuing parallel to suture, with lateral stripe, no subapical fascia; first (sutural) inter- val not raised or denticulate; posterior tibia flat- tened and weakly arcuate on posterior margin: @ostagRical sso. = 2 — Elytral pattern of more or less complete stripe P. liebermanorum, n. sp. of setae on sixth interval parallel to lateral mar- gin and transverse subapical fascia; apical half of the first (sutural) interval raised and dentic- ulate; posterior tibia mostly terete and straight, bo laterally compressed only at base ......... ie) Posterior /; of elytra almost completely cov- ered by setae; sides of pronotum with setae to top of ocular lobes; tooth on anterior femora =< 03 mm) long, acute; Panama, -../..5.. Seance ae pee eRe Bene Uae P. woldai, n. sp. — Elytra with posterior transverse subapical fas- cia but with large glabrous area; sides of pron- otum with setae only below ocular lobes; tooth — on anterior femora > 0.1 mm long, acuminate 3. Stripe of setae on sixth elytral interval incom- plete; first (sutural) interval with setae only at apex and at posterior transverse fascia; fifth and seventh intervals not raised; Panama Oto Ray ear P. cecropiae, Nn. sp. - Stripe of setae on sixth elytral interval com- plete; apical half of first (sutural) interval with stripe of setae; fifth and seventh intervals raised, faintly carinate; Nicaragua to Panama P. otidocephalinus Champion Ptous liebermanorum Hespenheide and LaPierre, new species (Figs: 1,4) Holotype male.—Black, moderately shining, relatively slender, slightly obovate; above glabrous except oblique stripe of small white setae from base of fifth elytral interval to third interval at about basal %, continuing to apex, on first and second ely- tral intervals for middle 4%, for entire length of ninth interval, basal % of tenth, apical 7, of eighth, and apical ¥, of seventh inter- val; stripes of white setae on pronotum an- terior to base of elytral stripes parallel to lateral margin reaching almost to apex; be- neath and at base of rostrum with moder- ately dense white setae, somewhat con- densed above anterior coxae, but glabrous above base of ocular lobes; legs with very small white setae, denser on dorsal and ven- tral surfaces; 5.0 mm long, 1.9 mm wide. Head finely, densely punctate above, with eyes large, rather narrowly separated, ros- trum 1.3 mm long, antennae inserted just above middle of rostrum. Pronotum 1.4 mm long slightly wider than long, widest just be- fore base, sides weakly arcuate, weakly con- vex in cross section, nearly flat in lateral view, moderately densely, finely punctate. Elytron 3.2 mm long; somewhat matte, reg- ularly convex, more weakly so in lateral view, basal ¥; of third interval somewhat raised and minutely denticulate. Metaster- num and first abdominal sternite very shal- lowly, broadly concave. Legs long, anterior femur with acute triangular tooth, posterior femur with tiny acute tooth, Aedeagus as in Higees: Allotype female.—As holotype, except metasternum and base of first abdominal sternite weakly convex, 6.8 mm long. 858 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1—4. woldai. 3, P. cecropiae. 4, P. otidocephalinus. Holotype—COSTA RICA: Heredia Pr., Est. Biol. La Selva, 50-150 m, 10°26'N 84°01'W, 09.11.1998, L.M. LaPierre, #98.277 (INBC). Allotype.—Same data as holotype, ex- cept #98.277-1 (INBC). Paratypes.—COSTA RICA: Alajuela Pr., 20 km S Upala, 01-20.11.1990, ED. Parker (1, CWOB); Heredia Pr., same data as ho- lotype, except #98.278.1; 24.01, 14.02, 03.2000, L.M. LaPierre, #00.649, 650, 651, reared from Cecropia obtusifolia wood (8, BMNH, CHAH, CWOB, USNM). Discussion.—Specimens vary in size Dorsal and lateral habitus and posterior tibia of species of Prous: 1, P. liebermanorum. 2, P. from 4.1—6.8 mm in length (mean = 5.52 mm, N = 12). In addition to this species, material has been seen that appears to rep- resent three additional species of Prous in the group with oblique elytral fasciae. Two of these species are from Panama, one is from Bolivia. Each is represented by only a single female specimen, all in the collec- tion of Charles W. O’Brien. We prefer to defer description of these until more mate- rial, including males, becomes available. This species is dedicated to Drs. Milton and Diana Lieberman in warm appreciation of their generosity and advice to LML VOLUME 104, NUMBER 4 7h Figs. 5-8. otidocephalinus. while he worked at San Luis Biological Sta- tion. Ptous woldai Hespenheide and LaPierre, new species (Rigs: 2; 16) Holotype male.—Black, moderately shining, relatively slender, slightly obovate; above glabrous except stripe of white setae on sixth elytral interval from base to apex and on apical 4 of all other intervals; some- What broad and irregular stripes of white setae on pronotum interior to base of elytral stripes, parallel to lateral margin reaching almost to apex and few white setae along mideline of pronotum at apex; white setae 0.5mm Male aedeagus of species of Prous. 5, P. liebermanorum. 6, P. woldai. 7, P. cecropiae. 8, P. at base of rostrum and along midline of head above eyes; moderately dense white setae on sides of pronotum to top of ocular lobes, and on meso- and metasterna; ab- dominal sternites glabrous; legs with uni- form white setae, denser on femora; 4.4 mm long, 1.7 mm wide. Head finely, densely punctate above, with eyes large, rather narrowly separated, rostrum 1.0 mm long, antennae inserted just above middle of rostrum. Pronotum 1.25 mm long slightly wider than long, widest at basal %, sides arcuate, weakly convex in cross section and in lateral view, moderate- ly densely, finely punctate. Elytron 2.8 mm long; rather strongly shining, evenly convex 860 in lateral view, basal % of third interval and apical % of first (sutural) interval raised and denticulate, fifth and seventh intervals raised, fifth more strongly so; coarsely punctate. Metasternum and base of first ab- dominal sternite very weakly, broadly con- cave. Legs moderately long, anterior femur with small acute triangular tooth, posterior femur with tiny acute tooth; posterior tibia terete, basally somewhat arcuate. Aedeagus as in Fig. 6. Allotype female.—As holotype, except metasternum and base of first abdominal sternite weakly convex, 5.1 mm long. Holotype. —PANAMA, Barro Colorado Is:, UV trap 1G im high); 09:07.19775 Eb Wolda (CWOB). Allotype.—Same data as holotype, ex- cept 13.04.1977 (CWOB). Paratypes.—Same data as holotype, ex- cept Ss 22101976. 13045 wis.a27 05, OS-O7 1977, 212-08 1978. 27-09 M9 Sas 08.08.1983 (8, CWOB); same data as ho- lotype, except UV trap 3 (26 m high), 21, 28:06; 14.075 525-9, 155 nko10s;, 20-09, C2009 19762 UI035) 18h 2a. 29.06% 19° 2006; 07-07 102: 7977. 24.045 07; 2ee0 55. 25-0604. MIRO. 230 7S, 10.091979, 04—05, 09.11.1980 (31, BMNH, CHAH, CWOB, STRI, USNM). Discussion.—Specimens vary in size from 3.6—5.0 mm in length (mean = 4.22 mm, N = 40). This species is named for Henk Wolda whose extensive sampling of insects on Barro Colorado Island (Wolda et al. 1998) and elsewhere in Panama has been an im- portant early step in the understanding of the biodiversity and seasonal phenology of Neotropical insects. Ptous cecropiae Hespenheide and LaPierre, new species (Figs. 3, 7) Holotype male: Black, strongly shining, relatively slender, narrowly ovate; above glabrous except stripe of white setae on basal ¥; of sixth elytral interval and apical 4% of second and fifth interval, and oblique PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON transverse fascia at apical ¥;; small patches of white setae on pronotum just anterior to base of elytral stripes; beneath and at base of rostrum with moderately dense white se- tae, somewhat more condensed on sides, but glabrous above base of ocular lobes and on abdominal sternites; legs with very small white setae on tibiae and dorsal por- tions of middle and posterior femora; 5.0 mm long, 1.9 mm wide. Head finely, densely punctate above, with eyes large, rather narrowly separated, rostrum |.3 mm long, antennae inserted just above middle of rostrum. Pronotum 1.35 mm long slightly wider than long, widest at basal 4%, sides rounded-angulate, weakly convex in cross section, nearly flat in lateral view, sparsely, finely punctate. Elytron 3.35 mm long; evenly convex, more weakly so in lateral view, basal 4 of third interval and apical 4 of first (sutural) interval raised and denticulate, fifth interval slightly raised. Metasternum and first abdominal sternite deeply, longitudinally concave. Legs long, anterior femur with long, acuminate tooth, posterior femur with shorter acuminate tooth; posterior tibia terete, basally some- what laterally compressed. Aedeagus as in Fig. Allotype female.—As_ holotype, except metasternum flat and base of first abdomi- nal sternite weakly convex, 6.8 mm long. Holotype —PANAMA, Chiriqui, Fortu- na (08°44’N 82°15’W) 1,050 m, UV trap, 19.10.1976 (CWOB). Allotype.—Same data as holotype, ex- cept 31.10.1977 (CWOB). Paratypes. PANAMA, Cocle Prov., Cerro Goital, 10-12.06.1985, E.G.Riley & D. Rider (1, CWOB); Panama Prov., Cerro Campana, 05.07.1974, C.W. & L. O’Brien & Marshall (1, CWOB), Cerro Campana, 850 m, 08°40'N 79°56’W, 29.05.1970, H.A. Hespenheide, on Cecropia (1, CHAH), 29.04.1973, H.A. Hespenheide (1, BMNH), 08.02.1972, W. Bivin (1, CMNC). Discussion.—Specimens vary in size from 3.9—6.3 mm in length (mean = 5.18 mm, N = 7). VOLUME 104, NUMBER 4 The specific refers to the adult and pu- tative larval host of this and other Prous, species of plant genus Cecropia. Ptous otidocephalinus Champion 1906: 633. (Figs. 4, 8) Diagnosis.—Female: Black, strongly shining, relatively slender, narrowly ovate; above glabrous except complete stripe of white setae on sixth elytral interval, on ba- ses of fourth and fifth intervals, and along apical % of suture, and broad oblique trans- verse fasciae at apex and apical /; narrow lines of white setae on pronotum anterior to base of elytral stripes; beneath and at base of rostrum with moderately dense white se- tae, somewhat more condensed on sides, but glabrous posterior to ocular lobes and on abdominal sternites; legs with very small white setae on tibiae and dorsal por- tions of femora; 5.4 mm long, 2.1 mm wide. Head finely, densely punctate above, with eyes large, rather narrowly separated, rostrum 1.5 mm long, antennae inserted just above middle of rostrum. Pronotum 1.4 mm long slightly wider than long, widest at bas- al %4, sides rounded-angulate, weakly con- vex in cross section, nearly flat in lateral view, sparsely, finely punctate. Elytra 3.7 mm long; evenly convex, more weakly so in lateral view, basal /; of third interval and apical %4 of first (sutural) interval raised and denticulate. Metasternum and first abdomi- nal sternite weakly convex. Legs long, an- terior femur with acute tooth, posterior fe- mur with very small acute tooth; posterior tibia terete, basal 4% somewhat laterally compressed. Male.—As female, except metasternum and base of first abdominal sternite deeply, longitudinally concave. Aedeagus as in Bis: 8: Specimens examined.—COSTA RICA: Puntarenas Pr., San Luis de Monteverde & vic., 1,000—-1,200 m, 10°17’N 84°49’W, 19.08.1998, L.M. LaPierre, #San97.93-01- 20 (CHAH, USNM, INBC), 1,100—1,400 sol m, 21-25.07, 02-09.08.1996, L.M.La Pierre, Cecropia (CHAH); Heredia Pr., Est. Biol. La Selva, 50-150 m, 10°26’N 84°01’'W, 09.11. 1998.MevBaPieme,, 798.2732; 24:01, 14:02; +02; 03;,.09:05.2000;. LM. LaPierre, #00.649, reared from Cecropia obtusifolia wood (8, BMNH, CHAH). NIC- ARAGUA, Granada, Sallé (holotype, BMNH). PANAMA: Chiriqui, 1300 m., Alto Lino nr. Boquete; UV traps 25:05:1977, 19.04.1978, H. Wolda (CWOB). Discussion.—Ptous otidocephalinus 1s very similar to P. cecropiae, but can be sep- arated by differences in the male genitalia and the characters given in the key. Speci- mens vary in size from 4.2—5.9 mm in length (mean = 5.05 mm, N = 36). The holotype is a female. ECOLOGY OF PTOUS Species of Ptous were reared by LaPierre as part of a general survey of wood-boring and other insects feeding on members of the Cecropiaceae at La Selva Biological Station and at San Luis de Monteverde, both in Costa Rica. Branches with maximum basal diameter of about 15 cm were clipped from mature trees of Cecropia obtusifolia Ber- toloni (La Selva and San Luis) and C. in- signis Liebmann, Coussapoa villosa Poep- pig and Endlicher, and Pourouma bicolor scobina (Benoist) C.C. Berg and van Heus- den (all at La Selva only) and left below the trees from which they were cut. After two months sections of the branches were removed and placed in rearing containers; sections were 30—40 cm in length and rep- resented a range of diameters from 4—15 cm. Ptous liebermanorum and P._ otidoce- phalinus were reared from branches of Cec- ropia obtusifolia and C. insignis at La Sel- va, and P. otidocephalinus from C. obtusi- folia at San Luis. Concurrent rearings from the other genera of the Cecropiaceae at La Selva (Coussapoa, Pourouma) did not pro- duce any Ptous although other species of Coleoptera were shared among the samples. Based on these rearing data and hand col- 862 lections of adults, it appears that Prous is a specialist on senescing wood of Cecropia species. Virtually all specimens of Prous we have seen for which there is collection data have been either reared from or collected on Cec- ropia or have been collected in UV light traps. It is interesting that no specimens of Ptous have been collected by the general sampling methods (Malaise traps, canopy foggings, UV light traps) of the Arthropods of La Selva (ALAS) Project. Although spe- cies of Cecropia have not been among the species fogged, other samples have been taken over a range of habitats at La Selva, and Cecropia species are widely distributed among these habitats. Cecropia specialists in other weevil genera (Lechriops, Lissod- eres, Pseudolechriops) have been collected in standardized samples, however. Other cryptorhynchine weevils reared from Cecropia by LaPierre include Coelos- ternus variisquamis (Champion) and Co- phes gibbus Champion. A complete sum- mary of the results of these rearings will appear elsewhere (LaPierre, in preparation). PTOUS AND MIMICRY Champion (1906) named Prous otidoce- phalinus for its striking resemblance to spe- cies of Otidocephalus (= Myrmex). The se- nior author has suggested elsewhere (Hes- penheide 1987, 1995) that this resemblance is a case of mimicry and that it involves a number of other taxa, including other spe- cies in several subfamilies of the Curcu- lionidae, Buprestidae, Cleridae, and Rhyn- chitidae. It is possible that species of Myr- mex are in fact the distasteful models for this system, in view of their species rich- ness and relative commonness, but no ex- perimental feeding trials have been made to support this hypothesis. It is interesting that both Prous and the zygopine weevil Lissod- eres share this habitus (Hespenheide 1987) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and that both use Cecropia species as hosts. Just as restriction of a species’ activity af- fects its ability to be sampled by general- ized sampling methods (see above), micro- habitat is also important to the effectiveness of mimicry (Hespenheide 1996). ACKNOWLEDGMENTS Chris Lyall and Sharon Shute assisted during a visit to The Natural History Mu- seum, London. Charles W. O’Brien shared expertise and generously loaned material from his personal collection. LML is grate- ful to Kirsten Querbach for the help she af- fably provided in monitoring rearing con- tainers at San Luis and to Pamela Wright for comments on the manuscript. Fieldwork by LaPierre at La Selva and San Luis was supported by a fellowship from the Depart- ment of Organismic Biology, Ecology and Evolution, UCLA. Hespenheide has been supported through the ALAS Project at La Selva (National Science Foundation grants BSR 9025024, DEB 9401069, DEB 9706976, and DEB-0072702) and grants from the UCLA Academic Senate. Both au- thors also were supported by personal funds. Margaret Kowalczyk prepared the figures. LITERATURE CITED Champion, G. C. 1904-1906. Curculionidae, Curcu- lioninae, Cryptorrhynchina. /n Biologia Centrali- Americana, Insecta, Coleoptera, IV, 4: 314—715. Hespenheide, H. A. 1987. A revision of the genus Lis- soderes Champion (Coleoptera: Curculionidae: Zygopinae). Coleopterists Bulletin 41: 41—55. . 1995. Mimicry in the Zygopinae (Coleoptera: Curculionidae). Memoirs of the Entomological Society of Washington 14: 145-154. . 1996. The role of plants in structuring com- munities of mimetic insects, pp. 109-126. In Gib- son, A., ed. Neotropical Biodiversity and Conser- vation, Los Angeles, CA: Mildred Mathias Botan- ical Garden. Wolda, H., C. W. O’Brien, and H. P. Stockwell. 1998. Weevil diversity and seasonality in tropical Pan- ama as deduced from light-trap catches. Smith- sonian Contributions in Zoology 590: 1-79. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 863-867 TWO NEW SPECIES OF THE CHEWING LOUSE GENUS GLIRICOLA MJOBERG (PHTHIRAPTERA: GYROPIDAE) FROM PERUVIAN RODENTS ROGER D. PRICE AND ROBERT M. TIMM (RDP) Department of Entomology, University of Minnesota, St. Paul, MN 55108, U.S.A. (Current address) 4202 Stanard Circle, Fort Smith, AR 72903-1906, U.S.A. (e- mail: rpricelice@aol.com); (RMT) Natural History Museum & Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS 66045-7561, U.S.A. (e- mail: btimm@ku.edu) Abstract.—Two new species of chewing lice (Phthiraptera: Gyropidae) from high-ele- vation Peruvian rodents are described and illustrated: Gliricola cutkompi from Cuscomys ashaninka Emmons (Abrocomidae) and G. brooksae from Dactylomys peruanus (J. A. Allen) (Echimyidae). The specimen of Cuscomys ashaninka that yielded the series of G. cutkompi also was the source of the type series of the previously described Abrocomo- phaga emmonsae Price and Timm. This finding reconfirms that single individual cavio- morph rodents may harbor two different genera of lice of the family Gyropidae. Resumen.—Se describen y se ilustran dos especies nuevas de piojos suramericanos (Phthiraptera: Gyropidae) de roedores peruanas de alta montana: Gliricola cutkompi par- asito de Cuscomys ashaninka Emmons (Abrocomidae) y G. brooksae parasito de Dac- tylomys peruanus (J. A. Allen) (Echimyidae). La serie tipica de G. cutkompi fue colectada del mismo especimen de Cuscomys ashaninka del cual se colect6 la serie tipica de Abro- comophaga emmonsae Price y Timm, anteriormente descrita por nosotros. Tal descubri- miento reconfirma que un solo individuo de roedor caviomorfo puede hospedar a dos generos distintos de la familia Gyropidae. Key Words: chewing lice, Gliricola, Gyropidae, Phthiraptera, Rodentia The chewing louse genus Gliricola Mj6berg (Phthiraptera: Gyropidae) contains 38 recognized species, with 29 of these in the nominate subgenus and 9 in the sub- genus Hutiaphilus Price and Timm. The members of the subgenus Gliricola are known from the Central and South Ameri- can caviomorph rodent families Caviidae and Echimyidae. Members of Hutiaphilus are restricted to the caviomorph family Capromyidae, the West Indian hutias that are found on the islands of the Greater and Lesser Antilles and the Bahamas. In our de- scription of the subgenus Hutiaphilus, we provide a discussion of the features of the genus Gliricola as well as both subgenera (Price and Timm 1997). Detailed character- ization of members of the family Gyropidae and subfamily Gliricolinae have been pre- sented by Clay (1970) and Price and Timm (2000). For brevity, these will not be re- peated here. We recently obtained two series of Glir- icola from two poorly known high-eleva- tion caviomorph rodents from Peru—the re- cently described abrocomid Cuscomys as- haninka Emmons (Abrocomidae) and the montane bamboo rat Dactylomys peruanus 864 (J. A. Allen) (Echimyidae). Each of these series represents a species new to science in the nominate subgenus. Furthermore, these hosts represent new generic records for Gliricola, and one, Cuscomys ashanin- ka, represents a new family record, the Abrocomidae. It is our intent to describe and illustrate these new species here. In the following descriptions, all mea- surements are in millimeters. The holotypes and some paratypes of both new species will be deposited in the Museo de Historia Natural, Universidad Nacional de San Mar- cos, Lima, Peru; other paratypes will be dis- tributed to the National Museum of Natural History, Smithsonian Institution (Washing- ton, D.C.), the K. C. Emerson Entomology Museum, Oklahoma State University (Still- water, Oklahoma), and the University of Minnesota (St. Paul, Minnesota). The hosts are deposited in the Museo de Historia Nat- ural, Universidad Nacional de San Marcos (MUSM) and the National Museum of Nat- ural History, Washington (USNM). Abbre- viations used for dimensions are given in the first description. For split drawings with a median vertical line, dorsal is to the left and ventral to the right. Gliricola (Gliricola) cutkompi Price and Timm, new species (Figs. 1-3) Type host.—Cuscomys ashaninka Em- mons. Female.—As in Fig. 1. Abdomen broad, with sparse chaetotaxy. Marginal tergal se- tae: I, 5—6; II-VIII, 10—17. Anterior tergal setae: I, 0; I-III, 2-7; IV—VIII, 4—14. Pleu- ral setae: H—VII, 10-14; VIII, 7-9; each side of pleura II-III with long seta, VI-— VIII with very long seta. Sternum II with 6—8 total setae. Marginal sternal setae: II, 8; IV—V, 9-13: VI-VII, 15—17. Anterior sternal setae mostly lateral: II, O—3; [V—V, 2-5; VI, 4-7; VII, 7-10. Tergum VII short- ened, terminal tergum large. Sterna [V—V short, VI medially enlarged, and VII en- larged, subrectangular. Each gonapophysis with 10—11 slender marginal setae. Dimen- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sions: temple width (TW), 0.27—0.29; head length (HL), 0.24—0.27; prothorax width (PW), 0.22—0.23; metathorax width (MW), 0.24—0.27; abdomen width at segment V (AWV), 0.49—0.56; total length (TL), 1.17— haSK0)s Male.—As in Fig. 3. Head and thorax near those of female. Abdomen with denser chaetotaxy and terga and sterna all of ap- proximately same length. Marginal tergal setae: I, 4—6; HI—VII, 15-22; VII, 13-16. Anterior tergal setae: I, 0; H, 4-8; HI-VII, 13—23. Pleural setae: H—III, 11—15; IV—VII, 11-17; VII, 10-14. Sternum I with 6—7 total setae. Marginal sternal setae: III, 8—10; IV—VII, 13-19. Anterior sternal setae even- ly across segment: III, 4—8; IV, 9-13; V, 11-15; VI, 14-19; VII, 17—22. Genitalia as in Fig. 2. Dimensions: TW, 0.25—0.27; HL, 0.22—0.23; PW, 0.20-—0.22; MW, 0.22-0.23; AWV, 0.37—0.39; TL, 1.01—1.06; genitalia width at paramere base, 0.06—0.07; genita- lia paramere length, 0.09—0.10; genitalia length, 0.38—0.41. Type material.—Holotype female, ex Cuscomys ashaninka, Pert: Cuzco, Cordil- lera Vilcabamba (11°39'36"S, 73°38'3 1" W), el355505m:, 15. June, 19977 colle Bourse Er Emmons #1359; in collection of the Museo de Historia Natural, Universidad Nacional de San Marcos, Lima, Peru, MUSM 12715 2 (also see Emmons 1999). Paratypes: 4 @, 5 6d, same data as holotype. Diagnosis.—The wide female head and abdomen, with the gross modifications in sizes of certain abdominal terga and sterna, set this species apart from all other known Gliricola. Additionally, the female gonapo- physis setae do not include any broad flat- tened setae, a condition that virtually all other species show. The male genitalia of G. cutkompi are unique, but show a general overall similarity to those of five other spe- cies—G. decurtatus (Neumann) from Kan- nabateomys amblyonyx (Wagner), G. fon- secait Werneck from Echimys dasythrix (Hensel), G. maculatus Werneck from Proechimys theringi Thomas, G. humilis Werneck from Proechimys albispinus (1. VOLUME 104, NUMBER 4 Figs. 1-5. G. brooksae. 4, Female ventral terminalia. 5, Female, dorsoventral. Geoffroy), and G. paraensis Werneck from Echimys grandis (Wagner). However, there are distinct differences in genitalic details and the females of all five species are quite different from those of G. cutkompi. We are uncertain about our interpretation of the unique segmentation of the female dorsal terminalia. There is the possibility that what we present in the female descrip- tion as a shortened tergum VII and much enlarged terminal tergum may actually be a situation in which tergum VII is transverse- 1-3, Gliricola cutkompi. 1, Female, dorsoventral. 2, Male genitalia. 3, Male, dorsoventral. 4—5, ly divided into two parts. If that is the case, then what we here consider terga VII and VIII may actually be a case in which both parts originate with tergum VII and tergum VIII may be fused with IX to form the en- larged terminal tergum. The location of the pleura and sterna seem to support the latter interpretation. Whichever may be correct, the details illustrated in Fig. 1 clearly im- part the essentials necessary for the correct application of the data in the description. Etymology.—This species is named for 866 Laurence K. Cutkomp, Department of En- tomology, University of Minnesota, in rec- ognition of his lengthy and productive ca- reer in teaching and research on insect tox- icology and the over 45 years he has been a close friend and colleague of the senior author. Gliricola (Gliricola) brooksae Price and Timm, new species (Figs. 4—5) Type host.—Dactylomys peruanus (J. A. Allen). Female.—As in Fig. 5. Abdomen narrow, without modifications of tergal and sternal sizes, and with many marginal tergal and sternal setae. Total tergal setae on I, 11—14. Marginal tergal setae: II-V, 25-34; VI-VII, 24-29; VIII, 20-25. Anterior tergal setae: II, 0-1; WI-VI, 4-8; VH-VIII, 5-11. Pleu- ral setae: II-VII, 7-14; VIII, 3—9; each of spiracle-bearing pleura III—VI with slender lateroposterior process; each side of pleuron II with 1 very long lateral seta and VIII with 2 such setae. Sternum II with 4—8 total setae. Marginal sternal setae: III, 8—10; IV, 13-18; V—VII, 17-21. Anterior sternal se- tae: III, 0; IV, 0-5; V—VI, 3-7; VII, 7-11. Each gonapophysis (Fig. 4) with 17-21 prominent marginal setae. Dimensions: TW, 0.19—0.20; HL, 0.20—0.21; PW, 0.15—0.16; MW, 0.19—-0.21; AWV, 0.37—0.43; TL, Lo L7=1.29: Male.—Unknown. Type material.—Holotype female, ex Dactylomys peruanus, Pera: Junin, Cordil- lera Vilcabamba (11°31'35’S, 73°38'31"W), el. 2,015 m, 23 June 1997, coll. Louise H. Emmons #1374; in collection of the Museo de Historia Natural, Universidad Nacional de San Marcos, Lima, Peri, MUSM @°. Paratypes: 2 2, same data as holotype; 5 @, same except (111°33'35"S; 73°38’ 28" Ww): el: 2,050 m, 2 July 1997, Louise H. Emmons #1398, USNM 582148 @. Diagnosis.—The female is unique among all known species of Gliricola in having the slender lateroposterior process on each of pleura III—VII and the large number of mar- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ginal setae surrounding each gonapophysis. No females of the other taxa have any sug- gestion of such a pleural process or of gon- apophysis chaetotaxy approaching that of G. brooksae. Etymology.—This species is named for Marion A. Brooks, Department of Ento- mology, University of Minnesota, in rec- ognition of her lengthy and productive ca- reer in teaching and research on insect mi- crobiology and the over 45 years she has been a close friend and colleague of the se- nior author. DISCUSSION The specimen of Cuscomys ashaninka that yielded the series of lice used here in the description of the new species Gliricola cutkompi also was the source of the series of Abrocomophaga used by Price and Timm (2000) as the basis for the new spe- cies description of A. emmonsae. Thus, this single individual was the host for lice of two different genera in the family Gyropi- dae. Where accurate records are available, we find gyropid lice to be extremely host-spe- cific ectoparasites, with speciation of lice closely paralleling speciation of their mam- malian hosts (Price and Timm 1997, 2000). Where we have a solid understanding of both host and louse species, only a single species of louse in the subgenus Gliricola occurs on a single host individual, whereas two (or in one case three) species of lice in the subgenus Hutiaphilus may be present on an individual host (Price and Timm 1997, 2000). As species of Gliricola are known to occur on a host species through- out its range, we suspect that G. cutkompi will be found on Cuscomys ashaninka and G. brooksae on Dactylomys peruanus throughout their respective ranges. Members of the genus Gliricola are now known from the Central and South Ameri- can caviomorph rodent families Caviidae (5 Gliricola known) and Echimyidae (25 Glir- icola known), the endemic South American family Abrocomidae (1 Gliricola known), VOLUME 104, NUMBER 4 and the endemic West Indian family Cap- romyidae (9 Hutiaphilus known). Because of the diverse radiation of the caviomorph rodents in these families (Abrocomidae, 5 species; Capromyidae, 13 species; Cavi- idae, 14 species; and Echimyidae, 70+ spe- cies), the host specificity of Gliricola, and the paucity of gyropid lice currently avail- able from these rodents, we strongly sus- pect that numerous new species of Gliricola have yet to be discovered. Clearly, species of Gliricola are widely distributed on the caviomorph rodents and are host specific, attesting to an ancient host—parasite asso- ciation. Much remains to be learned about the systematics and host relationships in this extremely diverse and speciose genus of chewing lice. ACKNOWLEDGMENTS We thank Linda Gordon and Louise Em- mons, National Museum of Natural Histo- ry, Washington, D.C., and Victor Pacheco, Museo de Historia Natural, Universidad 867 Nacional de San Marcos, Lima, for allow- ing us to remove specimens of lice from mammal specimens housed in their respec- tive collections, thereby making the discov- ery of these two new species possible. Rob Anderson translated our Abstract into Span- ish for the Resumen included herein. This work was partially sponsored by the Agri- cultural Experiment Station, University of Minnesota. LITERATURE CITED Clay, T. 1970. The Amblycera (Phthiraptera: Insecta). Bulletin of the British Museum (Natural History), Entomology 25: 73-98. Emmons, L. H. 1999. A new genus and species of abrocomid rodent from Peru (Rodentia: Abrocom- idae). American Museum Novitates 3279: 1-14. Price, R. D. and R. M. Timm. 1997. A new subgenus and four new species of Gliricola (Phthiraptera: Gyropidae) from Caribbean hutias (Rodentia: Capromyidae). Proceedings of the Biological So- ciety of Washington 110: 285-300. . 2000. Review of the chewing louse genus Abrocomophaga (Phthiraptera: Amblycera), with description of two new species. Proceedings of the Biological Society of Washington 113:210—217. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 868-883 ENIGMATIC TREEHOPPER GENERA (HEMIPTERA: MEMBRACIDAE): DEIRODERES RAMOS, HOLDGATIELLA EVANS, AND TOGOTOLANIA, NEW GENUS JASON R. CRYAN AND LEwis L. DEITZ (JRC) Laboratory for Conservation and Evolutionary Genetics, New York State Mu- seum, Albany, NY, 12230 U.S.A. (e-mail: jcryan@mail.nysed.gov); (LLD) Department of Entomology, Box 7613, North Carolina State University, Raleigh, NC 27695-7613, U.S.A. (e-mail: lewis-deitz @ncsu.edu) Abstract—Two poorly known Neotropical treehopper genera, Deiroderes Ramos and Holdgatiella Evans, are revised and redescribed based on adult morphology. The Carib- bean genus Deiroderes (unplaced within the subfamily Stegaspidinae) has three valid species including D. inornatus, new species (Jamaica). The Neotropical genus Holdga- tiella (currently unplaced within Membracidae) has two valid species including H. stria, new species (Venezuela). In addition, a previously unknown Caribbean genus is here described, Togotolania, new genus, with two species: T: longicorna, new species (Do- minican Republic) and 7. brachycorna, new species (Guadeloupe). All species included in these three genera are illustrated, and keys are given for the identification of adults. Key Words: Membracidae, Deiroderes, Holdgatiella, Togotolania, Caribbean, taxonomy Treehoppers (Hemiptera: Membracidae) are traditionally recognized by the enlarged pronotum, which in many species is con- spicuously ornamented with stalks, spikes, or bulbs. In most genera, there is at least a posterior pronotal process extending over (or even concealing) the scutellum. How- ever, the pronota of the Neotropical tree- hopper genera Deiroderes Ramos and Holdgatiella Evans are remarkably unre- markable. A third genus, Togotolania, de- scribed here as new, also has a simple pron- otum, adorned only with a median horn. These three genera are probably not closely related, indicating that enlarged pronotal or- namentation was likely gained or lost mul- tiple times in Membracidae. Other treehop- per genera lacking a posterior pronotal pro- cess occur in the subfamilies Nicomiinae, Endoiastinae, Centrotinae, and Stegaspidi- nae (fossils), though these may be cases of secondary loss. Following the most recent reclassifica- tions of Membracidae (Deitz and Dietrich 1993a, Dietrich et al. 2001a), Deiroderes and Holdgatiella were among several gen- era that remained unplaced within the tax- onomic framework. Although these two genera were originally described in the sub- family Centrotinae (Ramos 1957a, Evans 1962a), emerging morphological (Dietrich et al. 2001a, Cryan et al. in press) and mo- lecular (Cryan et al. 2000a, Cryan, unpub- lished data) evidence is revealing support for new phylogenetic placements within Membracidae, as discussed below. MATERIALS AND METHODS Protocols used during this work follow the materials and methods described by Cryan and Deitz (1999a). The following codens are used to refer to the collections in which relevant specimens are located or VOLUME 104, NUMBER 4 have been deposited. Arnett et al. (1993a) listed the full postal addresses for most of the institutions; those not found in that pub- lication are indicated by a dagger (7+) fol- lowing the coden. CIRAD?: Labo Entotrop (Faunistique-Tax- onomie), Montpellier, France. CNCI: Canadian National Collection of In- sects, Agriculture and Agri-Food Cana- da, Ottawa, Canada. JARC?: J.A. Ramos Collection, University of Puerto Rico, Mayagtiez, Puerto Rico. JWEC?7: J.W. Evans Collection, c/o Austra- lian Museum, Sydney, Australia. MNHN: National Collection of Insects, Muséum National D’ Histoire Naturelle, Paris, France. NCSU: North Carolina State University In- sect Collection, Department of Entomol- ogy, North Carolina State University, Ra- leigh, North Carolina, USA. SHMC?+y: S.H. McKamey Collection; cur- rent address: Systematic Entomology Laboratory, USDA, % National Museum of Natural History, Smithsonian Institu- tion, Washington, D.C., 20560-0168, USA. USNM: National Museum of Natural His- tory, Smithsonian Institution, Washing- ton DiC.. USA: Genus Deiroderes Ramos 1957 Deiroderes Ramos 1957a: 96. Type species: Deiroderes inermis Ramos 1957a: 96, by original designation. Diagnosis.—The genus Deiroderes in- cludes small stegaspidine treehoppers with simple pronota (lacking well-developed su- prahumeral horns and posterior pronotal process) and dark tubercles along the veins of the forewings. Adult.—Dimensions (mm): total length (from head to apex of forewings at rest) 3.2—3.9. Structure: Head (Figs. 1-2, 8-9, 14—15): broad, wider than long, with fine pubescence; ocelli not raised, above centro- ocular line; foliate lobes small, rounded or quadrate; postclypeus weakly or strongly 869 trilobed. Thorax: Pronotum (Figs. 2, 9, 15): simple, convex; either low or elevated; su- prahumeral horns represented by low cari- nae, or absent; posterior pronotal process absent. Pronotal surface sculpturing: finely punctate, each pit associated with one short setae clubbed, normal, or flattened. Scutellum (Figs. 2, 9, 15): short, acuminate; anterior half slightly raised; posterior half seta; flattened, with slight median depression. Legs (Figs. 3, 10): metathoracic femur with or without dorsal row of cucullate setae; metathoracic tibia with enlarged setal rows I, Hf, and III, all with cucullate setae; meta- thoracic tarsomere I with | apical cucullate seta. Forewing (Figs. 4, 11, 16): punctate, coriaceous basally; venation simple; | [rarely, 2] rm and | m-cu crossvein pre- sent; dark tubercles, each bearing one stout seta, spaced irregularly along veins and crossveins. Hindwing: venation essentially identical to Microcentrus hindwing vena- tion as illustrated by Deitz (1975a, fig. 37c) and Dietrich et al. (2001a, fig. 2B). Geni- talia: 2 2™4 valvulae (Figs. 5, 17) narrow throughout, with distinct dorsal serrations; d lateral plates (Figs. 6, 12) free, without hook; aedeagus and styles (Figs. 7, 13) var- iable; aedeagus with anterior face of pos- terior arm not denticulate. Range.—Caribbean, from Virgin Islands to Cuba. Remarks.—Ramos (1957a) originally de- scribed Deiroderes as a member of the sub- family Centrotinae, indicating the genus was closely related to Lophyraspis Stal (now in Aetalionidae: Biturritiinae: Bitur- ritiini) and Tolania Stal (now in Membra- cidae: Nicomiinae: Tolaniini). Deiroderes was later removed from Centrotinae and considered unplaced within Membracidae (Deitz and Dietrich 1993a). Dietrich et al. (2001a) found that Deiroderes was included within a monophyletic Stegaspidinae in some of the most-parsimonious cladograms resulting from a more detailed morphology- based phylogenetic analysis of the family Membracidae. Nevertheless, evidence for placement of Deiroderes remained equivo- 870 cal, and therefore the genus remained un- placed in Membracidae. In a phylogenetic analysis of basal tree- hopper groups, Cryan et al. (in press) found that Deiroderes groups with the genus An- tillotolania Ramos at the base of Stegaspi- dinae. Morphological features that placed the genus within Stegaspidinae included the forewing’s distal m-cu crossvein (fused ba- sad of the fork of vein M) and hooklike apex of the male styles. Within Stegaspidinae, some features— forewing with one r-m and one m-cu cross- vein; vein R,,, fused basally with R,; and male lateral plates free, lacking posteroap- ical hooks—suggest that Deiroderes is more closely related to the tribe Stegaspi- dini (reviewed in Cryan and Deitz 1999a, b, 2000a). Nevertheless, phylogenetic anal- yses based on morphological (Dietrich et al. 2001a, Cryan et al. in press) and molecular (Cryan et al. 2000a, Cryan, unpublished data) evidence suggest either that Deirod- eres might be more closely related to Mi- crocentrini or that placement is equivocal. Therefore, pending more definitive resolu- tion, we refer Deiroderes to Stegaspidinae but propose that the genus remains un- placed to tribe. When creating a generic name that refers to a feature of the prothorax, it is customary to modify the Greek noun “‘dere”’ to “‘de- res, thereby making the name masculine (G. Kuschel, personal communication). The first part of the compound name, “‘deiro-,” means to behead something, or to cut the neck; thus, ““Deiroderes” best translates as the ““beheaded”’ treehopper, probably refer- ring to either the lack of pronotal horns or the relatively flat head. KEY TO THE SPECIES OF ADULT DEIRODERES 1. Forewing vein M branching at, or immediately distad of, crossvein rm, M,,, extending pos- teriorly at right angle to M (Fig. 4); metatho- racic femur with dorsal and ventral rows of cu- culllatetsetaen(Eic= 3) is sere D. inermis Ramos — Forewing vein M branching distinctly distad of crossvein r-m, M,,, extending obliquely to PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON wing margin (Figs. 11, 16); metathoracic femur without rows of cucullate setae (Fig. 10) ... 2 2. Pronotal metopidium produced dorsally, with well-developed median carina; pronotum with suprahumenall\canmae (Bis iS) a) 22 ae ee D. punctatus (Metcalf and Bruner) — Pronotal metopidium low, dorsal median carina not well developed; pronotum without supra- humeral carinae or horns (Fig. 9) ....... Sedive- ube Teanga ine ores D. inornatus, new species Deiroderes inermis Ramos (Figs. 1—7) Deiroderes inermis Ramos 1957a: 96. Type locality—Guanica, Puerto Rico. Diagnosis.—Deiroderes inermis has the forewing with vein M branching immedi- ately distad of the r-m crossvein, M,,, ex- tending directly to the posterior margin of wing (nearly forming a right angle with M). Adult.—Dimensions (mm): total length 2 3.2-3.5, 5 3.5; width between humeral angles 2 1.2-1.4; ¢ 1.3; pronotal length ¢ 0.8-1.0, do 0.9; wing length 2 2.6—2.8, 6 2.4; width of head including eyes 2 1.1— 1.3, 6 1.2. Coloration: body tan to dark brown; forewing dark basally, dark colora- tion sometimes extending % of wing length, with dark spot at distal end of vein Cu. Structure: Head: dorsal projections (Fig. 1) indistinct. Thorax: Pronotum (Fig. 2): metopidium low; humeral angles short, blunt; middorsal crest weakly produced; su- prahumeral horns and carinae absent. Legs (Fig. 3): metathoracic femur with dorsal and ventral rows of cucullate setae; meta- thoracic tibiae with cucullate setae row I less robust than setae in II or Il. Forewing (Fig. 4): M branching immediately distal to rm crossvein, M,,, extending to posterior edge of wing, at right angle to M. Genita- lias 2 23) valvulae) (is. oS) marco throughout, apical % with distinct dorsal serrations. ¢: lateral plates (Fig. 6) free, lacking hooks, sparsely setose; styles (Fig. 7) with small hook apically; aedeagus (Fig. 7) with posterior arm straight. Nymph.—Unknown. Distribution.—British Virgin Islands: Guana Island [NCSU], Virgin Gorda VOLUME 104, NUMBER 4 871 3mm y= = A. po = y We es PY, vy, oe er eee fe g eT ele Eveetad Figs. 1-7. Deiroderes inermis. Structures of the holotype (male genitalia illustrated from a non-type speci- men). 1, Head, anterior aspect (face). 2, Head, pronotum, and scutellum, anterolateral aspect. 3, Left metathoracic femur, tibia, and tarsus, ablateral aspect. 4, Right forewing. 5, Female second valvulae, lateral aspect. 6, Male left lateral plate, lateral aspect. 7, Male aedeagus and left style, lateral aspect (illustrated with connective tissue). Abbreviations: A,, anal vein(s); ae, aedeagus; Cu,, cubital vein(s); cs, claval suture; ha, humeral angles; m-cu,, medial-cubital crossvein(s); M,, medial vein(s); mc, median carina; me, metopidium; sc, scutellum; st, style; r, radial crossvein; r-my,, radial-medial crossvein(s); R,, radial vein(s); I, Hl, III, enlarged setal rows. 872 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON [NCSU], Tortola [NCSU]; U.S. Virgin Is- lands: St. John [JARC]; Puerto Rico [NCSU]. Material examined.—Holotype [2] [NCSU, deposited on indefinite loan to USNM] with labels: ‘“‘Insular Forest/Guan- ica PR./3-Aug. 1945./On Capparis nitida.,”’ “j.A. Ramos/Collector,~ “Holotype; *‘Deiroderes/inermis Ramos/145.”’ Paratype [2; forewings missing] [NCSU] with la- bels: ““Cambalache/PR. 11-7-47,” ‘‘Cald- well &/Martorell/Collectors,” “‘Paratype,” ‘“Deitz Research/83-227e @, ‘“JARC,” and ‘“‘PARATYPE/Deiroderes/inermis/Ra- mos.’ Other specimens: | 2 (Deitz Re- Search #652271 2)! trom JARC. Qc (Cryan Research #94-286a 3 and Cryan Research #94-273b 3) from NCSU; 1 @ from USNM. Remarks.—The holotype of D. inermis was collected on Capparis sp. (Cappara- ceae), though there is a discrepancy con- cerning the host species. In the original de- scription of D. inermis (Ramos 1957a), the holotype is listed as “‘on Capparis indica (L.) Fawe and Rendle’’; however, the ho- lotype label information indicates the host as C. nitida (C. indica and C. nitida are not synonyms). This insect’s Latin specific epithet translates directly to ““unarmed” or ““weaponless,”’ presumably referring to the complete absence of pronotal projections (suprahumeral horns and posterior pro- cess). A specimen of D. inermis from the Brit- ish Virgin Islands was included in a mo- lecular systematic investigation of mem- bracid relationships (Cryan et al. 2000a). Results based on parsimony and maximum likelihood analyses of DNA nucleotide se- quence data from two nuclear genes (EF- la and 28S rDNA) indicated that Deirod- eres was a member of a clade that also included Antillotolania Ramos and Micro- centrus Stal. Deiroderes inornatus Cryan and Deitz, new species (Figs. 8-13) Type locality —Jamaica. Diagnosis.—Deiroderes inornatus has a simple, unadorned pronotum, strongly de- flected postclypeus and beak, and sinuate aedeagus. Adult ¢.—Dimensions (mm): total length 5.1—6.0; width between humeral an- gles 1.9—2.1; pronotal length 1.4—1.5; wing length 4.0—4.9; width of head including eyes 2.1—2.3. Coloration: body tan to dark brown; forewing tan or dark brown basally, light brown apically. Structure: Head: dor- sal projections (Fig. 8) small or indistinct; postclypeus and anteclypeus strongly de- flected under thorax. Thorax: Pronotum (Fig. 9): metopidium low; humeral angles weakly produced; mid-dorsal crest weakly produced; suprahumeral carinae and horns absent. Legs (Fig. 10): metathoracic femur without rows of cucullate setae. Forewing (Fig. 11): vein M branching well after rm crossvein; M,,, extending obliquely to pos- terior margin of wing; some specimens with 2 r-m crossveins on one or both forewings. Genitalia: lateral plates (Fig. 12) free, without hooks; styles (Fig. 13) sinuate, hooked apically; aedeagus (Fig. 13) with posterior arm = sinuate, narrowing apical- ly.2: unknown. Nymph.—Unknown. Distribution.—Jamaica [USNM, CNCI]. Material examined.—Holotype [4] [USNM] with labels: ‘‘Jamaica/19-VII-35,”’ “Sta 16/Chapin and/Blackwelder,” ‘‘Vouch- er Specimen/Cuticular Hydrocarbon/Anal- ysis Study/Jason R. Cryan, 1995,” and “HOLOTYPE 6/Deiroderes/inornatus/ Cryan & Deitz.”’ Paratype [¢] [CNCI] with labels: “JAMAICA, 4000'/Hardwar Gap/ VII-23-1966/Howden & Becker,” “Speci- men data/captured—NMNH/Biota (Col- well) file,” and ““PARATYPE 4d/Deirod- eres/inornatus/Cryan & Deitz.”’ Paratype [d, dissected; head and pronotum miss- ing] [USNM] with labels: “Jamaica/19- VIH-35,” “Sta 16/Chapin and/Blackweld- er,” “CRYAN RESEARCH/#94-273a 6,” and ““PARATYPE 46/Deiroderes/inornatus/ Cryan & Deitz.” Other specimens: 1 d from CNCI; 1 ¢ from USNM. Remarks.—Deiroderes inornatus 1s de- VOLUME 104, NUMBER 4 873 Figs. 8-13. otum, and scutellum, anterolateral aspect. 10, Left metathoracic femur, tibia, and tarsus, ablateral aspect. 11, Right forewing. 12, Male left lateral plate, lateral aspect. 13, Male aedeagus and left style, lateral aspect (illus- trated with connective tissue). scribed from five males; females are un- known. Minor color variation is evident among males, and while some specimens have 2 r-m crossveins in one or both fore- wings, no other substantial variation in morphology of males is apparent. Further, although some species in the subfamily Ste- gaspidinae exhibit sexual di- or polymor- phism (Cryan et al. in press), both sexes of D. inermis are morphologically similar. This fact, combined with the apparent al- lopatric distribution of Deiroderes species, makes it unlikely that D. inornatus is ac- Deiroderes inornatus. Structures of the holotype. 8, Head, anterior aspect (face). 9, Head, pron- tually the male of D. punctatus (for which males are unknown). The specific name “inornatus” is Latin for “unadorned,” referring to the lack of su- prahumeral horns and posterior pronotal process in this species. Deiroderes punctatus (Metcalf and Bruner) (Figs. 14-17) Tolania punctata Metcalf and Bruner 12s baZ 1S. Eustollia punctata: Goding 1926a: 105. 874 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 3mm Figs. 14-17. Deiroderes punctatus. Structures of a female paratype. 14, Head, anterior aspect (face). 15, Head, pronotum, and scutellum, anterolateral aspect. 16, Right forewing. 17, Female second valvulae, lateral aspect. Deiroderes punctatus: Ramos 1957a: 96. Microcentrus punctata: Hamilton 1971b: DOs: Type locality—Camagtiey, Cuba. Diagnosis.—Deiroderes punctatus has an elevated pronotum with an expanded mid- dorsal carina and suprahumeral ridges. Adult 2°.—Dimensions (mm): _ total length 3.6—3.8; width between humeral an- gles 1.3—1.4; pronotal length 0.9—1.1; wing length 3.0—3.1; width of head including eyes 1.3—1.4. Coloration: body light brown; pronotum with dark brown patches; forewing brown in basal 4%, with dark spot at distal end of vein Cu. Structure: Head: dorsal projections (Fig. 14) distinct, partial- ly hidden by anterior pronotal shelf; post- clypeus prominent, trilobed. Thorax: Pron- otum (Fig. 15): metopidium elevated; su- prahumeral carinae present; middorsal ca- rina strongly = elevated; ~— supraocular callosities evident. Legs: metathoracic fe- mur without rows of cucullate setae. Fore- wing (Fig. 16): vein M branching well be- yond r-m crossvein; M,,, extending obliquely to posterior margin of wing. Gen- italia: @: 2" valvulae (Fig. 17) narrow throughout, distal % with distinct dorsal ser- rations. 6: unknown. Nymph.—Unknown. Distribution.—Cuba [NCSU]. Material examined.—Paratype [@, dis- sected] [NCSU] with labels: ““Camaguey, @uba/Col> J. Acuna/Julie 15, 19232 Ons “Guaitajan,” “190b,”” “CUBA/5,” “Para- type,” ““Deitz Research/73-107c 2,” ““To- lania/punctata/Metc. & Brun./det. Z.P.M.,” and **Deiroderes/punctata/(Metcalf &/Bru- ner)/det. L.L. Deitz 1983.’’ Paratype [2] [NCSU] with labels: ““906I,” ““Camaguey, Cuba/Col. J-Acuna/Julies IS 192 3% ‘*Paratype,”’ and ‘*Tolania/punctata/Metc. & Brun./det. Z.P.M.”’ VOLUME 104, NUMBER 4 Remarks.—Although included in four dif- ferent genera by four different authors, the overall morphology of D. punctatus (espe- cially of the forewing) places it in the genus Deiroderes. The specific epithet 1s an adjec- tive and must agree in gender with Deirod- eres (masculine); thus the correct spelling is D. punctatus. Other than the cryptic refer- ence of one specimen being found *‘‘on Guaitajan” (see Material Examined, above; we found no reference to this in any botan- ical literature consulted), no information is available concerning the life history, ecolo- gy, or behavior of this species. Genus Holdgatiella Evans 1962 Holdgatiella Evans 1962a: 515. Type spe- cies: Holdgatiella chepuensis Evans 1962a: 516, by original designation. Diagnosis.—The forewing of Holdgatiel- la has vein M 3-branched; the male aedea- gus is tentaculate. Adult.—Dimensions (mm): total length (from head to apex of forewings at rest) 6.1=—7.9.. Structure: Head (Figs. 18, 25): dorsal projections weakly developed. Tho- rax: Pronotum (Figs. 19, 26): suprahumer- al horns and posterior process absent; me- topidium steeply declivous. Pronotal sur- face sculpturing: finely punctate, each pit associated with one short seta. Scutellum (Figs. 19, 26): swollen basally, flattening to acuminate apex. Legs (Figs. 20, 27): meta- thoracic tibiae with cucullate setae in rows I, II, and III; metathoracic tarsomere I with | apical cucullate seta. Forewing (Figs. 21, 28): apical limbus broadened proximally along costal margin, vein M with 3 branch- es, | rm crossvein present (distad of first branch of vein M). Hindwing: venation es- sentially identical to Microcentrus hind- wing venation as illustrated by Deitz (1975a, fig. 37c) and Dietrich et al. (2001, fig. 2B); apex with arcuate chaetoids (Die- trichs and Deitz. 1993.45, Dietrich et alt 2001a). Genitalia: 2 24 valvulae (Figs. 22, 29) slightly broadened at or beyond mid- point, with distinct dorsal serrations; d lat- 875 eral plates (Figs. 23) fused to pygofer; apex of aedeagus tentaculate (Fig. 24). Range.—Chile, Venezuela. Remarks.—As Evans (196la) observed, Holdgatiella superficially resembles Dei- roderes, differing chiefly in wing venation and morphology of the male and female genitalia. Evans placed Holdgatiella in the subfamily Centrotinae, while noting that the genus 1s closely related to the complex comprising Aetalionidae, Nicomiidae (now Membracidae: Nicomiinae), Biturritidae (now Aetalionidae: Biturritiinae), and the leafhopper subfamilies Ulopinae and Ma- cropsinae. Linnavuori and DeLong (1978a) wrote, “Holdgatiella, an undoubted Meso- zoic relic, evidently belongs to Membraci- dae as suggested by Evans, although it ap- parently has rather an isolated position within this family. Its relations to the other Membracidae remain open until a modern revision of the neotropical representatives of the family has been done.” Indeed, this genus was among seven membracid genera unplaced to subfamily in Deitz and Die- trich’s (1993a) reclassification of Membra- coidea. The morphology-based phylogenet- ic analyses of Dietrich et al. (2001la) sug- gest that Holdgatiella is the sister group to Nicomiinae, and the authors noted that an expanded concept of Nicomiinae (to in- clude Holdgatiella and other groups) might be warranted. In those analyses, Holdga- tiella is closely allied with Nicomia and To- lania, based primarily on forewing vena- tion: vein M with three branches and vein R with four branches (only in H. stria, new species, but apparently the plesiomorphic condition). The tentaculate male aedeagus of Holdgatiella, however, sets it distinctly apart from these genera. Results from a sep- arate morphology-based phylogenetic in- vestigation of basal treehopper groups (Cryan et al. in press) agreed with the al- liance of Holdgatiella with Nicomiinae. A more detailed analysis of relationships within Nicomiinae and related groups is un- derway (Albertson and Dietrich, personal communication). 876 KEY TO THE SPECIES OF ADULT HOLDGATIELLA 1. Scutellum (Fig. 19) without mid-dorsal groove; metathoracic femur (Fig. 20) with ventral row of cucullate setae; forewing (Fig. 21) with 2 or 3 m-cu crossveins; frons with foliate lobes weakly developed (Fig. 18) — Scutellum (Fig. 26) with mid-dorsal groove; metathoracic femur (Fig. 27) without ventral cucullate setae; forewing (Fig. 28) with only | m-cu crossvein; frons with foliate lobes well developed (Fig. 25) H. chepuensis Evans H. stria, new species Holdgatiella chepuensis Evans (Figs. 18—24) Holdgatiella chepuensis Evans 1962a: 516. Type locality—Chepu, Chile. Diagnosis.—Holdgatiella chepuensis has a ventral row of cucullate setae on the meta- thoracic femur. Adult.—Dimensions (mm): total length 2 6.5-7.0, 3 6.1—6.4; width between hu- tal length @ 1.2-1.4, d 1.3-1.4; wing length 2 5.5—5.9, d 5.2—5.4; width of head including eyes 2 2.4—2.6, 6 2.3—2.4. Col- oration: 2: face and body tan with dark brown markings, legs tan, forewing tan ba- sally, hyaline apically; d: face and body fuscous with black markings, abdominal sternites yellow, abdominal tergites yellow with brown transverse stripes, legs fuscous dorsally, tan ventrally, forewing brown ba- sally, anterior half of membranous area hy- aline, posterior half brown. Structure: Head: face (Fig. 18) with weakly devel- oped foliate lobes; postclypeus ovoid. Tho- rax: Pronotum (Fig. 19): humeral angles produced, blunt; median dorsal carina weak; posterior margin without triangular projections. Pronotal surface sculpturing: punctate; pits shallow, associated with a single short, apically curved seta. Scutellum (Fig. 19): slightly swollen basally, flatten- ing to acuminate apex. Legs (Fig. 20): metathoracic femur with dorsal and ventral rows of cucullate setae (20—26 and 15-18 setae, respectively). Forewing (Fig. 21): basal %—% coriaceous; branching pattern of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON vein R variable; 2 or 3 m-cu crossveins pre- sent, distal m-cu crossvein distad of first branch of vein M. Genitalia: 9: 2" val- vulae (Fig. 22) abruptly broadened in distal ¥%: dorsal ridge of broadened area weakly serrate. ¢: lateral plate (Fig. 23) fused to pygofer; styles (Fig. 24) elongate, ventrally curved to axe-like apical lobe; aedeagus (Fig. 24) L-shaped, anterior arm irregularly shaped, posterior arm slender; anterior face of posterior arm apically tentaculate (5 pairs of short, tentacle-like appendages). Nymph.—Unknown. Distribution.—Chile: Aucar [JWEC]; Cautin [JARC]; Chepu [NCSU]; Llanqui- hue [JARC]; Osorno [JARC]. Material examined.—Holotype not ex- amined. Other specimens: 2 2 and 4 ¢d from JARC, | ¢ from JWEC, and | 6 from NCSU. Remarks.—The aedeagal structure of this species differs greatly from that of other treehoppers. Not only are the apical lobes of the styles uniquely shaped, but also the presence of five pairs of apical appendages on the aedeagal corona sets H. chepuensis apart. Linnavuori and DeLong (1978a) il- lustrated various anatomical structures of H. chepuensis, including the hindwing, which appears nearly identical in venation to Mi- crocentrus hindwings (Deitz 1975a, fig. 37c; Dietrich et al. 2001a, fig. 2B). The spe- cific name “‘chepuensis” refers to the type locality, Chepu, Chile. Holdgatiella stria Cryan and Deitz, new species (Figs. 25-29) Type locality—20 km SE of Azulita, Mérida, Venezuela. Diagnosis.—Holdgatiella stria has the pronotum with posterolateral processes, the scutellum with a median dorsal groove, and the forewing with | m-cu crossvein. Adult: (2:==Dimensions Guim): sstotal length 7.9; width between humeral angles 2.6; pronotal length 2.1; wing length 7.0; width of head including eyes 2.5. Colora- tion: face tan with fuscous punctures and VOLUME 104, NUMBER 4 877 Figs. 18-24. Holdgatiella chepuensis. 18, Head, anterior aspect (face). 19, Head, pronotum, and scutellum, anterolateral aspect. 20, Left metathoracic femur, tibia, and tarsus, ablateral aspect. 21, Right forewing. 22, Female second valvulae, lateral aspect. 23, Male left lateral plate and pygofer, lateral aspect. 24, Male aedeagus and left style, lateral aspect (illustrated with connective tissue). foliate lobes, body tan with dark markings, clypeus weakly trilobed. Thorax: Prono- pronotum tan with brown metopidial tum (Fig. 26): humeral angles short, stout; crown, legs tan, forewing tan basally, hya- median dorsal carina well developed; pos- line apically. Structure: Head: face (Fig. terior margin with triangular projections on 25) with well-developed foliate lobes; post- either side of scutellum. Scutellum (Fig. 878 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 25—29. otum, and scutellum, anterolateral aspect. 27, Left metathoracic femur, tibia, and tarsus, ablateral aspect. 28, Right forewing. 29, Female second valvulae, lateral aspect. 26): slightly produced basally, flattening to acuminate apex, with median dorsal groove. Legs (Fig. 27): metathoracic femur with dorsal row of 15—18 cucullate setae; meta- thoracic tibia with paired cucullate setae in distal % of row II. Forewing (Fig. 28): bas- al %4 coriaceous; vein R with 4 branches (R,, R;, R3, R45); 1 m-cu crossvein present, distad of first branch of vein M. Genitalia: 2™ valvulae (Fig. 29) slightly broadened midway; dorsal ridge of distal % serrate. ¢: unknown. Nymph.—Unknown. Distribution.— Venezuela: Mérida [SHMC/ USNM]. Material examined.—Holotype [°@] [SHMC, deposited on indefinite loan to USNM], with labels ““VEN: Merida, 20 Holdgatiella stria. Structures of the holotype. 25, Head, anterior aspect (face). 26, Head, pron- km/SE Azulita, La/Carbonera 27.VII-/ 3.VII.89 S&J Peck,” “?,’’ “Holdgatiella sp./det. S. McKamey 1991,” and *““HOLO- TY PE/Holdgatiella/stria/Cryan & Deitz.” Remarks.—Although the scutellum is not emarginate apically, the presence of a me- dian dorsal groove suggests that the ances- tors of H. stria had a posterior pronotal pro- cess that was secondarily lost. The specific name “stria’’ is Latin for “furrow” of ‘““sroove,” referring to the median dorsal groove of the scutellum. Genus Togotolania Cryan and Deitz, new genus Type species.—Togotolania longicorna, new species. Diagnosis.—Togotolania has the follow- VOLUME 104, NUMBER 4 ing unique combination of characters: head with large dorsal projections, pronotum with a median horn, and forewings each with 3 r-m and | m-cu crossveins. Adult.—Dimensions (mm): total length (from head to apex of forewings at rest) 6.4—7.2. Structure: Head: dorsal cranial projections large, either pyramiform (Fig. 30) or emarginate (Fig. 35); ocelli on or above centro-ocular line. Thorax: Prono- tum (Figs. 31, 36): metopidium elevated; supraocular callosities indistinct; posterior process lacking, suprahumeral horns absent or represented by low carinae at base of median dorsal horn. Scutellum (Figs. 31, 36): short with swollen base, flattening to acuminate apex. Legs (Fig. 32): metatho- racic tibia with enlarged setal rows I, HI, and III, all with reduced cucullate setae. Fore- wing (Figs. 33, 37): 3 r-m crossveins pre- sent; vein R,,; fused to R basad of fork of R, and R,,,; 1 m-cu crossvein present, dis- tad of fork of vein M. Hindwing: venation essentially identical to Microcentrus hind- Wing venation as illustrated by Deitz (1975a, fig. 37c) and Dietrich et al. (2001, fig. 2B). Genitalia: 2 2™ valvulae (Fig. 34) broadened at midpoint; d lateral plates (Fig. 38) free, lacking apical hook; styles (Fig. 39) slender, with apical hook; aedea- gus (Fig. 39) U-shaped, with posterior arm tapering sharply at apex. Range.—Dominican Republic, Guade- loupe. Remarks.—Included in the phylogenetic analysis of Dietrich et al. (2001a) as ““New Genus T”, Togotolania was placed in a largely unresolved lineage basal to the clade comprising Nicomiinae + Centrodon- tinae + Centronodinae. Results of a sepa- rate morphology-based analysis (Cryan et al. in press) were concordant, with Togo- tolania placed at the base of a lineage com- prising Holdgatiella, Tolania, and Centron- odini. Although we here leave Togotolania unplaced in the family Membracidae, we acknowledge that this genus is probably al- lied with Nicomiinae. The generic name “Togotolania”’ translates as “‘cloaked To- 879 lania”’ and refers to the hoodlike pronotal structure. It is formed from “‘toga”’ [Latin] and “‘tolania” [origin unknown], joined by the vowel “‘o”’ for euphony (Brown 1956). KEY TO THE SPECIES OF ADULT TOGOTOLANIA 1. Median pronotal horn (Fig. 31) large, laterally compressed; head (Figs. 30-31) with pyrami- form dorsal projections; forewing (Fig. 33) with crossvein r-m, distad of fork of vein M aH ance dander ices chen T. longicorna, new species — Median pronotal horn (Fig. 36) small, not lat- erally compressed; head (Figs. 35-36) with emarginate dorsal projections; forewing (Fig. 37) with crossvein r-m, basad of fork of vein T. brachycorna, new species Togotolania longicorna Cryan and Deitz, new species (Figs. 30-34) Type locality.—2 km east of Boca Chica, Dominican Republic. Diagnosis.—Togotolania longicorna is easily recognized by its large, median, an- terior pronotal horn; the pronotum is uni- formly black. Adult 2.—Dimensions (mm): total length 6.4; width between humeral angles 1.9; pronotal length 1.6; wing length 5.1; width of head including eyes 1.9. Colora- tion: face tan with dark brown ovate mark- ings; body fuscous with pronotum black; forewing brown with dark venation; legs brown. Structure: Head: face (Fig. 30) strongly sulcate between pyramiform dorsal projections; postclypeus weakly trilobed, each lateral lobe with 3 small callosities; foliate lobes strongly developed. Thorax: Pronotum (Fig. 31): humeral angles well developed, blunt; median dorsal horn lat- erally compressed; suprahumeral horns re- duced to low carinae at base of median horn. Scutellum (Fig. 31): apex acuminate, lacking median dorsal groove. Legs (Fig. 32): metathoracic femur with | dorsal cu- cullate seta; metathoracic tibia with weakly cucullate setae in rows I, II, and III. Fore- wing (Fig. 33): r-m, and r-m, distad of fork of vein M. Genitalia: 2"! valvulae (Fig. 34) broadened at midpoint, tapering to apex; 880 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 30-34. pronotum, and scutellum, anterolateral aspect. 32, Left metathoracic femur, tibia, and tarsus, ablateral aspect. 33, Right forewing. 34, Female second valvulae, lateral aspect. dorsal ridge of broadened area serrate. d: unknown. Nymph.—Unknown. Distribution.—Dominican [SHMC/USNM]. Material examined.—Holotype [@, dis- sected] [iSHMC,, on andefinite, loan to USNM] with labels “DOM. REP, Dis. Nac.,/2 km. E. Boca Chica/August 11, 197 9/E.W. © Brien,’ VOUCHER. SPEG- IMEN/Cuticular Hydrocarbon/Analysis Study/Jason R. Cryan, 1995,” ‘“HOLO- TY PE/Togotolania/longicorna/Cryan & Deitz;.and)SHM-=’ Republic Togotolania longicorna. Structures of the holotype. 30, Head, anterior aspect (face). 31, Head, Remarks.—Togotolania longicorna, de- scribed from a single female specimen, is unique in its large, median dorsal horn on the crest of the pronotal metopidium. The suprahumeral carinae are located at the base of this horn; in some other treehopper groups with a dorsal pronotal elongation, the suprahumeral horns or carinae are at or near the apex (for example, the genus Ly- coderes Germar; see Cryan and Deitz 1999b: fig. 2). The Latin specific name ‘“longicorna”’ translates as “long horned,” referring to the median anterior pronotal horn. VOLUME 104, NUMBER 4 881 Figs. 35-39. Togotolania brachycorna. Structures of the holotype. 35, Head, anterior aspect (face). 36, Head, pronotum, and scutellum, anterolateral aspect. 37, Right forewing. 38, Male left lateral plate and pygofer, lateral aspect. 39, Male aedeagus and left style, lateral aspect. Togotolania brachycorna Cryan and Diagnosis.—The median, anterior pro- Deitz, new species notal horn of Togotolania brachycorna is (Figs. 35—39) short and weakly developed; the pronotum Type locality—Riv. Corossol, Guade- is mottled brown and tan, with a pale stripe loupe. encircling the median horn. 882 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Adult ¢.—Dimensions (mm): total length 7.2; width between humeral angles 2.3; pronotal length 1.9; wing length 5.6; width of head including eyes 2.2. Colora- tion: face, body, pronotum, and legs mot- tled brown and tan; pronotum with pale strip encircling median horn; basal % and venation of forewing dark brown. Struc- ture: Head: face (Figs. 35-36) strongly sulcate between large, emarginate dorsal projections; postclypeus unilobed; foliate lobes rounded. Thorax: Pronotum (Fig. 36): humeral angles well developed, blunt; median dorsal horn weakly developed; su- prahumeral horns absent. Scutellum (Fig. 36): apex acuminate, lacking median dorsal groove. Legs: metathoracic femur with | dorsal cucullate seta; metathoracic tibia with minute cucullate setae in rows I, II, and Ill. Forewing (Fig. 37): r-m, and r-m, basad of fork of vein M. Genitalia: lateral plates (Fig. 38) large, free, lacking apical hook; styles (Fig. 39) slender, hooked api- cally; aedeagus (Fig. 39) strongly U- shaped, with posterior arm tapered and lacking denticles on anterior face. 2: un- known. Nymph.—Unknown. Distribution.—Guadeloupe MNHN]. Material examined.—Holotype [d, dis- sected, with hindwings and | leg mounted beneath specimen] [CIRAD, deposited at MNHN] with labels ““GUADELOUPE/For- ét, PETIT-BOURG/Riv. Corossol/20-X- 1962/J. Bonfils,” “*3, and ““HOLOTYPE/ Togotolania/brachycorna/Cryan & Deitz.” Remarks.—Togotolania brachycorna, described from a single male specimen, has a much shorter median pronotal horn rela- tive to its congener (described above). The suprahumeral horns are lacking, and the pronotal coloration differs greatly from that of T. longicorna. Based on morphological variation and geographical separation, we believe the possibility 1s remote that 7. bra- chycorna is the male of 7. longicorna. The Latin specific name ““brachycorna”’ trans- [CIRAD/ lates as “short horned,”’ referring to the me- dian anterior pronotal horn. DISCUSSION AND SUMMARY The genus Deiroderes (with three spe- cies) is potentially allied to Antillotolania within the subfamily Stegaspidinae, al- though tribal association is unclear. Despite some similarities with the genera Melizo- deres and Llanquihuea (family Melizoder- idae; Linnavuori and DeLong 1978a, Deitz and Dietrich 1993a), Holdgatiella and To- gotolania (each with two species) are ap- parently among the basal lineages within Membracidae, probably allied with the sub- family Nicomiinae (Dietrich et al. 2001a; Cryan et al. in press). Further studies may confirm that one or two new family group taxa are needed to accommodate these tree- hoppers. ACKNOWLEDGMENTS We are grateful to R. L. Blinn, H. H. Neunzig, B. M. Wiegmann, and C. H. Die- trich, who offered many helpful sugges- tions, and to S. H. McKamey for bringing new taxa to our attention. G. Kuschel kind- ly provided etymological assistance regard- ing “Deiroderes.”’ For lending specimens, we are indebted to: the late J. A. Ramos (JARC); the late J. W. Evans (JWEC); R. L. Blinn (NCSU); S. H. McKamey (SHMC); the late R. C. Froeshner (USNM); and K. G. A. Hamilton (CNCI). This work is based in part on portions of the 1995 thesis submitted by J.R.C. in par- tial fulfillment of his M.S. degree in ento- mology (North Carolina State University). The material is based upon work supported in part by the North Carolina Agricultural Research Service (NCSU, Raleigh, North Carolina) and by the National Science Foundation Grants DEB-9815867 and DEB-9978026. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the authors and do not necessarily reflect the views of the North Carolina Agricultural Research Ser- vice or the National Science Foundation. VOLUME 104, NUMBER 4 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. 2nd Edit. Flora & Fauna Handbook 11. Sandhill Crane Press, Gainesville. 310 pp. Brown, R. W. 1956. Composition of Scientific Words. Smithsonian Institution Press, Washington, D.C. 882 pp. Cryan, J. R. and L. L. Deitz. 1999a. Review of the New World treehopper tribe Stegaspidini (Hemip- tera: Membracidae: Stegaspidinae): I: Bocydium Latreille, Lirania Stal, and Smerdalea Fowler. Proceedings of the Entomological Society of Washington 101(3): 469-489. . 1999b. Review of the New World treehopper tribe Stegaspidini (Hemiptera: Membracidae: Ste- gaspidinae): IH: Lycoderes Germar, Oeda Amyot and Serville, and Stegaspis Germar. Proceedings of the Entomological Society of Washington 101(4): 760-778. ——.. 2000a. Review of the New World treehopper tribe Stegaspidini (Hemiptera: Membracidae: Ste- gaspidinae): Ill: Flexocentrus Goding, Stylocen- trus Stal, and Umbelligerus Deitz. Proceedings of the Entomological Society of Washington 102(1): 82-98. Cryan, J. R., J. A. Robertson, and L. L. Deitz. In press. The New World treehopper tribe Microcentrini (Hemiptera: Membracidae: Stegaspidinae): Mono- graphic revision and phylogenetic position. Thom- as Say Publications, Entomological Society of America. Cryan, J. R., B. M. Wiegmann, L. L. Deitz, and C. H. Dietrich. 2000a. Phylogeny of the treehoppers (In- secta: Hemiptera: Membracidae): evidence from two nuclear genes. Molecular Phylogenetics and Evolution 17: 317-334. Deitz, L. L. 1975a. Classification of the higher cate- gories of the New World treehoppers (Homoptera: Membracidae). North Carolina Agricultural Ex- periment Station Technical Bulletin 225: [i-iv], 1— iT: 1989a. Bibliography of the Membracoidea (Homoptera: Aetalionidae, Biturritiidae, Membra- cidae, and Nicomiidae) 1981-1987. North Caro- 883 lina Agricultural Research Service Technical Bul- letin 290: 1-31. Deitz, L. L. and C. H. Dietrich. 1993a. Superfamily Membracoidea (Homoptera: Auchenorrhyncha). I. Introduction and revised classification with new family-group taxa. Systematic Entomology 18: 287-296. Deitz, L. L. and D. D. Kopp. 1987a. Bibliography of the Membracoidea (Homoptera: Aetlionidae, Bi- turritiidae, Membracidae, and Nicomiidae) 1956— 1980. North Carolina Agricultural Research Ser- vice Technical Bulletin 284. [ii] + 39 pp. Dietrich, C. H. and L. L. Deitz. 1993a. Superfamily Membracoidea (Homoptera: Auchenorrhyncha). II. Cladistic analysis and conclusions. Systematic Entomology 18: 297-311. Dietrich, C. H., S. H. McKamey, and L. L. Deitz. 2001a. Morphology-based phylogeny of the tree- hopper family Membracidae (Hemiptera: Cicado- morpha: Membracoidea). Systematic Entomology 26: 213-239. Evans, J. W. 1962a [**19617"]. Leafhoppers from Chile collected by the Royal Society Expedition to Southern Chile, 1958/59 (Homoptera, Cicadello- idea). Annals and Magazine of Natural History (13)4: 513-517. Goding, F W. 1926a. New genera and species of Mem- bracidae. Transactions of the American Entomo- logical Society 52: 103-110. Hamilton, K. G. A. 1971b. Placement of the genus Microcentrus in the Aetalionidae (Homoptera: Ci- cadelloidea), with a redefinition of the family. Journal of the Georgia Entomological Society 6(4): 229-236. Linnavuori, R. E. and D. M. DeLong. 1978a. The tree- hoppers (Homoptera: Membracidae) known to oc- cur in Chile. Brenesia 14-15: 171-194. 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 S. C. Bruner. 1925b. Membracidae of Cuba. Brooklyn Entomological Society Bulle- tin 20: 203-214. 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. Ramos, J. A. 1957a. A Review of the Auchenorhyn- chous [sic] Homoptera of Puerto Rico. Journal of Agriculture of the University of Puerto Rico 41(1): 38-117. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 884—895 THE SAWFLY GENUS KAMBAITINA MALAISE (HYMENOPTERA: TENTHREDINIDAE) IN INDIA MALKIAT S. SAINI, DAVID R. SMITH, AND V. VASU (MSS, VV) Department of Zoology, Punjabi University, Patiala-147002, India; (DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, Smithsonian Institution, Wash- ington, DC 20560-0168, U.S.A. Abstract.—The southeastern Asian genus Kambaitana is reviewed. Five new species from India are described: K. clypeata, K. lineata, K. impunctata, K. pup, and K. cardiaca. Kambaitana fulvipicta Malaise from Myanmar and India is redescribed. Males of the genus are characterized for the first time. A key for all seven known species, including K. albipicta Malaise from Myanmar, is provided. Key Words: The genus Kambaitina is known only from Myanmar and India. It was described by Malaise (1961) who included two new species, K. fulvipicta and K. albipicta, both from Myanmar. Saini and Deep (1994) first recorded the genus and K. albipicta from India. The genus belongs in the subfamily Allantinae as defined by Smith (1979), and in which it was subsequently included by Abe and Smith (1991). Here we describe five additional species from India, describe the males of the genus for the first time, and give a key to all seven species. Host plants are not known for the genus. Species may be associated with rosaceous plants such as Rubus spp. or Fragaria spp. as are many related allantines in the North- ern Hemisphere. Types of new species are deposited in Pusa National Collections, Division of En- tomology, Indian Agricultural Research In- stitute, New Delhi. Paratypes are in Punjabi University, Patiala, India (PUP) and the Na- tional Museum of Natural History, Smith- sonian Institution, Washington, DC (USNM). Kambaitina, India, Myanmar, Tenthredinidae, Allantinae Abbreviations used in text are: EL = eye length; [ATS = inner apical tibial spur; [CD = intercenchri distance; IDMO = intero- cular distance at level of median ocellus; ITD = intertegular distance; LID = lower interocular distance; MB = metabasitarsus; OATS = outer apical tibial spur; OCL = oculooccipital distance; OOL = oculoocel- lar distance, POL = postocellar distance. Kambaitana Malaise Kambaitina Malaise 1961: 253—255; Mal- aise 1963: 192 (in key); Abe and Smith 1991: 41, 103. Type species: Kambaitina fulvipicta Malaise, by original designa- tion. Description.—Antenna long, 2.7—3.2X head width, hardly tapering towards apex; third segment equal to or shorter than fourth segment; scape longer than its apical width; pedicel almost as long as broad. Inner mar- gins of eyes subparallel, scarcely converg- ing below; lower interocular distance great- er than eye length (Fig. 2). Clypeus deeply, roundly incised % to % its medial length, with blunt lateral lobes (Fig. 2); frontal area VOLUME 104, NUMBER 4 885 roundly elevated above a tangent touching both eyes; postocellar area subconvex, broader than long (Fig. 3). Malar space about equal to diameter of front ocellus. Genal carina developed only near base of mandible. Mandibles asymmetrical, right mandible with a large subapical tooth, left mandible with minute subapical tooth, al- most simple. Metabasitarsus shorter than following segments combined; tarsal claws with long subapical tooth, slightly shorter than outer tooth; basal lobe absent. Fore- wing (Fig. 1) with crossvein 2r present; Rs present, therefore with 4 cubital cells; anal cross vein (a) oblique (70°). Hindwing (Fig. 1) with cell 1M closed, cell IRs open; anal cell (1A) petiolate, petiole shorter than cell width; cu-a meeting petiole of anal cell at a slightly oblique angle. Ovipositor well sclerotized, long and slender (Fig. 4). Discussion.—Kambaitana belongs to the group of genera Malaise (1961) called the Monostegidea group, which includes Em- pronus Malaise, Monostegidea Rohwer, and Kambaitia. All are eastern Asian and Ori- Forewing and hindwing of Kambaitina cardiaca. ental. This group is characterized by the asymmetric mandibles, the right mandible with a large subapical tooth and the left mandible with only a minute tooth, almost simple (in other genera of Allantinae with asymmetric mandibles the left mandible has a large tooth and the right mandible is sim- ple); the long malar space; the closed cell M in the hindwing; the short metabasitar- sus; and the long antennae with a short third segment. Malaise (1961) gave a key to these four genera and defined Kambaitina by roundly incised clypeus with blunt lat- eral teeth; the hind orbits carinate only near the base of the mandibles; the scape and pedicel both longer than broad; the tarsal claw with inner tooth about as long as outer one and without a basal lobe; the subconvex mesoscutellum situated on the same level as all three mesonotal lobes; the hindleg with the tibia longer than the tarsus; and the lat- eral furrows of the head reaching the back of the head. The latter four characters dis- tinguish Kambaitana from Kambatia, the genus to which it is most similar. 886 3 front Figs. 2-3. view. 3, Head, dorsal view. Kambaitina cardiaca. 2, Head, Based on the tarsal claw character given by Malaise (1961), the placement of K. al- bipicta from Myanmar is not clear. He stat- ed that K. albipicta has “‘claw without basal lobe, with a very small one, then with a bristle.” Because species of Kambaitina lack a basal lobe, we believe K. albipicta may belong in Kambatia. Although this de- cision will require study of the type, we in- clude K. albipicta in the following key to species. Malaise (1961) did not have males of e1- ther of the two species he described. Be- cause we have available males of most spe- cies, we feel that the generic character “scape and pedicel distinctly longer than they are broad at apex,”’ should be changed to “scape distinctly longer than broad at apex.” In all males, the pedicel is almost as long as broad at its apex. i) . Antenna entirely black KEY TO SPECIES OF KAMBAITINA Antenna with a white band, segments 6 to 7 or more whitish yellow Clypeus roundly, narrowly, and deeply incised (Fig. 8); median fovea broad, deep and ditch- like; antennal segments 5—9 compressed (male genitalia as in Figs. 28, 34) .. Clypeus broadly, roundly, or subsquarely in- cised (Figs. 9-10); median fovea shallow, in form of narrow streak; antennal segments 4—9 COMpPressed’ "Le Gis paras Nes cs eee ee Postocellar area broader than long as 3:2, with a distinct medial longitudinal furrow on its an- clypeus subsquarely incised, with crenulate anterior margin (Fig. 10); labrum with rounded anterior margin (Fig. 10); inner half of tegula and lateral deflexed parts of ter- gite 2 whitish yellow; all sternites black; frons terior 34; minutely, densely punctured (male genitalia as in Figs. 29, 36) Postocellar area broader than long as 2:1, with- out median furrow; clypeus roundly incised, with smooth anterior margin (Fig. 9); labrum with pointed anterior tip (Fig. 9); tegula entire- ly black, lateral deflexed sides of tergites 2—4 and sternites 2—5 entirely whitish yellow; frons impunctate (female lancet as in Fig. 24; male cenitaltaas dn) Piss! 30545) tae oe einen Propodeum entirely black; antennal segments 6 and 7 whitish yellow, segments 8 and 9 com- pressed; clypeus roundly incised up to % of its medial length (Fig. 7) (female lancet as in Fig. 22 male cenitalia ‘asin Eigs4 27. 33) eee Posterior margin of propodeum whitish yellow; antennal segments with at least 6—8 whitish yellow, segments 6—9 compressed; clypeus roundly or subsquarely incised up to 2 of its medial length (Figs. 5—6) . Postocellar area broader than long as 2:1; lat- eral furrows not reaching hind margin of head; clypeus subsquarely incised (Fig. 6); only inner half of tegula whitish yellow; antennal seg- ments 3 and 4 equal (female lancet as in Fig. 25-amalecenitaliavassin) Bigs 265.52) ieee Postocellar area broader than long as 5:4; lat- eral furrows reaching hind margin of head; clypeus roundly incised (Fig. 5); tegula whitish and 4 as 5:6 ... Narrow posterior margins of tergites 2, 6—8 and yellow; antennal segments 3 broad posterior margin of tergite 9, antennal segments from middle of 5 to 8, and spot on temple whitish yellow; wing hyaline, tarsi 2—4 uniformly fuscoferruginous; tarsal claw with- K. clypeata, n. s K. lineata, n. K. impunctata, n. K. pup, n. K. cardiaca, n. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON i) sp. sp. sp. sp. VOLUME 104, NUMBER 4 887 Fig. 4. out basal lobe (female lancet as in Fig. 24; male genitalia as in Figs. 25, 31) cE) Ge uta ero ae eer aoa K. fulvipicta Malaise — Broad posterior margins of all tergites and an- tennal segments from middle of 4 to 8 whitish yellow; spot on temple fuscous; wings faintly yellowish; metatarsi 2—4 black with a white longitudinal stripe on posterior surface; tarsal claw with indistinct basal lobe ......... rs CRC ee cana re K. albipicta Malaise Kambaitina clypeata Saini, Smith, and Vasu, new species (Figs. 8, 11, 28, 34) Female.—Unknown. Male.—Color: Black with labrum, clyp- eus except extreme base, lower orbits, ex- treme posterodorsal and posterolateral mar- gins of pronotum, extreme apices of coxae, and trochanters whitish yellow; spot on me- soscutellum, femora, tibiae, protarsus, me- sotibia and mesotarsus fuscoferruginous. Wings hyaline; venation including costa, subcosta, and stigma piceous. Structure: Length 7.0 mm. Antennal length 3 head width, antennal segments 3 and 4 equal, apical 4 segments gradually decreasing in length as 8:7:6:4, segments 5— 9 compressed. Clypeus (Fig. 8) narrowly and roundly incised up to % its medial length with broadly rounded lateral teeth; Apex of abdomen, sawsheath, and ovipositor of Kambaitina fulvipicta. labrum (Fig. 8) with deflexed rounded an- terior margin, broader than long as 3:2. Su- praclypeal area triangularly raised with a sharp longitudinal carina; malar space 0.75 diameter of median ocellus; lower margin of eye at level of lower rim of an- tennal socket; LID:IDMO:EL = 3:3:2. Su- praclypeal and supraantennal pits well marked; supraantennal tubercles well de- veloped and confluent with similarly ele- vated frontal ridges; median fovea in form of prominent, broad groove, clearly reach- ing median ocellus; post-, inter-, and cir- cumocellar furrows sharp and distinct; lat- eral postocellar furrows deep, parallel, and abruptly ending just before hind margin of head; postocellar area broader than long as 4:3: head slightly widened behind eyes; OOL:POL:OCL = 3:2:3. Mesoscutellum subconvex, without carina, its appendage without carina; [CD:ITD = 1:2. Tarsal claw (Fig. 11); metabasitarsus subequal to fol- lowing segments combined as 6:7; IATS: MB:OATS = 4:12:3. Genitalia: Penis valve (Fig. 28), genital capsule (Fig. 34). Sculpture and pubescence: Head shining with few, sparse, fine, inconspicuous wrin- kles confined to frontal area. Thorax with minute scattered punctures except posterior 888 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5—20. 5—10, Clypeus and labrum. 5, Kambaitina fulvipicta. 6, K. cardiaca. 7, K. pup. 8, K. c/ypeata. 9, K. impunctata. 10, K. lineata. 11-16, Tarsal claw. 11, K. clypeata. 12, K. fulvipicta. 13, K. impunctata. 14, K. cardiaca. 15, K. pup. 16, K. lineata. 17-20, Hypopygium. 17, K. fulvipicta. 18, K. impunctata. 19, K. cardiaca. 20, K. pup. slope of mesoscutellum bearing numerous large, shallow punctures, surface shining. Abdomen impunctate, surface subshining. Body covered with blackish pubescence. Type.—Holotype 6, India, Manipur, Ukhruls 1, 800)m, 23.1%.1992: Distribution.—India: Manipur. Remarks.—The black antennae and char- acteristic shape of the clypeus which is roundly, narrowly, and deeply incised sep- arates this species from other species of this genus. Etymology.—The species name reflects the characteristic shape of the clypeus. Kambaitina lineata Saini, Smith, and Vasu, new species (Figs. 10, 16, 29, 36) Female.—Unknown. Male.—Color: Body black with labrum, clypeus, a square spot on supraclypeal area, inner orbit, lower hind orbit, spot on temple close to hind margin of head, broad poster- odorsal and posteroventral margins of pron- otum, most of tegula except an outer spot, posterior triangular apex of mesonotal mid- dle lobe, top of mesoscutellum, parapterum, broad spot in middle of mesepisternum, spot on metepisternum, deflexed lateral margin of tergite 2, procoxa except extreme base, apical % of mesocoxa, outer stripe and apical % of metacoxa, and all trochanters and adjacent parts of femora whitish yel- low; femora and tibiae and tarsi of all legs fuscoferruginous. Wings hyaline; venation including costa, subcosta, and stigma pice- ous. Structure: Average length 6.5mm. An- tennal length 3.2 head width, antennal segments 3 and 4 subequal as 8:9, apical 4 segments subequal as 6:5:4:4, segments 4— 9 compressed. Clypeus (Fig. 10) subrectan- VOLUME 104, NUMBER 4 gularly incised up to % its medial length with crenulate anterior margin and blunt lateral teeth. Labrum (Fig. 10) with de- flexed, rounded anterior margin, broader than long as 5:3. Supraclypeal area trian- gularly raised with a faint longitudinal blunt carina; malar space 0.75 diameter of front ocellus; lower margin of eye below level of antennal socket; LID:IDMO:EL = 3:3:2. Supraclypeal and supraantennal pits deep and well marked; supraantennal tubercles moderately raised and confluent with simi- larly raised frontal ridges; median fovea in form of shallow ditch with a very fine streak in middle, posteriorly not extending to median ocellus; post-, inter-, and circu- mocellar furrows sharp and distinct; lateral furrows deep, bulging medially and reach- ing hind margin of head, postocellar area subconvex, with shallow but distinct me- dian longitudinal furrow on anterior %4, broader than long as 3:2. Head parallel be- hind eyes; OOL:POL:OCL = 3:2:3. Me- soscutellum subconvex, ecarinate, its ap- pendage ecarinate; ICD:ITD = 1:2. Tarsal claw (Fig. 16); metabasitarsus shorter than following segments combined as 2:3; IATS: MB:OATS = 5:16:4. Genitalia: Penis valve (Fig. 29), genital capsule (Fig. 36). Sculpture and pubescence: Frontal area with minute irregular, scattered punctures, rest of head shining with dense, minute punctures. Thorax almost impunctate ex- cept few shallow punctures on _ posterior slope of mesoscutellum, surface shining with general oily lustre. Abdomen impunc- tate, shining. Body covered with brownish black pubescence. Types.—Holotype 6d India, Nagaland, Pfutsero, 2,000 m, 20.v. 1993. Paratypes: 1 3d, India, Nagaland, Vizho-Razho, 1,800 m, 11.v.1993; 2 5, same data as holotype. Distribution.—India: Nagaland. Remarks.—A combination of the follow- ing characters separate K. lineata from oth- er species of the genus: antenna and ster- nites entirely black; postocellar area with a distinct medial longitudinal furrow on an- terior %4, clypeus subsquarely incised; inner 889 half of tegulae whitish yellow; and antennal segments 4—9 distinctly compressed. Etymology.—The species name is based on the presence of a distinct medial longi- tudinal furrow on the anterior %4 of posto- cellar area. Kambaitina impunctata Saini, Smith, and Vasu, new species (Figs. 9, 13, 18, 24.730%35) Female.—Color: Black with labrum, clypeus, triangular spot on supraclypeal area, basal 4’ of mandible, narrow inner or- bit, spot on mesonotal lateral lobe before mesoscutellum, spot on mesoscutellum, broad transverse stripe on lower half of me- sepisternum narrowly extending to coxal rim, lateral deflexed sides of tergites 2—4, sternites 2—5, and coxae and _ trochanters whitish yellow; femora, tibiae, and tarsi ful- vous. Wings hyaline; venation including costa, subcosta, and stigma piceous. Structure: Length 7.0 mm. Antennal length 3.2 head width, segments 3 and 4 subequal as 5:6, length of apical four seg- ments gradually decreasing as 8:7:6:5, fla- gellar segments 4—9 faintly compressed. Clypeus (Fig. 9) roundly incised up to % its medial length with blunt lateral teeth. La- brum (Fig. 9) with deflexed pointed anterior tip, broader than long as 3:2. Supraclypeal area protuberant; malar space 0.75 diam- eter of front ocellus; lower margin of eye below level of antennal socket; LID: IDMO: EL = 3:3:2. Supraclypeal and supraanten- nal pits deep and well marked, supraanten- nal tubercles moderately raised and conflu- ent with similarly roundly raised frontal ridges; median fovea in form of deep, nar- row, streak extending to median ocellus, post-, inter-, and circumocellar furrows sharp; lateral furrows deep and ending abruptly just before hind margin of head: postocellar area broader than long as 2:1. Head parallel behind eyes; OOL:POL:O0CL = 3:2:3. Mesoscutellum subconvex, ecari- nate, its appendage ecarinate; ICD:ITD = 2:5. Tarsal claw (Fig. 13); metabasitarsus equal to following joints combined; [ATS: 890 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON - \ 29) aL, Figs 21-24. Lancet. 21, Kambaitina fulvipicta. MB:OATS 5:14:4. Lancet (Fig. 24) broad, with 18 serrulae, annular hairs dis- tinct full width of lancet. Hypopygium as in Fig. 18. Sculpture and pubescence: Head, thorax and abdomen impunctate and shining with general oily lustre. Body covered with brownish pubescence. Male.—Length 6.5 mm. Similar to fe- K. pup. 23, K. cardiaca. 24, K. impunctata. male except ventral surface of scape, lower orbits, ventral half of hind orbits, parapter- um, broad transverse band extending from anterior to posterior margin along convexity of mesepisternum whitish yellow. Lateral deflexed sides of tergite 4 and all sternites entirely black. Genitalia: Penis valve (Fig. 30), genital capsule (Fig. 35). Types.—Holotype ¢, India, West Ben- VOLUME 104, NUMBER 4 gal, Darjeeling, 2,280 m, 7.v.1993. Para- type: | d same data as holotype, deposited im PUP. Distribution.—India: West Bengal. Remarks.—Kambaitina impunctata is separated from other species by the follow- ing combination of characters: antenna black; clypeus roundly incised; labrum with pointed anterior tip; tegulae black; lateral deflexed parts of tergites 2—4 and sternites 2—5 entirely whitish yellow; and body im- punctate. The female lancet also is distinct from other known females of the genus; it is broader and has distinct annular hairs ex- tending its full width (Fig. 24). Females of the other species with black antennae are unknown, but it is possible this type of lan- cet is characteristic of species of this group, Whereas the lancets of the species with white-banded antennae are much more slen- der and lack annular hairs. Etymology.—The species name pertains to the impunctate body. Kambaitina pup Saini, Smith, and Vasu, new species (Riese 7 los, 20.22) 2 7.733) Female.—Color: Black, with antennal segments 6 and 7, labrum, clypeus, smail square spot on supraclypeal area, broad lower and narrow upper orbits discontinu- ous in middle, broad spot on lower hind orbits, spot on hind margin of head behind temple, broad posterodorsal and posterolat- eral margins of pronotum, tegula, apex of mesonotal middle lobe, anterior mesoscu- tellum, parapterum, broad spot in middle of mesepisternum, spot on posteroventral tip of mesepimeron, spot on metepisternum, deflexed lateral sides of all tergites, poste- rior margin of last sternite, procoxa except extreme base, apical % of meso- and me- tacoxae, and all trochanters and adjacent parts of all femora whitish yellow; femora, tibiae and tarsi fuscoferruginous. Wings hy- aline; venation including costa, subcosta, and stigma piceous. Structure: Average length 6.5 mm. An- tennal length 2.7 head width; segments 3 89] and 4 equal, apical four segments subequal as 8:7:6:5, segments 8 and 9 compressed. Clypeus (Fig. 7) roundly incised up to % of medial length with blunt lateral lobes. La- brum (Fig. 7) with deflexed rounded ante- rior margin, broader than long as 3:2. Su- praclypeal area protuberant; malar space equal to diameter of median ocellus; iower margin of eye below level of antennal sock- et; LID: IDMO:EL = 6:6:3. Supraclypeal and supraantennal pits deep and well marked; supraantennal tubercles moderate and confluent with similarly roundly raised frontal ridges; median fovea in form of broad and shallow groove marked by lon- gitudinal fine streak in its middle, posteri- orly shallowly reaching front ocellus; post-, inter-, and circumocellar furrows sharp and distinct; lateral furrows deep, protuberant, sunken and ending just before hind margin of head; postocellar area broader than long as 3:2. Head parallel behind eyes; OOL: POL:OCL = 3:2:2. Mesoscutellum subcon- vex, ecarinate, its appendage ecarinate. Tar- sal claw (Fig. 15); metabasitarsus shorter than following joints combined as 4:5; IATS:MB:OATS = 3:8:2. Lancet (Fig. 22) with 22 serrulae on about apical half of lan- cet; without annular hairs. Hypopygium as in Fig. 20. Sculpture and pubescence: Frontal area with dense, minute, irregular punctures in- termingled with network of irregular grooves and ridges, rest of head with fine, shallow punctures, surface shining. Thorax impunctate except posterior slope of me- soscutelium bearing few minute shallow punctures, surface shining with general oily lustre. Abdomen impunctate, subshining. Body covered with brownish pubescence. Male.—-Average length 6.5 mm. Similar to female except broad front and hind orbits whitish yellow and continuous with temple spot. Deflexed lateral sides of tergites 2—4 fuscous. Genitalia: Penis valve (Fig. 27), genitai capsule (Fig. 33). Types.—Holotype 2, India, Manipur, Ukhrul 1,700 m, 23.v.1993. Paratypes:1 ¢, Manipur, Zunheboto, 1,800 m, 14.V.19953; 892 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 25-36. Male genitalia. 25-30, Penis valve. 25, Kambaitana fulvipicta. 26, K. cardiaca. 27, K. pup. 28, K. clypeata. 29, K. lineata. 30, K. impunctata. 31-36. Gonoforceps. 31, K. fulvipicta. 32, K. cardiaca. 33, K. pup. 34, K. clypeata. 35, K. impunctata. 36, K. lineata. 1 2, 16.v.1993; 1 6, Nagaland, Vinzho-Ra- zho, 1,800 m, ll.v.1993; 1 od, Satakha, 12500) mane cli5v519935) ion? ose Piutser 2,100 m, 20.v.1993. Distribution.—India: Manipur, Nagaland. Variation.—The extreme base of anten- nal segment 6 and extreme tip of segment 7 may be fuscous; the deflexed lateral sides of tergites 2—4 in the male may be fuscous, and the outer margins of the tegulae may be piceous. Remarks.—Kambaitana pup is distin- guished from other species of the genus by the following: Antennal segments 6 and 7 whitish yellow; clypeus roundly incised for one-third its medial length; propodeum en- tirely black; and antennal segments 8 and 9 distinctly compressed. Etymology.—The species name is de- rived from the acronym of Punjabi Univer- sity, Patiala (PUP), where revisionary work on the taxonomy of sawflies is being carried out. It is used as a noun in apposition. Kambaitina cardiaca Saini, Smith, and Vasu, new species GBiess. 1—3,.65 145) 199 23426952) Female.—Color: Black with antennal segments 6—8, labrum, clypeus, square spot on supraclypeal area; broad inner orbit, tri- angular spot on hind margin of head behind temple, broad posterodorsal and posterolat- eral margins of pronotum; inner half of te- gula, apex of mesonotal middle lobe, top of mesoscutellum, broad spot on metascutel- lum, parapterum, broad spot on lower % of mesepisternum, spot on posterior tip of me- sepimeron, paired spot on each metepister- num and metepimeron, spot on metaster- num, extreme posterior margin of propo- deum and tergites 2, 8 and 9, deflexed lat- eral sides of all tergites, broad medial spot on sternites 2—6, sternite 7 more or less, pro- and mesocoxae except extreme bases, outer surface and apical % of metacoxa, and all trochanters and adjacent parts of all fem- VOLUME 104, NUMBER 4 ora whitish yellow; femora, tibiae, and tar- sal segments fuscoferruginous. Wings hya- line; venation including costa, subcosta and stigma piceous. Structure: Average length 9.5 mm. An- tenna 2.7 head width, segments 3 and 4 equal in length, apical 4 segments subequal as 8:7:6:5, segments 6—9 compressed. Clyp- eus (Figs. 2, 6) subsquarely incised up to 4% its medial length with blunt lateral teeth. Labrum (Figs. 2, 6) with deflexed rounded anterior margin, broader than long as 3:2. Supraclypeal area protuberant with faint longitudinal carina; malar space equal to di- ameter of median ocellus, lower margin of eye below level of antennal socket; LID: IDMO:EL = 6:6:3. Supraclypeal and su- praantennal pits well marked; supraantennal tubercles moderately raised and confluent with similarly raised frontal ridges; frontal anterior to front ocellus broadly and round- ly heart shaped, gradually and only very faintly depressed towards center, with me- dian fovea in form of narrow streak shal- lowly reaching median ocellus; postocellar furrows sunken; inter- and circumocellar furrows sharp and distinct; lateral furrows deep and abruptly ending well before hind margin of head; postocellar area broader than long. Head parallel behind eyes; OOL: POL:OCL = 3:2:3. Mesoscutellum subcon- vex, ecarinate, its appendage ecarinate; ICD:ITD = 2:5. Tarsal claw (Fig. 14); me- tabasitarsus subequal to following segments combined as 8:9; IATS:MB:OATS = 3:8:2. Lancet (Fig. 23) with 23 serrulae on about apical half of lancet; annular hairs absent. Hypopygium as in Fig. 19. Sculpture and pubescence: Frontal area with dense, minute, irregular punctures, rest of head with sparse, fine, punctures, surface shining. Thorax impunctate except posterior slope of mesoscutellum bearing few minute punctures, surface shining with general oily lustre. Abdomen impunctate, dull. Body covered with fuscous pubescence. Male.—Average length 7.5 mm. Similar to female except deflexed lateral margin of 893 tergites 5—9 black. Genitalia: Penis valve (Fig. 26); genital capsule (Fig. 32). Types.—Holotype: @, India, Manipur, Ukhrul, 1,700 m, 21.v.1993. Paratypes: 4 2, 6 3, Manipur, Ukhrul, 1,700 m, 21.v: 1993; 1 2, 2 3, Nagaland, Vizho-Razho, 1,800 m, Il.v.1993; 1 ¢d, Zunheboto, 1,800 m, 14.v.1993; 6 2, 2 6, Satakha, 1,500 m, [I5.V.1993;"3, 2, 246, Prutsero; 2000 m: 9 v-1993, 15,4 95:3 SJ 20n 1998: Distribution.—India: Manipur, Nagaland. Variation.—Antennal segments 7 and 8 may be more or less fuscoferruginous. Remarks.—Kambaitana cardiaca 1s dis- tinguished from other species of the genus by antennal segments 6—8, the inner half of the tegulae, the posterior margins of tergites 2, 8 and 9, and the medial spot on sternites 2—6 whitish yellow; the subsquarely incised clypeus with the emargination half its me- dial length; and the lateral furrows ending before the hind margin of the head. Etymology.—The species name is de- rived from the peculiar shape of the frontal area. Kambaitina fulvipicta Malaise (Bigs. 4.9, 27s 2 25.51) Kambaitina fulvipicta Malaise 1961: 254— 255; Saini and Deep 1994: 49. Female.—Color: Black with antennal segments 6—8, labrum, clypeus, square spot on supraclypeal area, basal ’% of mandible, inner orbits, lower half of hind orbit, spot on temple, broad posterodorsal and postero- lateral margins of pronotum, tegula, apex of mesonotal middle lobe, top of mesoscutel- lum, broad spot on metascutellum, parap- terum, broad transverse spot on lower half of mesepisternum, posterior margin of pro- podeum, extreme posterior margins of ter- gites 2 and 6—8 and broad posterior margin of tergite 9, deflexed lateral sides of all ter- gites, sternites 2—7 more or less, pro- and mesocoxae and trochanters, apical /% of me- tacoxa, and metatrochanter whitish yellow; femora, tibiae, and tarsi fuscoferruginous. 894 Wings hyaline; venation including costa, subcosta, and stigma piceous. Structure: Average length 8.0 mm. An- tennal length 3.2 head width; segments 3 and 4 subequal as 5:6, length of apical four segments gradually decreasing as 8:7:6:5, apical segments 6—9 faintly compressed. Clypeus (Fig. 5) roundly incised for % its medial length with blunt lateral lobes. La- brum (Fig. 5) with deflexed rounded ante- rior margin, broader than long as 3:2. Su- praclypeal area protuberant; malar space 1.25 diameter of front ocellus; lower mar- gin of eye below level of eye; LID: IDMO: EL 3:3:2. Supraclypeal and supraantennal pits well marked; supraantennal tubercles moderately raised and confluent with simi- larly roundly raised frontal ridges; median fovea in form of narrow depressed area with a pit on its anterior half and posteriorly only shallowly reaching front ocellus; post- , Inter-, and circumocellar furrows distinct; lateral furrows deep, bulging medially and reaching hind margin of head; postocellar area broader than long as 5:4. Head parallel behind eyes; OOL:POL:OCL = 6:4:7. Me- soscutellum aimost flat, ecarinate, its ap- pendage ecarinate; ICD:ITD = 1:3. Tarsal claw (Fig. 12); metabasitarsus shorter than following segments combined as 3:4; [ATS: MB:OATS = 8:20:7. Lancet (Figs. 4, 21) with 10 serrulae on only apicai half; annular hairs absent. Hypopygium as in Fig. 17. Sculpture and pubescence: Frontal area Opaque with dense and minute punctures, rest of head shining with scattered minute punctures. Thorax and abdomen shining, impunctate with general oily lustre. Body covered with blackish pubescence. Male.—Average length 7.5 mm. Simiiar to female except antennal segment 6 and 7 fuscoferruginous. Genitalia: Penis valve (Fig. 25), genital capsule (Fig. 31). Types.— (holotype), | 2 (paratype) at Naturhistoriska Riksmuseet, Stockholm. Specimens examined.—2 2, 1 6, India, Himachal Pradesh, Kasol, 1,600 m, 29.v.1984; 2 2, West Bengal, Darjeeling, 2,280 m, 2.v.1986; 1 6, Uttar Pradesh, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ghangria, 3,000 m, 27.vi.1987; 1 2, Chop- ta, 3,000 m, 25.vi.1992; 1 92, Arunachal Pradesh, Nine mile, 2,000 m, 24.5.1993. Distribution.—Myanmar; India: Hima- chal Pradesh, West Bengal, Uttar Pradesh, Arunachal Pradesh. Variation.—The apical half of antennal segment 5, the pronotum almost entirely, and the mesoscutellar appendage more or less may be whitish yellow. The apical half of antennal segment 8 may be black. Remarks.——This species is characterized by the whitish yellow antennal segments 6— 8, tegulae, narrow posterior margins of ter- gites 2 and 6—8, broad posterior margin of tergite 9, and sternites 2—7; roundly incised clypeus which is incised for half its medial length; subquadrate postocellar area; and lateral furrows reaching the hind margin of head. The female lancet is distinct from other known females by having only ten serrulae and only on the apical half rather than on most of its length (Fig. 21). The specimens studied go to Kambaitana in Malaise’s keys (1961, 1963) and agree with the original description (1961). The male is associated with the female for the first time. ACKNOWLEDGMENTS The financial assistance rendered by the US, PL-480, Project No. IN-ARS-418, in collaboration with ICAR, New Delhi, is thankfully acknowledged. Cathy Anderson, Systematic Entomology Laboratory, U.S.D.A., took the photos and prepared the plates. We thank N. M. Schiff, U.S. Forest Service, Stoneville, MS, and D. A. Nickle and J.W. Brown, Systematic Entomology Laboratory, Beltsville, MD, and Washing- ton, DC, respectively, for reviewing the manuscript. LITERATURE CITED Abe, M. and Smith, D. R. 1991. The genus-group names of Symphyta (Hymenoptera) and their type species. Esakia 31, 115 pp. Malaise, R. 1961. New Oriental saw-filies (Hymen.: Tenthr.). Entomologisk Tidskrift 82(3—4): 231}— 260. VOLUME 104, NUMBER 4 895 Malaise, R. 1963. Hymenoptera Tenthredinoidea, dia. Journal of the Bombay Natural History So- subfamily Selandriinae. key to the genera of the ciety 91: 47-50. World. Entomologisk Tidskrift 84(3—4): 159— Smith, D. R. 1979. Nearctic Sawflies IV. Allantinae: PSY: Adults and larvae (Hymenoptera: Tenthredinidae). Saini, M. S. and J. S. Deep. 1994. First record of Al- United States Department of Agriculture, Techni- lantinae (Tenthredinidae: Hymenoptera) from In- cal Bulletin 1595, 172 pp. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 896-911 SYSTEMATIC STUDIES ON THE ANTONINA CRAWI COCKERELL (HEMIPTERA: COCCOIDEA: PSEUDOCOCCIDAE) COMPLEX OF PEST MEALYBUGS DOUGLAS J. WILLIAMS AND DOUGLASS R. MILLER (DJW) Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.; (DRM) Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, Rm. 137, Bldg. 005, BARC-W, Beltsville, MD 20705, U.S.A. (e-mail: dmiller@sel.barc.usda.gov). Abstract.-Careful analysis of specimens identified as Antonina crawi Cockerell has shown that three very similar species are involved, i.e., A. crawi, A. nakaharai Williams and Miller, new species, and A. socialis Newstead (reinstated status). A key is provided for 11 species reported on bamboos and a description is given for a bamboo-infesting species from Taiwan, i.e., A. maai Williams and Miller, new species. Antonina crawi 1s reported from China, Japan, and the U.S. (California) but is suspected to have disappeared from the latter country. Key Words: Coccoidea, Pseudococcidae, new species, Antonina maai, Antonina naka- harai, redescribed species, Antonina crawi Cockerell, Antonina socialis Newstead, bamboo, key As part of a study on the mealybugs of southern Asia, the first author noted an un- usually large amount of morphological var- iation in Antonina specimens determined as A. crawi Cockerell. After careful analysis of these specimens it is evident that what had originally been considered a single tax- on is actually three different species. Be- cause mealybugs in the Crawi complex are primarily bamboo feeders and because bamboos are frequently used as ornamen- tals, we examined as many Antonina spec- imens as possible from that host and pro- vide a key to all of the species that occur on bamboo. Antonina includes 17 valid species, all of which are native to the Old World (Ben- Dov et al. 2001), particularly Asia. They are: A. australis Froggatt (Australia), A. bambusae Khalid and Shafee (India), A. crawi Cockerell (probably native to Japan and China), A. elongata Tang (China), A. graminis (Maskell) (possibly native to In- dia), A. maritima Ramakrishna Ayyar (In- dia and Sri Lanka), A. meghalayaensis Khalid and Shafee (India), A. natalensis Brain (possibly Africa), A. pretiosa Ferris (Asia), A. purpurea Signoret (Europe), A. tesquorum Danzig (northeastern Asia), A. thaiensis Takahashi (Malaysia and Thai- land), A. transcaucasica (Borchsenius) (Ar- menia and Azerbaijan), A. transvaalensis Brain (South Africa), A. vera Borchsenius (northeastern Asia), and A. zonata Green (southern Asia). Antonina indica panica Hall recently has been considered a valid species (Williams 2001), 1.e., A. panica. Several species have become invasive pests in various parts of the world, partic- ularly A. crawi (actually A. nakaharai Wil- liams and Miller, n. sp. and A. socialis Newstead), A. graminis, and A. pretiosa VOLUME 104, NUMBER 4 (Miller et al. 2002). A comprehensive book was written by Hendricks and Kosztarab (1999) on the legless mealybugs including detailed morphological treatments and a key for the adult females of all but 4 species of Antonina. They were unable to locate specimens of A. bambusae, A. elongata, A. meghalayaensis, and A. transcaucasica. The immature stages of several species were treated by Yang and Kosztarab (1967), and adult males were described by Afifi and Kosztarab (1967) and Beardsley (1965). We have found I|1 species that occur on bamboo including: A. bambusae (Bambu- sa), A. crawi (see later description for list of bamboo hosts), A. graminis (Bambusa sp.), A. maai Williams and Miller, n. sp. (see later description for lst of bamboo hosts), A. meghalayaensis (Bambusa sp.), A. nakaharai Williams and Miller, n. sp. (see later description for list of bamboo hosts), A. pretiosa (Arundinaria, Bambusa, Phyllostachys, Pleioblastus, Sasa, and Yu- shania), A. purpurea (Phyllostachys), A. so- cialis (see later description for list of bam- boo hosts and corrected status), A. thaiensis (Bambusa), A. zonata (Bambusa, Indoca- lamus, Semiarundinaria, Teinostachyum). It is interesting that 9 of these species are spe- cific to bamboos and only 2 species are in- cidental on bamboo. The latter 2 species (A. graminis and A. purpurea) are more general feeders occurring on a wide range of grass hosts. The three species encompassing the Cra- wi complex are here determined as A. cra- wi, A. nakaharai, and A. socialis. These species are considered a complex because they have so frequently been confused. There are no characters unique to this group that do not also occur in other species of Antonina. Although Antonina maai is sim- ilar to species in the Crawi complex, it is most similar to A. pretiosa. MATERIALS AND METHODS Depositories of specimens are: The Nat- ural History Museum, London (BMNH); California Department of Food and Agri- 897 culture, Sacramento (CDFA); Florida State Collection of Arthropods, Gainesville (FSCA); Muséum National d’ Histoire Na- turelle, Paris (MNHN); Taiwan Agricultural Research Institute, Entomology Collection, Taichung, Taiwan (TARI); National Muse- um of Natural History, Beltsville, MD (USNM); R. M. Bohart Museum, Univer- sity of California, Davis, CA (UCD). Abbreviations used in the Specimens Ex- amined sections are: ad. = adult; imm. = immature; sl. = slide. Measurements of the labium are the lon- gest dimension of the anterior to the pos- terior edges of the structure as it appears on slide-mounted specimens. In some instanc- es the labium is oriented with the apex pointing directly at the microscope slide (ventrally) rather than pointing posteriorly, as is the case for most mealybugs. Thus, the measurement in these instances is not al- ways taken of the lateral profile of the la- bium but instead is of the largest diameter of the structure. Measurements of the anal tube are taken on the dorsal surface from the anterior end of the anal ring to the open- ing of the anal tube nearest to the anal ring; this measurement is the shortest length. Also note that there is a band of pores and tubular ducts internally near the entrance of the anal tube. These structures are not shown in the illustrations. Inclusion of Antonina bambusae and A. meghalayaensis in the key to Antonina spe- cies that occur on bamboo is based on ex- amination of the original descriptions only; we have been unable to locate specimens of these species. Hendricks and Kosztarab (1999) questioned the validity of some of the statements made by Khalid and Shafee (1988) in their descriptions of these species, including the presence of a circulus in A. bambusae. RESULTS Antonina crawi Cockerell (Fig. 1) Antonina crawi Cockerell 1900:70. Antonina crawit: Ferris 1918:77 (misspell- ing). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 898 ( y ( +8 oe 1D) ae or Se) ne ° eye ee | es “° O Ove ° 6 a Ove Ouane O ive oltre} 7 vere © te Specimen from Japan, near Tokyo, on Arundinaria fastuosa. 1. , Antonina craw ener, ile VOLUME 104, NUMBER 4 Antonina bambusae: Borchsenius 1934:13 (misidentification). Type material._Lectotype adult female single specimen on slide labeled as follows: left label ‘“‘Antonina/crawi CkII/TY PE/ Bamboo/Japan (Craw) 1900/CkIl. Coll.;” right label “LECTOTYPE/Desig. H. Hen- dricks/1992/Antonina crawi CklIl./Re- mounted: Balsam.” In addition there is an additional slide containing 2 adult female paralectotypes (all in USNM). The lecto- type was designated by Hendricks and Kosztarab (1999). Description.—Adult female on micro- scope slide, elongate oval, 1.4—4.5(3.2) mm long, 0.7—2.9(1.8) mm wide; posterior apex of last 2 or 3 abdominal segments sclero- tized (more mature paratypes sometimes with more sclerotization); lateral margins of abdominal segments VII—VIII convex, rare- ly segment VI slightly convex, others straight sided. Antenna 2- or 3-segmented, 45—62(54) » long. Legs usually represented by inconspicuous sclerotized area or small dermal pocket, sometimes absent. Labium 55—90(80) long, between 2 and 3 times shorter than clypeolabral shield. Ostioles present on abdomen only. Circulus absent. Spiracles with 2 sizes of trilocular pores in sclerotized area surrounding atrium. Cerarii absent, but posterior abdominal setae en- larged, subconical. Anal ring invaginated in anal tube, 145—175(162) pw long; ring di- ameter 88—130(105) 2; anal tube internally with ring of multilocular pores and tubular ducts. With 28—66(42) multilocular pores on each side of body in area delimited by posterior edges of each anterior and poste- rior spiracle. Dorsal surface with short setae, decreas- ing in length and width anteriorly; longest seta on posterior apex 32—40(36) wp long. Trilocular pores of 1 size, absent from pos- terior 3 or 4 segments, absent along body margin except near spiracles, scattered over remainder of surface. Discoidal pores of same distribution pattern as triloculars, less abundant. Multilocular pores forming lon- 899 gitudinal band along body margin; with O— 7(3) pores on segment VII near interseg- mental line between VII and VIII. Oral-col- lar tubular ducts of | variable size, absent or rare on segment VIII except in anal tube, abundant Over remainder of surface. Ventral surface with setae similar to those on dorsum. Multilocular disc pores present in band along body margin, with 6— 24(14) pores on abdominal segment IV within area delimited by lateral margin of disc-like pore clusters on each side of seg- ment; with multilocular pores on segment VIII often spanning space between vulva and posterior apex of body, occasionally re- stricted to half or three-fourths of distance between vulva and posterior apex of body; dermal surface surrounding opening of anal tube with cluster of multilocular pores and oral-collar tubular ducts. Disc-like pores present in mediolateral areas from metatho- rax to segments IV, V, or rarely VI. Triloc- ular pores of 2 or 3 sizes, smaller size in plate outside of spiracular atrium; larger size scattered around mouthparts, medium size scattered over remainder of surface ex- cept absent or rare on posterior 3 or 4 seg- ments, most abundant near body margin. Oral-collar tubular ducts present over sur- face. Vulvar area with 1 pair of lateral apophyses and | pair of weakly indicated posterior apophyses. Notes.—This species is similar to Anton- ina nakaharai and A. socialis but differs by having: 6—24(14) multilocular pores on ab- dominal segment IV within the area delim- ited by the lateral margins of disc-like pore clusters on each side of the segment; disc- like pores present from abdominal segment II to segments IV, V, or VI; the row of dor- sal multilocular pores on segment VII, ad- jacent to the intersegmental line between segments VII and VIII, with O—7(3) pores on each side of body. Antonina nakaharai has: 0O—4(1) multilocular pores on abdomi- nal segment IV within the area delimited by lateral margins of disc-like pore clusters on each side of the segment; disc-like pores present from abdominal segment II to seg- 900 ments IV, V, or VI; the row of dorsal mul- tilocular pores on segment VII, adjacent to the intersegmental line between segments VII and VIII, with O—3(0) pores on each side of body. Antonina socialis has: 0—7(2) multilocular pores on abdominal segment IV within the area delimited by the lateral margins of disc-like pore clusters on each side of the segment; disc-like pores present from abdominal segment II to VII or VII; the row of dorsal multilocular pores on seg- ment VII, adjacent to the intersegmental line between segments VII and VIII, usu- ally with I—12(6) pores on each side of body. Although this species previously was re- ported to be widespread (Ben-Dov 1994), the species concept presented here limits its distribution to China, Japan, and the U. S. (California). The latter record is based on a single adult female that was collected in San Jose, California, in 1900; it has not been collected in the U. S. since and is as- sumed to have disappeared in the interven- ing period. Specimens examined.—CHINA: Bam- boo gardens, Lingnan University, Canton, on Phyllostachys nigra, date ?, F A. Mc- Clure (1 ad. 2, 2 imm. on | sl.) (USNM). JAPAN: Kagoshima, on Phyllostachys sp., I-11-1923, B. M. Broadbent (1 ad. 2) (USNM); Kuro, Chiku, Hotel Chiku, on Bambusa sp.,? -?-1889, collector ? (2 ad. @ on | sl.) (USNM); Shinohana-machi, Ko- hoku-ku, Kanagawa-ken, Yokohama, on Sasa sp., X-19-1941, K. Sato (9 ad. @ on 2 sl.) (USNM); Near Tokyo, on Arundinar- ia fastuosa cultivar Hasimoto, III-18-1917, collector ? (1 ad.2?) (USNM); locality ?, on bamboo, XI-?-1909, E. M. Ehrhorn (1 ad. 2) (USNM); locality ?, on bamboo, ?-?- 1900, Craw (1 lectotype ad. 2, 2 lectotype ad. 2 on 2 sl.) (USNM); [Authors’ note: An additional slide containing | ad. 2 from E. E. Green’s collection may also be from the type series, but associated information only indicates, Japan, from bamboo (BMNH)]. UNITED STATES: CALIFOR- NIA: San Jose, Japanese Nursery, on bam- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON boo, II-?-1900, collector ? (1 ad. 9, 12 first instars on 2 sl.) (USNM). Antonina maai Williams and Miller, new species (Fig? 2) Antonina pretiosa: Hendricks and Kosztar- ab 1999:115 (misidentification, in part) Type material—Holotype adult female mounted alone on a slide labeled as fol- lows: left label *““Taiwan/Hsinkien, Taipeh/ Hsien/Ex Bambusa/Oct 9, 1951/T Maa/Fer- ris No M331” right label ““Antonina maai/ Williams & Miller/HOLOTYPE, UCD.” In addition there are 55 paratypes deposited in the following museums: BMNH, CDFA, FSCA, MNHN, TARI, UCD, USNM. Description.—Adult female holotype on microscope slide, elongate oval, 1.5 mm long (paratypes 1.0—2.7(1.6) mm), 0.8 mm wide (paratypes 0.7—1.8(1.1) mm); posteri- or apex of last 3 abdominal segments scler- otized, lateral margins of all abdominal seg- ments sclerotized (more mature female paratypes sclerotized completely); without convex lateral margins of abdominal seg- ments; sometimes margin of segment VIII convex in very mature adults. Antenna 2- segmented, basal segment small, with 3 se- tae, apical segment larger with 5 sensory setae and 2 or 3 thinner setae, antenna 62 w long (paratypes 52—70(61) w). Legs ab- sent. Labium profile 70 long (paratypes 48—75(66) ww), about 3 times shorter than clypeolabral shield. Ostioles present on ab- domen only. Circulus absent. Spiracles only with smaller size of trilocular pore in atri- um. Cerarii absent, but posterior abdominal setae slightly enlarged. Anal ring invagi- nated in anal tube, 142 p long (paratypes 135—168(150) w); ring diameter 88 p long (paratypes 85—100(92) w); anal tube inter- nally with ring of small trilocular pores near entrance. With 5 multilocular pores on each side of body in area delimited by posterior edges of each anterior and posterior spiracle (paratypes 0—10(4) pores). Dorsal surface with short setae, decreas- 901 VOLUME 104, NUMBER 4 i _ ule o' /A\o toh \{ \x \vo ie \ Antonina maai. Specimen from Taiwan, Taipeh, Hsien, on Bambusa sp. Bigi2: 902 ing in length and width anteriorly; longest seta on posterior apex 28 pw long (paratypes 25—31(28) w). Trilocular pores of 2 sizes, smaller size scattered over most of surface, increasingly abundant anteriorly, absent on posterior 2 or 3 abdominal segments; larger size restricted to marginal areas of anterior thorax and head. Discoidal pores of same distribution pattern as triloculars, less abun- dant. Multilocular pores present in cluster near entrance of opening to anal tube, mul- tilocular pores absent elsewhere on holo- type, variable on paratypes from relatively abundant near body margin of abdominal segments, thorax, and head to only | or 2 along any area of body margin; without pores on segment VII near intersegmental line between VII and VIII (paratypes O-— 1(0) pores). Oral-collar tubular ducts of 2 sizes, larger size restricted to marginal areas of abdomen, usually absent from segment VIII, smaller size in medial and mediolat- eral areas of abdomen and posterior thorax, absent from head (most paratypes with some small ducts along body margin of tho- rax and head). Ventral surface with setae similar to those on dorsum. Multilocular disc pores present in diffuse band along body margin, without pores in medial and mediolateral areas of segment IV; usually without mul- tilocular pores on segment VIII, rarely with 1 or 2; dermal surface near anal tube sur- rounded by cluster of multilocular pores. Disc-like pores restricted to mediolateral cluster on abdominal segments I and II; of- ten tubular in structure. Trilocular pores scattered over surface, most abundant an- teriorly. Oral-collar tubular ducts of 2 sizes, larger size present laterally from metatho- rax or abdominal segment I to abdominal apex, smaller size in medial and mediolat- eral area, sometimes present laterally on head and thorax. Vulvar area with | pair lateral apophyses, | pair lateral apodemes, and | medial apophysis. First-instar (based on numerous embryos from Taiwan and | first instar from Canton, China) are not in sufficient condition to al- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON low a complete description, but some di- agnostic characters are as follows. Antenna 6-segmented. Mesosternal apophysis ab- sent. Small round circulus present on inter- segmental line between segments III and IV. Ostioles restricted to posterior pair. Tri- locular pores of 3 sizes: larger size present around body margin, usually with 17 on each side of body (1 on lateral margin of each abdominal segment, 2 on lateral mar- gin of each thoracic segment, and 3 on lat- eral margin of head); medium-sized trilo- culars usually associated with ventral me- diolateral setae on each side of segment VII and with | or 2 near mouthparts, about 1/3 of specimens with 4 such pores forming longitudinal line associated with ventral mediolateral setae of abdominal segments IV—VII; small-sized triloculars forming marginal line around body, associated with larger marginal triloculars, also with 2 in each sclerotized area associated with spi- racular atrium, | associated with each leg, 1 associated with each ostiole, and some- times with | or 2 in ventral mediolateral areas of segments II and III. Discoidal pores associated with ventral sublateral setae, with | on each side of each body segment. Cerarii present on lateral margins of seg- ments V or VI to VIII, those on segment V and VI with | conical seta, those on seg- ments VII and VIII with 2 conical setae. Dorsal setae arranged in 4 longitudinal lines on each side of body (1 submedial, 1 me- diolateral, and 2 lateral) with | seta in each position on each body segment except with- out mediolateral setae on segments IL and III; ventral setae arranged in 3 longitudinal lines on each side of body (1 submedial, | mediolateral, and sublateral) with | seta in each position on each abdominal segment. Posterior abdominal segments with slight lobe laterally. Posterior seta 56—65(65) long. Notes.—Adult females of this species are most similar to the adult females of Anton- ina pretiosa in possessing abdominal seg- ments which are plate-like and sclerotized. Antonina maai differs by having disc-like VOLUME 104, NUMBER 4 pores confined to a single submedian group behind each posterior spiracle and having a few multilocular pores scattered along body margin including near opening to anal tube; A. pretiosa has disc-like pores forming a band from abdominal segment II posteriorly around entire abdomen and lacks pores along the body margin except near opening to anal tube. Based on the descriptions and key pre- sented by Yang and Kosztarab (1967) the first instar of this species also seems to be most similar to A. pretiosa. However, we discovered a discrepancy in their descrip- tion that merits correction. All specimens of A. pretiosa, including those examined by Yang and Kosztarab contain a very small, but definite circulus. Antonina maai and A. pretiosa share the following characters: 3 sizes of trilocular pores; no longitudinal lines of triloculars on dorsum; cerarii with conical setae; body margin of posterior ab- dominal segments lobed; a small circulus. Antonina maai differs by having: line of small-sized trilocular pores around body margin and trilocular pore associated with each ostiole. Antonina pretiosa has: no line of small-sized trilocular pores around body margin and no trilocular pore associated with ostioles. Etymology.—This species is named after Tsing-chao Maa, the collector, formerly of the Agricultural Research Institute, Taipeh, Taiwan, who also collected many scale in- sects in China and other areas in South East Asia. His help during G. E Ferris’ expedi- tion to China in 1948-1949 was much ap- preciated by Ferris (see Ferris 1950). Other specimens examined (not para- types)—CHINA: Foochow (= Fuzhou), on, ?-?-1976, M. S. Yang (8 ad. 2, on 3 sl.) (BMNH); Lingnan University, Canton, on Bambusa multiplex, UX-18-1948 (Ferris Collection # 97) (11 ad. @, | first instar, on Gr!) (WED): Antonina nakaharai Williams and Miller, new species (Fig. 3) Type material—Holotype adult female center specimen on slide with 2 adult fe- 903 male paratypes labeled as follows: left label “Glendale, MD/V-18-77 on/Phyllostachys sp./leaf sheath by S./Nakahara 6-77-3/” right label “‘Antonina nakaharai/Williams & Miller/HOLOTYPE/PARATYPE”’ the right label also gives a map of the position of the holotype. In addition there are 111 adult female paratypes, 7 immature para- types (excluding first instars), and 155 first instar paratypes. Description.—Adult female holotype on microscope slide, elongate oval, 1.7 mm long (paratypes 1.4—3.7(2.6) mm), 0.7 mm wide (paratypes 0.4—1.8(1.1) mm); poste- rior apex of last 3 abdominal segments sclerotized (more mature paratypes some- times with more sclerotization); lateral mar- gins of abdominal segments VI-VII con- vex, others straight sided. Antenna 2-seg- mented (paratypes with 2 or 3 segments), 65 yw long (paratypes 52—85(62) pw). Legs usually represented by inconspicuous scler- otized area, sometimes with associated seta. Labium 60 pw long (paratypes 70—100(80) uw), 2-3 times shorter than clypeolabral shield. Ostioles present on abdomen only. Circulus absent. Spiracles with 2 sizes of trilocular pores in atrium. Cerarii absent, but posterior abdominal setae enlarged, subconical. Anal ring invaginated in anal tube, 150 pw long (paratypes 120—170(139) 2); ring diameter 85 p long (paratypes 88— 100(95) jw); anal tube internally with ring of small multilocular pores and_ tubular ducts. With 14 & 19 multilocular pores on each side of body in area delimited by pos- terior edges of each anterior and posterior spiracle (paratypes 10—50(25) pores). Dorsal surface with short setae, decreas- ing in length and width anteriorly; longest seta on posterior apex 42 p long (paratypes 30—52(40) w). Trilocular pores of | size, ab- sent from posterior 2 or 3 segments, scat- tered over remainder of surface, least abun- dant posteriorly. Discoidal pores of same distribution pattern as triloculars, less abun- dant. Multilocular pores forming longitu- dinal band along body margin; without pores on segment VII near intersegmental PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 904 Antonina nakaharai. Specimen from U.S., Maryland, Glendale, on Phyllostachys sp. Fig. 3. VOLUME 104, NUMBER 4 line between VII and VIII (paratypes O- 3(0) pores). Oral-collar tubular ducts of | size, absent from segment VIII except in anal tube, abundant over remainder of sur- face. Ventral surface with setae similar to those on dorsum. Multilocular disc pores present in band along body margin, with | pore on abdominal segment IV within area delimited by lateral margin of disc-like pore clusters on each side of segment (paratypes with O—4(1) pores); with multilocular pores on segment VIII restricted to posterior half or third of distance between vulva and pos- terior apex of body; dermal surface sur- rounding opening of anal tube with cluster of multilocular pores. Disc-like pores pre- sent in mediolateral areas from metathorax to segment VI (paratypes from metathorax to segments IV, V, or VI). Trilocular pores of 2 or 3 sizes, smaller size in plate outside of spiracular atrium and near leg areas; larger size scattered around mouthparts and spiracles, medium size scattered over re- mainder of surface except absent or rare on posterior 3 or 4 segments, most abundant near body margin. Oral-collar tubular ducts present over surface. Vulvar area with | pair of lateral apophyses and | pair of weakly indicated posterior apophyses. Notes.—For a comparison of this species with A. crawi and A. see: vthe *“Notes”” section of the former species. Etymology.—We have named this spe- cies after Sueo (Steve) Nakahara who col- lected some fine specimens of this species in the U.S. He has been an important col- league of both authors for many years. Specimens examined.—CHINA: Hong Kong, on Sinobambusa tootsit, 1-30-41, W. B. Wood (2 ad. @ on 2 sl.) (USNM) (not part of paratype series); Hong Kong, Bo- tanical Gardens, on /xora stricta (?), VIII- 1-1936, R. H. LePelley (1 ad. 2) (USNM); Koolongue, Fukien, on Phyllostachys aurea VII-26-1930, E A. McClure (2 ad.¢ on | slide) (USNM); locality ?, on Sinobambusa laeta, Gouldman (1 ad. 2) (USNM) (not part of paratype series); locality ?, on Bam- socialis 905 busa sp., IX-1-1936, E C. Chen (4 ad. 2 on 4 sl.) (BMNH). RUSSIA: Szocsi (= So- chi), on Bambusa sp., LX-5-1985 E Kozar (1 ad. 2) (BMNH). JAPAN: Kuro, Chiku, Hotel Chiku, on bamboo, 7?-?-1889 (2 ad. 2; 3 firstunstars “on 2 sli (USNM)2 Mt. Hicko, on bamboo, date and collector ? (1 ad. 2) (USNM); Sunza Village, on bam- boo, XI-6-1980, E. L. Paddock (5 ad. @, 1 imm. on 3 sl.) (CDFA); Tokyo on bamboo, date and collector. 2> (Grad. on isl.) (UCD); Yokohama, on bamboo “‘seeds,”’ I- ?-1908, (12 ad. 2, 7 first instars on 4 sl.) (USNM); Yokohama, on Arundinaria var- legata pygmaea, VII-11-1911, collector ? (1 ad. 2) (USNM); Imperial Plant Quarantine Station, Yokohama, on bamboo, date ?, S. I. Kuwana (1 ad. 2) (UCD); locality ?, on bamboo, XI-?-1909, E. M. Ehrhorn (1 ad. 2°) (USNM) (not part of paratype series); locality, host ?, TI-?-1912, collector ? (3 ad. 2 on 1 sl.) (UCD); locality ?, on Arundi- naria japonica, VII-14-1933, W. H. Wheeler (1 ad. 2) (USNM); locality ?, on bamboo stems and leaves, VIH-29-1934, W. J. .Ehinger@ sads 2 1 imim,; on. 2 :sl) (USNM); locality ?, on bamboo stems, VIUI-2-1934.2.VE, 1G. Taylor _.G: Wad. 2) (USNM); locality ?, on bamboo, I-3-1935, C. V. Scott (i ad. 2). (USNM); locality: ?; on bamboo, XI-11-1936, O. A. Hardy (2 ad. 2 on | sl) (USNM); locality ?, on bamboo, 1-29-1937, A. B. Wells (3 ad. 2 on 1 sl) (USNM); locality ?, on Bambusa sp., X-4- 1938, RR; EF; Wilbur i2sady -22con) i -’sl) (USNM); in quarantine at Seattle, WA, on bamboo, I-17-1946, Young and Smith (1 ad. 2) (USNM); in quarantine at Encinitas, San Diego Co., CA, on Sasa kurihensis chabomanba, 1-28-1991, K. Sims (5 ad. 2: ron Sas) (EG DEA)s sLATWAN:s corzlin- shan, 2,660 m. elevation, near Arisan, on pygmy bamboo, [X-29-1949, T. Maa (6 ad. 2 on 3 sl.) (UCD). UNITED STATES: HA- WAII: Honolulu, on bamboo, VIII-3-1959, S. Miyake (5 ad.@ on | sl.) (USNM); Hon- olulu, on bamboo, date and collector ? (1 ad. 2, 3 first instars on 2 sl.) (USNM); Ka- waihae, on bamboo, XII-29-1994, S. Na- 906 kahara (5 ad.2, 1 imm. on | sl.) (USNM) Oahu, on bamboo, IV-4-1961, J. W. Beards- ley (4 ad.2 on 2 sl.) (USNM). LOUISI- ANA: New Orleans, on bamboo, [X-29- 19249 Beek? Planks (6jades “Savon Iasi) (USNM). MARYLAND: Prince George’s Co., Glendale, on Phyllostachys sp., leaf sheaths, V-18-1977, S. Nakahara (8 ad. &, 2 imm. on 3 sl.) (BMNH, USNM). NEW JERSEY: Riverton, on Bambusa sp., VU- 15-1915, Ho By Weiss) (©) ade 22) mm: > 100s fisst anstars “ont7 sl) (USNM): SOUTH CAROLINA: Aiken, on bamboo, MMES=19 i AvEucthiss@ sad? onl si) (USNM). TEXAS: Houston, on bamboo, date ?, N. B. Zuber (5 ad. 2, 1 imm. on 1 sl.) (USNM); Houston, on bamboo, VIII-?- 1918, E. Teas (2 ad. @, 4 first instars on 2 sl.) (USNM). Antonina socialis Newstead, reinstated status (Figs. 4—5) This species was described by Newstead (1901) but was subsequently treated as a junior synonym of Antonina crawi by Wil- liams (1962) based on the illustration of Ferris (1953). Ben-Dov (1994) and Ben- Dov et al. (2001) concurred with Williams’ synonymy. Type material—Lectotype adult female right specimen on slide with 2 additional adult female paralectotypes labeled as fol- lows: left label “‘Broxbourne,/Herts; on Arun-/dinaria japonica/Ex Ed. Gard. ChronY/2631-99:/E-MiMrrvolt Mc p285*7; right label *“Antonina/socialis nsp/123 News/Cotype females/3 = lectotype/316” (BMNH). The lectotype was designated by Williams (1985). Description.—Adult female on micro- scope slide, elongate oval, 1.3—5.0(3.3) mm long, 0.7—2.2(1.7) mm wide; posterior apex of last 2 abdominal segments sclerotized (more mature paratypes sometimes with more sclerotization); lateral margins of ab- dominal segments VIH-VIII convex, others straight sided. Antenna 2- or 3-segmented, 75—105(103) ww long. Legs usually repre- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sented by inconspicuous sclerotized area or small dermal pocket, sometimes absent. La- bium 88—112(99) w long, between 2 and 3 times shorter than clypeolabral shield. Os- tioles present on abdomen only. Circulus absent. Spiracles with 2 sizes of trilocular pores in sclerotized area surrounding atri- um. Cerarii absent, but posterior abdominal setae enlarged, subconical. Anal ring invag- inated in anal tube, 127—200(167) pw long; ring diameter 85—132(120) w); anal tube in- ternally with ring multilocular pores and tu- bular ducts. With 14—96(50) multilocular pores on each side of body in area delimited by posterior edges of each anterior and pos- terior spiracle. Dorsal surface with short setae, decreas- ing in length and width anteriorly; longest seta On posterior apex 32—47(40) pw long. Trilocular pores of 1 size, absent from pos- terior 3 or 4 segments, absent along body margin except near spiracles, scattered over remainder of surface. Discoidal pores of same distribution pattern as triloculars, less abundant. Multilocular pores forming lon- gitudinal band along body margin; with |— 12(6) pores on segment VII near interseg- mental line between VII and VIII. Oral-col- lar tubular ducts of | variable size, absent or rare on segment VIII except in anal tube, abundant over remainder of surface. Ventral surface with setae similar to those on dorsum. Multilocular disc pores present in band along body margin, with O— 7(2) pores on abdominal segment IV within area delimited by lateral margin of disc-like pore clusters on each side of segment; with multilocular pores on segment VIII restrict- ed to posterior half or quarter of distance between vulva and posterior apex of body, occasionally with 1 to 3 such pores just posterior to vulva; dermal surface surround- ing opening of anal tube with cluster of multilocular pores and oral-collar tubular ducts. Disc-like pores present in mediolat- eral areas from metathorax to segments VII or VIII. Trilocular pores of 2 or 3 sizes, smaller size in plate outside of spiracular atrium; larger size scattered around mouth- 907 VOLUME 104, NUMBER 4 a, New Orleans, on bamboo. lan Antonina socialis. Specimen from U.S., Louis Fig. 4. 908 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Antonina socialis. Specimen from Bermuda, on bamboo. VOLUME 104, NUMBER 4 parts, medium size scattered over remainder of surface except absent or rare on posterior 3 or 4 segments, most abundant near body margin. Oral-collar tubular ducts present over surface. Vulvar area with | pair of lat- eral apophyses and | pair of weakly indi- cated posterior apophyses. Notes.—For a comparison of this species with A. crawi and A. nakaharai see the *“Notes” section of the former species. Original material of this species is not satisfactory for illustrating. Fig. 4 is reason- ably similar in appearance to the type se- ries. Fig. 5 shows a variant that has fewer multilocular pores. Specimens examined.—BERMUDA: lo- cality ?, on bamboo, date and collector ? (6 ad 2 on 4 sl.) (BMNH); locality ?, on Arun- dinaria sp., V-7-1955, F J. Simmonds (4 ad. 2 on 1 sl.) (USNM). CHINA: Hong Kong, on Bambusa sp., X-21-1921, A. S. Entchcock Gi - = extinct lineage nodes [_] = extant species | = evolution NORTH - diversification | | = colonization on Gramineae SOUTH Fig. 3. ing and well definite flange, and 1.60 times BL at most. Marginal papillae not present on ab- donmnalgscomentSs: ete os cee cleon sss 2 ase 4 4. Green and very convex aphids living on rosa- ceous herbs. Tergites of metathorax to ab.seg. VI form a continuous sclerotized plate, tergites of prothorax, mesothorax and ab.seg.VI and VIII free (Fig. 2A). Distal and external setae of h.tb. normally longer than 0.5 times h.tb. width at seta insertion point [On Acaena splendens. Tie- tra del Fuego, southern slopes of the Andes] Sion NEP oes EAS En CREE CS Cone oe er P. acaenae — Brown aphids living on grasses. Tergites of ab.seg.I to [V or V forming a continuous plate, while tergites of thoracic segments and ab.seg.VI to VIII free (Fig. 2B). Distal and ex- ternal setae of h.tb. normally shorter than 0.5 times h.tb. width at seta insertion point [On Alopecurus magellanicus and Hordeum como- sum. Tierra del Fuego, North of the Andes] P. fueguinus Pentamyzus tenuis has a body shape very different from the other species of the genus with no developed frontal tubercles, glan- dular formations, or pointed dorsal setae, but it has the five-segmented antennae, the clavate siphunculi, and lives on grasses which are reasons not to establish a new genus level taxon. The absence of winged viviparous fe- Hypothesis on the origin and diversification of Pentamyzus. males in the South American species could be related to the fact that they live in very windy areas. PHYLOGENY AND BIOGEOGRAPHY The presence and relative richness of Pentamyzus in the “‘far south” of South America (including the Falkland Islands, as they are situated on the continental plat- form) is evident. The presence of P. gra- minis in California shows a very enlarged discontinuous distribution of the genus. This known distribution could be unreal be- cause extended areas of South America are not well studied, although we have looked for aphids in Andean areas of several Ar- gentinean provinces and interesting moun- tain areas of Mexico are well surveyed (Re- maudiere, personal communication). In our opinion, there are three possibilities (Fig. 3) to explain the origin and diversification of Pentamyzus. (1) Pentamyzus appeared in South America as a member of a lineage of Ma- crosiphini which emigrated from North America and more recently one species (as far as we know) returned to North America, 926 to California. The primitive Pentamyzus may have been dioecious, as its ancestor. Later, it diversified into two monoecious sublineages, one rosaceous-feeding and an- other grass-feeding. The sublineage on Ro- saceae finally established itself on herba- ceous ones, because there are no other au- tochthonous rosaceans species in the far south (this is not surprising in monoecious aphids, for example Chaetosiphon fragae- folii (Cockerell 1901), Amphorophora gei (Borner 1939), or Acyrthosiphon boreale Hille Ris Lambers 1952 [Macrosiphini] liv- ing on Rosaceae herbaceous species), and P. acaenae belongs to this sublineage. The other known Pentamyzus species belong to the other sublinage; in this group the most different species are: (i) P. tenuis, which has a elongated body and is South Ameri- can, and (i1) P. graminis, which has several derivative characteristics such as small-cla- vate siphunculi, lack of complementary u.rs. setae, and lack of marginal papillae (this last characteristic is also present in other species of the genus) and belongs to the returned lineage. (2) The primitive dioecious Pentamyzus stock was North American, and host di- verging originated in North America. Later, two sublineages colonized South America via the Andes. Here we found two possi- bilities: (2a) The northern species on Ro- saceae disappeared, P. acaenae belongs to the rosacean sublineage established in the South, P. graminis belongs to the old north- ern grass sublineage, and P. falklandicus, P. fueguinus, and P. tenuis belong to the southern grasses sublineage; or (2b) all northern species disappeared and P. gra- minis returned from South America, such as in hypothesis 1. (3) Pentamyzus was a North American monoecious and grass-feeding genus. Later it colonized South America, and one sub- lineage (represented by P. acaenae) ac- quired Acaena as a host. As in hypothesis 2, there are two possibilities with regard to P. graminis: (i) it belongs to the non-dis- placed old-lineage (in Fig. 3), or (i) it is PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the last species of a lineage which returned from South America. The three hypotheses are concomitant with the present theory on the colonization of South America by the Aphidinae (the Holarctic origin of this subfamily and its subordinated taxa, such as Macrosiphini, is generally accepted, Heie 1996). South America was invaded in different waves by different lineages of this tribe (so Micro- parsus, Pentamyzus and siphunculi-reticu- lated aphids, such as Uroleucon, Nietona- friella, and Blanchardaphis, belong to dif- ferent generic groups and live on plants of different families, the first one on Fabaceae and last three on Asteraceae). The three hypotheses may be correct, but we prefer | or, to a lesser extent, 3, because they are the most parsimonious when ex- plaining the morphological similarities be- tween P. fueguinus and P. acaenae and the presence of two “special” grass-feeding species, P. tenuis (because of its body ap- pearance) and P. graminis (because its de- rivative characteristics and distribution). In either case, only molecular analysis could allow us to arrive at a conclusion as to which of these hypotheses is correct. ACKNOWLEDGMENTS We thank Roger Blackman and Paul Brown (The Natural History Museum, Lon- don) and Ole Heie (Holte, Denmark) for constructive criticism of the manuscript. We also thank to E. Méndez, E. Martinez and A. Dalmasso, Botanists of the IADIZA (Mendoza, Argentina) for identifying the plant species from Tierra del Fuego. LITERATURE CITED Brown, P. 1987. Key to the genus Pentamyzus Hille Ris Lambers (Homoptera, Aphididae), with a de- scription of a new species from the Falkland Is- lands. Systematic Entomology 12(1): 1—6. Eastop, V. E 1998. Why do aphids do that?, pp 37— 47. In Nieto Nafria, J. M. and A. FE G. Dixon, eds. Aphids in Natural and Managed Ecosystems. Universidad de Leon. Leon. Footit, R. G. and W. R. Richards 1993. The Insects and Arachnids of Canada. Part 22. The Genera of VOLUME 104, NUMBER 4 the Aphids of Canada (Homoptera: Aphidoidea and Phylloxeroidea). Agriculture Canada, Re- search Branch, publication num. 1885. Ottawa. 766 pp. Heie, O. E. 1992. The Aphidoidea (Hemiptera) of Fen- noscandia and Denmark. IV. Family Aphididae: Part | of tribe Macrosiphini of Subfamily Aphi- dinae. Fauna Entomologica Scandinavica 25: |— 188. . 1996. The evolutionary history of aphids and hypothesis on the coevolution of aphids and plants. Bolletino di Zoologia agraria e Bachicol- tura (Serie II) 29(2): 149-155. Hille Ris Lambers D. 1966. Notes on aphids, with descriptions of new genera and new species (Homoptera: Aphididae). Hilgardia 37(15): 569-623. . 1974. On American aphids, with descriptions of a new genus and some new species (Homop- tera: Aphididae). Tijdschrift voor Entomologie 117(4): 103-155. California 927 Nieto Nafria, J. M., M. A. Delfino, y M. P. Mier Du- rante 1994. La afidofauna de la Argentina, su con- ocimiento en 1992. Universidad de Le6n (Secre- tariado de Publicaciones). Le6n. 235 pp. Remaudiere, G. and M. Remaudiere 1997. Catalogue des Aphididae du Monde / Catalogue of the World’s Aphididae. Homoptera Aphidoidea. Insti- tut National de la Recherche Agronomique. Paris. 478 pp. Schouteden, H. 1904. Aphiden. Hamburger Magal- haensische Sammelreise 4: 3-6. Smith, C. EF and M. M. Cermeli 1979. An Annotated List of Aphididae (Homoptera) of the Caribbean Islands and South and Central America. North Carolina Agricultural Research Service Technical Bulletin 259; 1-131. von Dohlen, C. D. and N. A. Moran, 2000. Molecular data support a rapid radiation of aphids on the Cretaceous and multiple origins of host alterna- tion. Biological Journal of the Linnean Society 71: 689-717. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 928-940 DESCRIPTION OF FIVE NEW SPECIES OF TABANIDAE (DIPTERA) FROM COSTA RICA AND REVISED KEYS TO SPECIES FOR THE GENERA FIDENA WALKER, SCIONE WALKER, AND CHRYSOPS MEIGEN IN COSTA RICA JOHN E BURGER Department of Zoology, Spaulding Hall, University of New Hampshire, Durham, NH 03824 U.S.A. (e-mail: jfb @cisunix.unh.edu) Abstract.—Five new species of Tabanidae from Costa Rica in four genera are described and differentiated from related species in their respective genera occurring in Costa Rica: Fidena (Fidena) griseithorax Burger, new species, Scione albopilosus Burger, new species, Acanthocera (Polistimima) vespiformis Burger, new species, Chrysops alajue- lensis Burger, new species, and Chrysops costaricensis Burger, new species. The im- mature stages of Acanthocera (Polistimima) vespiformis also are described, the first rep- resentative of the genus Acanthocera Macquart for which the immature stages have been found. Revised keys to the species of Fidena Walker, Scione Walker, and Chrysops Meigen occurring in Costa Rica are presented. Key Words: Examination of the Tabanidae in the col- lection of the Instituto Nacional de Biodiv- ersidad (INBio), Santo Domingo de Heré- dia. Costa Rica, and selected specimens in the Florida State Collection of Arthropods (FSCA) and the Cornell University Insect Collection (CUIC) revealed five unde- scribed species from Costa Rica, including a striking wasp mimic in the subgenus Pol- istimima Fairchild, genus Acanthocera Macquart, the first representative of this ge- nus recorded from Central America. These species are described below, compared with related taxa, and revised keys to species of the genera Fidena Walker, Scione Walker, and Chrysops Meigen occurring in Costa Rica are adapted from Fairchild (1986). Fidena (Fidena) griseithorax Burger, new species (Figs. 1-2) Diagnosis.—A large grayish and black species with elongate, conical tomented Neotropical, keys, Tabanidae, Costa Rica, new species face, snow-white beard, gray dusted mesos- cutum, blackish and brown legs, uniformly yellowish brown infuscated wings and shin- ing black abdomen with white lateral hair tufts on tergites 2, 5 and 6. Female.—Head: Front narrow, height 8.75 times width at base, slightly diverging above, gray tomentose below, slightly dark- ened toward vertex, bearing dense black hairs, callus absent. Subcallus grayish brown tomentose, protuberant. Frontocly- peus conical, elongate, entirely light gray tomentose and bearing abundant black setae dorsally and laterally, setae especially dense anteriorly. Antenna with scape and pedicel grayish brown, flagellum dark brown. Max- illary palp with apical palpomere dark brown, long and slender, length 8.3 times greatest width, about %4 length of mouth- parts, black setose, except for bare area ba- sally on outer surface. Outer sheath of pro- boscis broken off and missing, stylets near- VOLUME 104, NUMBER 4 ly twice as long as height of head. Eye densely pale pilose. Beard snow-white. Thorax: Mesoscutum grayish brown me- dianly, gray laterally, overlain with hoary gray tomentum, bearing black and scattered white hairs. Notopleural lobe light gray to- mentose, bearing mostly long black hairs and scattered white hairs. Area above wing base with dense tuft of white and black hairs. Postalar lobe bearing long white and scattered black hairs. Scutellum light brown with some grayish tones medianly, bearing black hairs. Pleuron light brownish gray, bearing dense white hairs dorsally and pos- teriorly on the mesanepisternum, postero- dorsal area of the katepisternum, the mes- anepimeron and the anterior katatergite, and black hairs ventrally on the katepisternum, posteriorly on the Katatergite, and scattered black hairs intermixed among white hairs on the mesanepisternum below the wing base. Legs: Coxae grayish brown, fore coxa white haired on basal *%, black haired on apical %, mid and hind coxae black haired. Femora dark brownish black and wholly black haired. Fore tibia and tarsus brown, mostly black haired, with orange hairs ventrally on apical half of fore tibia and ventrally on all tarsomeres, mid tibia and tarsus dark brown, hind tibia and tarsus black, mid- and hind tibiae and tarsi wholly black haired. Wing: Pale yellowish brown tinted, costal cell strongly yellowish, vein R, without spur, cell r; closed and petiolate. Abdomen: Shining black and black pi- lose, except anterior portion of first segment dark brown, tergites and sternites 2, 5 and 6 with lateral patches of white hairs. Length: 19.2 mm, wing, 16 mm. Holotype.—Female, COSTA RICA: Es- tacion Esquinas, Peninsula de Osa, Prov- incia Puntarenas, 200 m, August 1993, J. Quesada (INBIOCRIOO1 157720). Deposit- ed in the Instituto Nacional de Biodiversi- dad (INBio), Santo Domingo, Herédia, Costa Rica. Etymology.—This species is named for the hoary grayish tomentum overlying the 929 gray and brownish color of the mesoscu- tum. Discussion.—Fidena griseithorax shares with Fidena flavipennis Kroéber in the Costa Rican fauna the narrow front, elongated, to- mented face, dark light) yellowish brown tinted wings with closed and petio- late cell r;, and coloration of the abdomen. legs, Fidena griseithorax is easily distinguished from F. flavipennis by the white beard, white hairs on the pleuron and the gray to- mentum and lateral white hairs on the me- soscutum. KEY TO SPECIES OF FIDENA OCCURING IN COSTA RICA 1. Legs prominently bicolored, femora black, tib- iae and tarsi yellow. Wing black or heavily in- fuscated basally to ends of cells br and bm .. 2 — Legs not bicolored, all black, brown or yellow- ish. Wing rarely heavily infuscated basally, usually evenly tinted. =e ee eae ee 3 2. Beard snow white. Sides of mesoscutum with a stripe of white hairs from before transverse suture to posterior margin. Abdomen shiny black, with small tufts of silver-white hairs in middle of tergites 2—5, and at sides of tergites 2, 5 and 6. Sternites 2—4 with white lateral hair tufts; 2 sometimes with white hind marginal band. Face partially denuded and shiny load ne Sane ae ws Ae ertomeroides (Lutz) — Beard black or dark brown. Mesoscutum with- out contrasting pale hairs. Abdomen shiny black or deep brown, with segments 4 or 5—7 clothed with pale straw yellow, rufous orange, brown or black hairs. Tergite 2 with a patch of white hairs at sides, and sternites 2—4 with small lateral white hair tufts. Rarely tergites 3 and 4 with small white hair tufts rhinophora (Bellardi) 3. Legs pale yellowish brown. Beard, pleuron and abdominal sternites pale yellowish pilose, con- trasting with dark dorsal surfaces ......... NN EERE Sse wes eae ys RN auribarba (Enderlein) = ess darkabrownito blacks 3 2% #2 8).s.94 2% Ss 4 4. Abdomen largely black pilose ........... 5 Abdomen extensively golden pilose ....... i 5. Beard snow-white. Pleuron mostly snow-white haired — Beard and pleuron dark brown to black haired ee Ee RS cae Mn beh as hen es griseithorax, Nn. sp. SON OPEC ALC SOEs Cac Chis ee Crea a CR CeO RCOES 6 6. Large species, generally over 18 mm in wing length. Front narrow, more than 4.5 times as high as wide, below. Antenna brownish _ black. convergent Face conically produced, 930 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Adult females of Fidena griseithorax and Scione albopilosus. 1, Fidena griseithorax, habitus. 2 F. griseithorax, lateral view of head. 3, Scione albopilosus, habitus. 4, S. albopilosus, lateral view of head. VOLUME 104, NUMBER 4 brown, wholly grayish tomentose. Wing uni- formly deep yellowish brown tinted. Abdomen dark brown to black, shiny, sparsely black pi- lose, with tufts of white hairs at sides of ter- Ene 225 oP clael! 6)" “5 Gia So oles flavipennis Krober — Small species, wing length generally less than 12 mm. Front broader, less than 3 times as high as wide, parallel-sided. Antenna bright yellow. Face less produced, yellow, with extensive bare patches laterally. Wing smoky hyaline, the costal cell blackish and end of cell br darkened. Abdomen black, the second segment sparsely white haired above and below, forming a more or less distinct hind marginal band widened at sides above. Tergites 5—7 sparsely white haired and often with median white triangles on ter- PIE Sa Hay eens is ah eats nat iy eee e® ans schildi (Hine) 7. Abdomen with pale yellow integument and all segments beyond black pilose first segment paleveoldenspilose=a =. | a ee bicolor Kréber — Abdomen with at least some segments beyond first not entirely golden pilose 8. Face relatively short, shorter than front and subcallus, largely shiny. Antenna short and un- usually broad, basal flagellomeres distinctly wider than long. Thorax black, generally with prominent white hair tufts on each side of the scutellum, rarely reduced to a few white hairs mixed with black. Abdomen largely bright golden pilose, but with first tergite black pi- lose, second with golden hairs forming a large median triangle and lateral patches, third with only dorsolateral anterior black patches, re- mainder wholly golden pilose. Wing yellowish tinted, the basal cells and costal cell darker; cell r; coarctate, rarely closed J EFAS Geel CMe ea en Ee tata eee ioe trapidoi Fairchild — Face relatively long, as long as or longer than front and subcallus, gray above, shiny laterally. Antenna long and slender, basal flagellomeres at least as long as wide. Thorax without white hair tufts beside scutellum. Ab- domen bright golden pilose from third to last segments only, but with a small patch of yel- low hairs on posterior lateral borders of second tergite. Wing evenly brownish yellow tinted, cell r; closed and petiolate ... . howardi Fairchild tomentose Scione albopilosus Burger, new species (Figs. 3-4) Diagnosis.—A small, dark blackish brown species with distinct dorsolateral me- soscutal stripes, snow-white haired beard, pleuron and postalar tufts, unicolorous grayish wings and yellowish brown and pale pilose sternum. Female.—Head: Front relatively broad, 93] twice as high as width at base, slightly di- verging below, black brown and black pi- lose, except lighter brown near eye mar- gins. Subcallus and frontoclypeus concolo- rous with front. Face relatively short, densely black pilose. Antennal scape and pedicel dark brown, scape bearing long black setae, basal four flagellomeres brown, apical four flagellomeres darker brown. Maxillary palpus dark brown, apical pal- pomere relatively broad and strongly flat- tened, strongly narrowed in apical half and sharply pointed, length about three times greatest width. Proboscis relatively long and slender, about 1.5 times height of head. Beard snow-white haired. Eye densely light brown pilose. Thorax: Mesoscutum and scutellum shining dark brown and black pilose except for large tufts of snow-white hairs above wing base and on postalar lobe. Mesoscu- tum bearing two distinct yellowish gray dorsolateral longitudinal stripes that fade out in posterior third, and a narrower gray median longitudinal stripe extending about half way to scutellum. Pleuron light gray tomentose and white pilose except for scat- tered black hairs intermixed on mesanepis- ternum. Legs: Fore coxa grayish brown, white pilose basally, black pilose apically. Fore legs dark brown, mid and hind legs black. Wing: Wholly grayish hyaline. Abdomen: Tergites 1—2 brown, remain- ing tergites darker brown to black-brown, tergites 1—5 with a middorsal row of yel- lowish white haired spots, and pale haired lateral margins, otherwise black pilose. Sternites pale yellowish brown, white pi- lose. Length: 11.0 mm, wing, 10.7 mm. Holotype.—Female, COSTA RICA: Provincia Carta, M N Guayabo, Turrialba, 1,100 m, 21 June 1994, coll. J. E Corrales (INBIOCRIO01973469). Deposited in the Instituto Nacional de Biodiversidad (INBio) collection, Santo Domingo, Herédia, Costa Rica. Etymology.—This species is named for the snow-white haired beard, postalar hair 932 tufts and pleuron strongly contrasting with the dark brown body. Discussion.—This species is closest to Scione costaricana Szilady, but differs in having much longer and more slender api- cal palpomere, the snow-white hairs of the head and thorax, dark brown and black haired scutellum, grayish hyaline wing, less distinct middorsal yellowish spots on the abdominal tergites, and the entirely pale yellowish brown and pale pilose sternum. KEY TO THE SPECIES OF SCIONE OCCURING IN COSTA RICA 1. Wing particolored, yellowish and gray, cross- veins with obscure dark clouds. Thorax striped with yellowish gray markings. Abdomen yel- lowish brown to brown, often with some darker color posteriorly, with a middorsal row of yel- low to white pilose triangles or spots on the posterior margin of tergites 1—6 (sometimes obscure to absent in abraded specimens) ... . Sys peop hae ees Stay eyo maculipennis (Schiner) — Wing hyaline or slightly tinted, not partico- lored and without clouds on the crossveins . . i) Nw . Mesoscutum with distinct grayish or yellowish brown dorsolateral stripes extending at least half way to scutellum. Abdomen with contrast- aver joeUKS WEN UIT agape e sue Sonos aoo bec 3 without strongly contrasting stripes. Abdomen without contrasting pale hair — Mesoscutum TURES erate en STM oe eee seis So ome fonek eager 4 3. Beard, pleural hairs and postalar hair tufts snow-white. Sternum pale yellowish brown, Witte spIlLOSe) pseceep sy swey eaictel tse albopilosus n. sp. — Beard pale yellow to golden and black haired to mostly black haired. Postalar hair tufts gold- en. Sternum with the anterior two segments brown, remainder black, black pilose except for white haired hind margins on sternites 2—5 costaricana Szilady 4. Beard, pleuron and sternum white pilose. Api- cal palpomere extremely short and blunt, only half as long as flagellum . . aurilans (Wiedemann) — Beard, pleuron and sternum black pilose. Api- cal palpomere sickle shaped, subequal in length to flagellum ...... fulvosericea (Krober) Acanthocera (Polistimima) vespiformis Burger, new species (Figs. 5-11) Diagnosis.—A large, wasplike blackish brown and black species with greatly elon- gated scape and pedicel, basal flagellomere PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with thumblike dorsal spine, greatly inflated maxillary palpi, wing with anterior third darkened, and abdomen strongly constricted basally. Female.—Head: Front parallel above, slightly divergent below near subcallus, in- dex 1.6, velvety black medianly, blackish brown laterally, bearing dense, very short bristly black hairs, except on callus and near eye margins, callus brown, triangular, rather poorly defined, widely separated from eye margins, with a narrow dorsal ex- tension half way to vertex. Vertex distinctly sunken below level of ocular margin, ocular margin with short orange and black hairs. Subcallus brownish black, with long yel- lowish and dark hairs dorsolaterally below junction with front. Frontoclypeus shining dark brown, strongly swollen. Gena black- ish gray tomentose. Antenna extremely long, equal in length to mesoscutum and scutellum combined, scape and pedicel shining dark brown, flagellum reddish brown, scape cylindrical, greatly elongate, twice length of pedicel and equal in length to flagellum, scape and pedicel together 1.5 times longer than flagellum, basal flagello- mere and apical flagellomeres subequal in length, basal flagellomere with thumblike dorsal spine extending %4 length of basal fla- gellomere, apical flagellomeres not hirsute, gradually tapered apically, terminal flagel- lomere sharply pointed. Maxillary palpus subshining dark brown, apical palpomere greatly inflated basally, strongly narrowed distally and sharply pointed apically, bear- ing short dark hairs. Labellum of proboscis partially sclerotized. Beard sparsely black haired. Thorax: Mesoscutum black medianly, blackish brown laterally, bearing very short recumbent black hairs and pale yellowish white hairs posterolaterally anterior to scu- tellum. Scutellum black medianly, brown laterally, with tuft of white hairs on lateral margins. Pleuron blackish above, with dark grayish bloom below, bearing short black and yellowish hairs and small tuft of white hairs on posterodorsal surface of mesane- VOLUME 104, NUMBER 4 933 Figs. 5-11. Acanthocera (Polistimima) vespiformis. 5, Adult female, habitus. 6, Adult female, lateral view of head. 7, Mature larva, lateral view. 8, Mature larva, lateral view of anal segment. 9, Pupa, lateral view. 10, Pupal aster, lateral view. 11, Frontal plate of pupa. 934 pisternum anterior to wing base. Legs: Coxae and femora blackish, except femora narrowly brown at extreme apices, coxae and fore and mid femora bearing golden hairs, hind femur black haired, fore and mid tibiae blackish brown and golden haired, hind tibia black and black haired, concolo- rous with hind femur, tarsi brown, golden haired. Wing: Anterior third strongly yel- lowish brown tinted, tinting fading out to- ward wing apex beyond fork of veins R, and R,;, and faint yellowish tint along vein Cu in cell bm, vein R, without spur. Abdomen: Segments 1—2 strongly com- pressed laterally, abdomen strongly arched dorsally, tergite 1 brown anteriorly, black laterally and posteriorly, tergite 2 black, with 2 brown patches sublaterally in middle of segment, tergites 1—2 with patch of sil- very hairs posterolaterally, remaining ter- gites black and black haired, sternite 2 ir- regularly brown and black, with complete transverse band of silvery hairs on posterior margin, remaining sternites black and black haired. Length: 19.2 mm, wing, 16 mm. Mature larva.—Length 26—28 mm, body creamy white to yellowish white. Head cap- sule 5.2 mm long, 1.2 mm wide. Respira- tory siphon 0.8 mm long, width and length subequal. Anal segment length and width subequal. Body fully striated, cuticular stri- ations most conspicuous on thoracic seg- ments, very indistinct on abdominal seg- ments, especially posteriorly, striations very narrowly spaced. Pseudopodia, if present, inapparent. Anterior third of thoracic seg- ments ornamented with extremely small, dense microspines. Anterior margins of ab- dominal I-VI with row of elongate, curved microspines entirely encircling each seg- ment, posterior margin of segment VII and anterior margin of anal segment broadly en- circled by dense, minute microspines. Pupa.—Length 18.4—20.8 mm. Color uniformly light brown. Frontal plate with numerous transverse dark cuticular stria- tions anteriorly and laterally. Antennal ridg- es very narrowly divided medianly into 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON strongly projecting, shelflike extensions an- teriorly, raised about 0.4 mm above surface of frontal plate, strongly rugose. Anterior and posterior orbital setae very short, borne on very small, irregular tubercles. Frontal tubercle absent. Callus tubercles very low and irregular, anterior and posterior margins sinuous, each bearing a single short seta. Antennal sheaths large and conspicuous, extended well beyond cephalothoracic su- ture, length about 1.4 mm. Thoracic spira- cles strongly recurved, earlike. Anterior mesonotal tubercle bearing 2 setae each. Abdominal spinous fringes biseriate dorsal- ly and laterally, uniseriate ventrally, ante- rior row very short, posterior row mostly short and stout, with scattered longer, more slender spines intermixed, and mostly long, slender spines mid ventrally. Dorsal preanal combs reduced to 3—4 spines borne on very large sublateral lobes and a single short spine on a median lobe. Ventral preanal comb with 5 short, stout spines on each side (2). Terminal aster with dorsal arms re- duced to minute spines, lateral and ventral arms very stout, lateral arms longest, pro- jecting posteriorly, ventral arms about half length of lateral arms, projecting postero- ventrally. Holotype.—Female, COSTA RICA: Puntarenas Province, Peninsula de Osa, Parque Nacional Corcavado, near Cerro Rinc6n, 700 m., coll. Alejandro Azofeifa Azofeifa, Jan. 1997. Paratypes.—3 2, same data as holotype, except coll. Jan. 1998. All specimens reared from larvae and pupae. Holotype 2 and 2 ? paratypes to be deposited in the Instituto Nacional de Biodiversidad (INBio), Santo Domingo, Herédia, Costa Rica; | 2 para- type in the University of New Hampshire Insect Collection (UNHC). Etymology.—This remarkable species 1s named for its striking wasplike appearance. Discussion.—The paratype females are very similar to the holotype, except 15.2 mm—15.4 mm. This striking wasp mimic was originally considered to belong to a new subgenus of Acanthocera, but the elon- VOLUME 104, NUMBER 4 Figs. 12-13. istimima) vespiformis. 12, Sap run from Cedro Marta Larval habitat of Acanthocera (Pol- tree (Calophyllum brasiliense) showing central cavity. 13, Pupal exuvia of A. vespiformis protruding from a sap run. gate scape combined with the relatively short flagellum with apical flagellomeres not greatly lengthed suggests that this spe- cies should be placed in the subgenus Pol- istimima, erected by Fairchild (1969) for Acanthocera (Polistimima) polistiformis Fairchild, 1961, from Amapa, Brasil. Acan- thocera (Polistimima) vespiformis differs from that species in color of the body, legs and wings, the much longer antennal scape and the basal flagellomere with a long thumblike dorsal spine. Unfortunately, the species from Brasil is known only from a single male. Thus, it is uncertain whether some of the differences of A. vespiformis may be due to sexual differences. Larvae and pupae of A. vespiformis were found inhabiting tunnels in sap runs (Figs. 12—13) associated with beetle attacks to Cedro Maria trees (Calophyllum brasiliense Cambess) (Clusiaceae). When the larvae are in the sap tunnels, they transform the 935 sap into a sticky, foul-smelling mass that attracts flies and wasps that become trapped in the sticky sap. The larvae prey on the trapped insects. The larvae always were found only in sap tunnels less than two me- ters from the ground. Prior to adult emer- gence, the pupa moves partially out of the tunnel (Fig. 13). Emergence occurs during the day. Developing larvae have been ob- served throughout the year (Zumbado, per- sonal communication). Length of the life cycle is unknown. In addition to the adults listed above, one deformed, shrivelled adult female did not harden properly, and is not included in the series of paratypes. This is the first species of the genus Acanthocera for which and pupae have been found and reared, so it is not known if other species of Acanthocera live in similar hab- itats. This also is the first record for the ge- nus Acanthocera in Central America., hith- erto known only in South America, as far north as Colombia. larvae Chrysops alajuelensis Burger, new species (Figs. 14-16) Diagnosis.—A yellowish gray and brown species with strongly inflated antennal scapes, blackish gray thorax, reddish to yel- lowish brown scutellum, wing crossband with irregular outer margin, abdominal ter- gites with pale yellowish gray median tri- angles and sublateral spots, leaving a darker bivittate submedian marking and sternites uniformly yellowish brown or faintly spot- ted. Female.—Head: Front yellowish gray tomentose with a dark subshiny area sur- rounding ocelli. Frontal callus crescent shaped, narrowed laterally, reddish brown with darker markings laterally on the upper margin. Frontoclypeus reddish brown with narrow yellowish gray tomentose stripe. Antenna with scape, pedicel and base of fla- gellum brown, remainder of flagellum black, scape and pedicel distinctly inflated. Maxillary palpus light brown. Thorax: Mesoscutum blackish gray with 936 Rap a ae Figs. 14-18. 18, C. costaricensis, abdomen, dorsal view. broad, indistinct submedian grayish longi- tudinal stripes and lateral margins. Scutel- lum reddish brown. Pleuron yellowish gray tomentose and yellowish white pilose. Legs: Femora reddish brown except ex- treme apices darker, tibiae and tarsi light brown. Wing: Dark pattern distinct, cells br and bm ¥; and % infuscated basally, respec- tively, crossband broad, outer margin irreg- ular, with an extension toward fork of veins PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Adult females of Chrysops alajuelensis and Chrysops costaricensis. 14, C. alajuelensis, lateral view of head. 15, C alajuelensis, wing. 16, C. alajuelensis, abdomen, dorsal view. 17, C. costaricensis, wing. R, and R; along vein R,,; and in cell m,, cell cua, with apical half hyaline, apical spot drop-shaped. Abdomen: Tergite 1 with a dark subme- dian marking adjacent to scutellum and pale sublateral spots, tergites 2—6 with pale me- dian triangles and sublateral spots leaving a darker bivittate submedian area. Sternites uniformly yellowish brown and pale whit- ish pilose. VOLUME 104, NUMBER 4 Length: 9.2 mm, wing, 7.6 mm. Male.—Head: Eyes holoptic, narrowly contiguous at mid-height, upper eye facets distinctly larger than lower facets. Ocellar tubercle subshiny blackish, with long black- ish hairs. Frontoclypeus shiny light brown with very narrow yellowish gray tomentose midstripe. Antennal scape and pedicel blackish brown, flagellum black except at extreme base. Maxillary palpus dark brown, apical palpomere short and slender, blunt apically. Thorax: Mesoscutum and scutellum sub- shiny blackish, pale stripes on mesoscutum evanescent. Pleuron concolorous with me- soscutum, black pilose. Legs: Similar to fe- male. Wing: Pattern similar to female ex- cept cells br and bm %4 infuscated basally. Abdomen: Black, tergites 2—5 with broad yellow sublateral spots and lateral margins of tergites 1—4 yellowish to yellowish brown. Sternites 1—5 yellowish, with broad median longitudinal black stripe and_nar- rower stripes sublaterally, sternites 6—7 black. Length: 7.2 mm, wing, 6.4 mm. Holotype.—Female, COSTA RICA: Guanacaste Province, 12 mi. SW of Canas, I{-27—1964, 25 ft., H. E. Evans (FSCA). Paratypes.—COSTA RICA: San José Province: 3 6, | 2, San Antonio-Desam- parados, 16-18 June 1964, C. O. Berg (2 Ge hea CUwIe ag ESCA); Alajuela Prov- ince: 1 ¢, San Josecito de Alajuela, 2 km. from road to Atenas, near. Rio Itiquis, 6 July 1969, Karl Valley, taken sweeping (CUIC); Guanacaste Province: 1 2, same data as holotype 2 (FSCA); 292, Estacion Pitilla, 9 km. S. Santa Cecilia, GNP Bio- diversity Survey, 85°25’40"W, 10°59'26'N, Jan. 1989 (INBIOCRI001047591) and Feb. 1989 (INBIOCRI000105975) (INBio). Etymology.—Named for Alajuela Prov- ince in Costa Rica. Discussion.—The five paratype females are 8.4—8.8 mm long. All the males are 7.2 mm long. The female paratypes are very similar to the holotype, except the posterior extensions of the crossband do not reach the 937 hind margin of the wing. Two of the male paratypes have very faint, small dark gray- ish median posterior triangles on tergites 2— 4. Chrysops alajuelensis is closely related to Chrysops scalaratus’ Bellardi, known from Mexico to western Panama, and was initially confused with that species. Females of C. alajuelensis differ from those of C. scalaratus, as follows: frontal callus rela- tively slender and reddish brown, antennal scapes distinctly stouter, frontoclypeus with distinct midfacial tomentose stripe, mesos- cutum yellowish gray with very indistinct paler stripes, scutellum reddish brown, ground color of abdominal tergites yellow- ish brown, sublateral spots relatively small, sternites unicolorous yellow brown or with very faint longitudinal stripes. C. scalaratus has the frontal callus broader and mostly black, antennal scape only slightly inflated, frontoclypeus bare and shiny, rarely with a short tomentose spur beneath antennae, me- soscutum shiny black with distinct gray longitudinal stripes, scutellum black, ab- dominal tergites with ground color blackish and sublateral pale spots very large, ster- num with distinct median black stripe. Males of C. alajuelensis differ from those of C. scalaratus as follows: antenna shorter and scape much stouter, mesoscutum with paler stripes evanescent or absent, abdom- inal tergites with broad black midstripe and large sublateral yellowish brown spots, ster- nites bearing broad median and narrow sub- lateral black stripes. Chrysops scalaratus males have the antenna distinctly longer and scape less inflated, mesoscutum with distinct longitudinal gray stripes, abdominal midstripe narrow, with distinct median grayish triangles on tergites 2—5 and narrow sublateral dark stripes on tergites 2—6, ster- nites without distinct sublateral black stripes. Chrysops costaricensis Burger, new species (Figs. 17—18) Diagnosis.—A yellow and black species with blackish thorax, conspicuously yellow 938 spotted pleuron, wholly black femora, nar- row apicocostal infuscation on the wing, and black abdomen with distinct rows of median and dorsolateral triangles. Female.—Head: Front yellowish gray tomentose with a large shiny black area sur- rounding ocelli. Frontal callus crescent shaped, broadest in middle, strongly nar- rowed near eye margins, brown with black upper margin. Frontoclypeus' shining brown, gena strongly inflated. Antennal scape light brown, relatively slender, pedi- cel darker brown to blackish at apex, fla- gellum black. Maxillary palpus dark red- dish brown. Thorax: Mesoscutum and scutellum shining black, mesoscutum with two broad yellowish submedian longitudinal stripes anteriorly that fade out about half way to the scutellum, and lateral margins narrowly yellowish. Pleuron shining black, proepis- ternum, proepimeron, upper area of mesa- nepisternum, mesanepimeron and meron strongly yellow spotted and pale haired. Legs: Femora wholly black, fore and hind tibiae black, mid tibia brown, all basal tar- someres brown, apical tarsomeres brown basally, black apically. Wing: Wing with distinct dark brown pattern and broad cross- band, cells br and bm 7% and Y; infuscated basally, respectively, apices also infuscated, crossband with slightly irregular outer mar- gin, divided posteriorly by hyaline area in cell cua, that does not encroach on cell m,, apicocostal infuscation narrow, filling half of cell r, and shghtly widened in cell ry, extending to wing apex. Abdomen: Tergite | with narrow, irreg- ular posterior and lateral margins yellow, tergite 2 yellow with pair of large, black submedian markings narrowly joined to smaller dorsolateral black spots, producing paired geminate markings, middorsal yel- low triangle large, extending to anterior margin, tergites 3—5 black with middorsal row and dorsolateral rows of distinct yellow triangles arising from hind margins, tergites 6—7 black. Sternites broadly black medianly and laterally, leaving narrow ventrolateral PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON yellowish brown patches, and hind margins narrowly yellowish. Length: 7.3 mm, wing, 6.6 mm. Male.—(Headless). Thorax and abdomen similar to female, but more extensively black. Submedian pale stripes on mesos- cutum evanescent. Pleuron almost entirely black except for small gray tomentose spots on proepisternum, mesanepisternum and meron. Wing with cells br and bm entirely infuscated except for small subapical hya- line spots, and infuscation broadly invading cell a,;. Abdominal tergite 2 more exten- sively black, median yellow triangle small, equilateral, not extending to anterior mar- gin. Holotype.—Female, COSTA RICA: Provincia Herédia, Estacion Magsasay, Parque Nacional Braulio Carrillo, 200 m., January, 1991, coll. A. Fernandez (INBI- OCRI000627622) (INBio). Paratypes—COSTA RICA: 1 2, Herédia Province: Estaci6n El Ceibo, Parque Nacion- al Braulio Carrillo, 400—600 m, March 1990, coll. C. Chaves (INBIOCRIO00167501) (IN- Bio); 2, same data, except January 1990 (IN- BIOCRIO00201892) (INBio). Puntarenas Province: 2, Rancho Quemado, Peninsula de Osa, 200 m., November 1991, coll. E Que- sada (INBICRI000462501) (UNHC); &, same data, except December 1992 (INBIOC- RIO00905966) (INBio); d, same data except December 1991 (INBIOCRI000342968) (IN- Bio). San José Province: 2, EF Cementario de la Maquina, Parque Nacional Chirripo, 2,100—2,500 m., 2 March 1993, coll. M. Zumbado (INBIOCRIO01305778) (INBio). Discussion.—The paratype females are 6.6—7.3 mm long. The black markings on tergite 2 are narrowly or broadly confluent, producing paired geminate dark markings, and the ventrolateral yellowish-brown markings on sternites 3—5 are absent. Chrysops costaricensis appears to be closest to Chrysops chiriquensis Fairchild, 1939, known from Mexico (Chiapas) to Co- lombia, but differs in the following char- acteristics: females are somewhat smaller than C. chiriquensis females; frontal callus VOLUME 104, NUMBER 4 reddish brown with black upper margin, frontoclypeus bare and shiny, pleuron strongly yellow spotted, femora black, wing with apical infuscation narrow, joined to crossband, median yellow triangle on ter- gite 2 open to anterior margin. Females of C. chiriquensis have the frontal callus largely black, frontoclypeus with tomentose midstripe, mesoscutum without yellowish lateral stripe, pleuron dull grayish spotted, femora brownish with black apices, apical infuscation drop shaped, separated from crossband, median triangle on tergite 2 not open to anterior margin. The male of C. costaricensis has femora black, wing with apical infuscation joined to crossband, and tergites 2—5 with median and sublateral yel- low triangles arising from posterior margin. Males of C. chiriquensis have reddish brown femora, apical infuscation separated from crossband and tergites 2—4 with yel- low sublateral stripes. KEY TO THE SPECIES OF CHRYSOPS OCCURRING IN COSTA RICA An asterisk (*) indicates not yet recorded from Costa Rica, but possibly may occur there. 1. Frontoclypeus with a median tomentose stripe 3 — Frontoclypeus entirely bare and shiny (rarely Wathva:shorttomentosesspun)) 4 eae ee 3 i) Mesoscutum and scutellum shiny black. Apical infuscation of wing separated from crossband. Abdominal tergites black with pale yellowish markings, tergite 1 entirely black except for ex- treme apex, tergites 3—5 with median and sub- lateral yellowish triangles arising from hind margins. Sternum black with yellowish gray sublateral stripes — Mesoscutum grayish brown tomentose. Scutel- lum yellowish brown. Apical infuscation of wing joined to crossband. Abdominal tergites mostly yellowish brown with yellowish brown median triangles and rounded sublateral spots not arising from hind margins. Sternum uni- formly yellowish brown to faintly striped me- dianly and sublaterally 3. Discal cell of wing largely hyaline, at least a central spot or hyaline streak — Discal cell of wing wholly infuscated 4. Abdomen largely yellow, with two dorsolateral chiriquensis Fairchild alajuelensis, n. sp. o dark stripes from the second to third tergites, generally divided into two stripes on third and succeeding segments. Legs and frontal and fa- cial callosities largely yellow. Thorax yellow to brown, with yellow tomentose stripes. Wing pattern yellowish brown Abdomen with prominent gray tomentose transverse bands on posterior margins of all tergites. Wing pattern of two dark bands cross- ing the wing at the ends of cells br and bm and beyond discal cell. Apical spot a faint, narrow infuscation along fore border. Apices of femora and at least bases of tibiae yellow ........ . Apical spot of wing a slender band or slightly widened, not extending beyond apex and en- croaching into cell r, only at extreme apex Apical spot of wing extensive, filling most of wing apex and leaving but a narrow and irreg- ular hyaline crescent or series of spots between it-and.crossbandis, saree ea nec. oe Cell cua, entirely infuscated. Cell m, usually with a small hyaline spot. Crossband broad and intensely black. Thorax black, without dorsal stripes or lateral spots. Abdomen black, sides of the second tergite with small, dull yellow triangles, and with small, dull yellow median triangles on terpites: 2—Ae 7 ae eee cee Cell cua, always with at least the apex hyaline, often over half hyaline. Cell m, with or without a hyaline spot Mesoscutum and pleuron blackish, without stripes or spots. Wing with hyaline area in cell cua, clearly extending into cell m,, but not crossing it. Abdomen black with small dull yel- low triangles laterally on tergite 2 and median yellow triangles on tergites 2—5 Mesoscutum strongly striped or at least weakly striped anteriorly, pleuron striped or spotted .. . Abdomen with second tergite largely yellow except for two large, submedian black mark- ings and small dorsolateral spots, yellow me- dian triangle reaching anterior margin. Tergites 3-5 black with prominent median and dorso- lateral yellow triangles arising from posterior margin. Sternites 1—4 with large median black PatCheSpasee yrectect reas eee Abdomen without prominent yellow dorsolat- eralptmianglesvonetersites*3=5° Hage. clea Abdomen with a dorsolateral series of pale dashes, forming a pair of interrupted stripes, as well as a row of pale median triangles on ter- gites 2—5. Sides of tergites 1—3 or 4 yellowish. Abdomen yellowish ventrally, with a broad, black median longitudinal stripe. Crossband of wing with outer border irregular, with exten- sion toward fork of veins R, and R; along vein costaricensis, N. 939 . variegatus (DeGeer) alleni Fairchild 6 leucospilus Wiedemann* . melaenus Hine sp. 9 940 R,.;, fading posteriorly and failing to reach hind margin in cell m, ..... scalaratus Bellardi — Abdomen without dorsolateral interrupted stripes or even ventral stripe. Outer margin of crossband straight or slightly concave, cross- band tfillmevapexcoficelliim zs See 2 ee 10 10. Second tergite black, broadly yellow at the sides, rarely with a small median yellow spot, generally without any median marks. Tergites 3—4 black, with equilateral yellow triangles whose apices may reach anterior borders, ter- gite 5 with a yellow hind border widened into a low triangle in middle. Wing with a narrow- er crossband, the hyaline area in posterior margin of crossband large, filling all but base of cell cua, and entirely crossing cell m, in the middle. Cell r, with a small hyaline streak aw DASE) Ae ese <5 Ail: ed oed A auroguttatus Krober — Second tergite black with yellow median hourglass-shaped mark extending longitudinal width of tergite, sides broadly yellow. Third to fifth tergites black with yellow hind bor- ders, widened into narrow median triangles which usually reach anterior border of seg- ments. Wing with a broad, dark crossband, without a hyaline fenestra in cell r;, and with hyaline area in the posterior margin of the crossband confined to cell cua, .-......-. 11 11. Apical spot black, extending as a clearly de- fined narrow costal stripe of even width to wing apex varians var. tardus Wiedemann — Apical spot dilute, poorly defined, spread as a brownish shade over much of wing apex Bur raid cana pein eee ste varians vargus Fairchild 12. Hind femur at least one-third black basally — Hind femur red or yellow, at most extreme apexublacks sys Qhmticsiey metric eaeri ese mere 14 13. Abdomen black, with the second segment an- teriorly narrowly, laterally broadly yellow, and with a broad yellow median triangle which rarely extends through black to join an- terior yellow band. Tergites 3—5 or 6 with nar- row yellow hind borders, widened into me- dian triangles of variable size on 3 and often 4. Pleuron usually with a spot of yellow pile, rarely lacking. Wing with crossband joined to a cloud on fork of veins R, and R; by a dark bridge above the vein, leaving a small hyaline spot between cloud and crossband on vein EE an eRe eats i Ge eOe ret rare calogaster Schiner — Abdomen wholly black, or rarely with pattern of calogaster faintly indicated in grayish to- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mentum. Pleuron always entirely black. Cloud on fork of veins R, and R; never joined by a complete bridge to crossband, rarely al- MOsStyomed by ASpUtS ete ena soror Krober 14. Cloud on fork of veins R, and Rs joined to crossband by a broad spur of dark color. Mid tibia black or blackish, contrasting with pale femora. Abdomen with second tergite narrow- ly white to yellow anteriorly, widened to full length of segment at sides and with a broad, small median yellow triangle. Third and fourth tergites rarely with faint vestiges of small, pale median triangles . . nexosus Fairchild — Cloud on fork of veins R, and R; unconnected with crossband. Mid tibia at most slightly brownish, not strongly contrasting with pale femora. Abdomen as above, but pale triangle on second tergite usually taller and narrower, occasionally joined to anterior yellow by a narrow, median yellow stripe. Third tergite usually with a dull yellowish median streak, sometimes lacking, and fourth tergite rarely with a minute paler streak .. mexicanus Kroéber ACKNOWLEDGMENTS I thank the curators of Diptera for the Cornell University Insect Collection and the Florida State Collection of Arthropods for the loan of specimens; Tess Feltes, Ports- mouth, New Hampshire, for the illustrations of the new species; and special thanks to Manuel Zumbado, Instituto Nacional de Biodiversidad (INBio), Santo Domingo, Costa Rica, for arranging visits to study the collection of Tabanidae at INBio, assistance in working with the collection, and for in- formation regarding the habits and habitat of the immature stages of Acanthocera (Polistimima) vespiformis. LITERATURE CITED Fairchild, G. B. 1961. Insecta Amapaensia.—Diptera: Tabanidae. Studia Entomologica 4: 438—448. ———. 1969. Notes on Neotropical Tabanidae. XII. Classification and distribution with keys to genera and subgenera. Arquivos de Zoologia, Sao Paulo 17: 199-255. 1986. The Tabanidae of Panama. Contribu- tions of the American Entomological Institute 223) 139" PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 941-947 A REVISION OF THE NEOTROPICAL BUTTERFLY GENUS SECO HALL AND HARVEY (LEPIDOPTERA: RIODINIDAE) JASON P. W. HALL AND DONALD J. HARVEY Department of Systematic Biology—Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0127, U.S.A. (e-mail: jpwhall@ hotmail. com) Abstract.—The South American riodinid genus Seco Hall and Harvey is revised. The taxonomy and biology of its species are discussed, locality data and range maps are given, and the adults and male and female genitalia (where known) are illustrated for all species. Key Words: A recent phylogenetic review of the Neo- tropical riodinid genera Charis Htibner [1819] and Calephelis Grote and Robinson 1869, showed that the heterogeneous Char- is was paraphyletic with respect to Cale- phelis, prompting the division of Charis into three monophyletic genera (Hall and Harvey 2002). One of these genera, Detri- tivora Hall and Harvey 2002 (containing the cleonus and gynaea groups), has been revised already by Hall and Harvey (2001) and Harvey and Hall (2002). The purpose of this paper is to revise the remaining ge- nus, Seco Hall and Harvey 2002, a small group of rare and very poorly known riod- inine butterflies that are uniquely restricted to xeric habitats around the periphery of South America. The adults and genital structures of all species (where known) are illustrated, and a distribution map is pre- sented based on an extensive compilation of specimen data. METHODS Dissections were made using standard techniques, abdomens being soaked in hot 10% potassium hydroxide solution for ap- proximately five minutes, and subsequently stored in glycerol. Morphological terms for Charis, Neotropics, Riodinini, Seco, xeric habitats genitalia follow Klots (1956) and Ehot (1973), and the terminology for wing ve- nation follows Comstock and Needham (1918). The protocol for listing material ex- amined follows Hall (1999). Seco specimens have been examined and their locality data recorded from the follow- ing collections: AME Allyn Museum of Entomology, Florida Museum of Natural His- tory, Sarasota, FL, USA AMNH American Museum of Natural His- tory, New York, NY, USA BMNH The Natural History Museum, London, UK CMNH Carnegie Museum of Natural His- tory, Pittsburgh, PA, USA DA Collection of David Ahrenholz, St. Paul, MN, USA JHKW _ Collection of Jason Hall & Keith Willmott, Washington, DC, USA MUSM Museo de Historia Natural, Univ- ersidad Nacional Mayor de San Marcos, Lima, Peru SME Senckenberg Museum, Frankfurt, Germany USNM_ National Museum of Natural His- tory, Smithsonian Institution, Washington, DC, USA 942 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bice le Adults of Seco species (dorsal surface on left, ventral surface on right, unless otherwise stated). A, 3 S. aphanis, lguazi, Argentina (BMNH). B, 2 S. aphanis, La Soledad, Argentina (BMNH). C, ¢ S. calagutis, nr. Macara, Ecuador (JHKW). D, 2 S. calagutis, Salto de Napac, Ecuador (USNM). E, ¢ S. ocellata, dorsal surface, Puerto Colombia, Colombia (CMNH). FE ¢ S. ocellata, ventral surface. ZMHU Museum fiir Naturkunde, Hum- boldt-Universitat zu Berlin, Ger- many REVISION OF SECO Seco Hall and Harvey 2002 (Figs. 1—4) Seco Hall and Harvey 2002: 415-417. Type species by original designation: Charis calagutis Hewitson 1871: pl. 46, figs. 11, 12: Diagnosis and systematic position.— Seco species are small riodinids (forewing length 10-12 mm) with compact wing shapes and rounded wings. The sexes are monomorphic. The dorsal surface is brown with prominent discal and postdiscal lines, the latter containing individual spots that VOLUME 104, NUMBER 4 are unusual in being outwardly curved, and a single submarginal silver line (rays along veins in aphanis). The fringe varies from entirely brown (ocellata) to entirely white (calagutis). The ventral surface differs only by having additional silver markings in the forewing apex (calagutis and ocellata only). This arrangement of submarginal sil- ver markings is unique in the family. All members of Detritivora and Charis have two dorsal submarginal silver lines and lack the elongate silver markings in the ventral forewing apex (Hall and Harvey 2002). The male genitalia of all Seco species possess the deep notch in the anterior mar- gin of the tegumen characteristic of the tribe Riodinini (Harvey 1987). The uncus is rectangular, the falces and tegumen are of average size and shape for the tribe, and the vinculum is evenly narrow and sinuate. The aedeagus is narrow and sigmoidal with a pointed tip that opens broadly; no cornuti are present. The pedicel is highly modified into the posteriorly elongate structure with fine acanthae (acellular projections) restrict- ed to a raised distal area (termed a “‘vogel- kopf” by Stichel 1910-11) that is so com- mon in the Riodinini. The valvae consist of a typically elongate and somewhat mem- branous lower process and a short upper process that 1s joined above the aedeagus to form a transtilla. The transtilla is developed into a pair of lateral upwardly curved pro- jections similar to those found in Chalodeta Stichel 1910 (Hall 2002), but in Seco spe- cies the transtilla is short instead of mark- edly elongate and downwardly curved, and the lateral projections are asymmetrical. The female genitalia are interspecifically variable. The corpus bursae is elongate, the signa small spine-like invaginations (cala- gutis) or very small sclerotized pads (aphanis), the ductus bursae membranous with only the very distal portion sclero- tized, the ductus seminalis membranous, the ostium bursae a simple hole either centrally positioned (calagutis) or displaced to the right (aphanis), and only abdominal sternite eight is sclerotized. 943 Seco appears to be most closely related to Detritivora, Charis and Calephelis, but its exact phylogenetic relationship to them and other closely related genera such as Chalodeta is not certain. Depending on how certain characters were coded and which outgroup(s) was used in the analysis, Hall and Harvey (2002) found Seco to var- iably be sister to Detritivora, Charis and Calephelis, sister to Charis and Calephelis or sister to Chalodeta and Detritivora. The first of these evolutionary scenarios seems the most plausible (Hall and Harvey 2002). KEY TO THE SPECIES OF SECO 1. Outer silver submarginal line on all wings complete and parallel to outer wing margin GEES) iC AIR) ot, gree Stites egestas cet vo raneyrs 1 2 — Outer silver submarginal line on all wings con- sists of rays along veins (Figs. 1A, B) .. aphanis 2. Postdiscal line displaced distally, submarginal black spots on all wings of even size (Figs. 1C, | D) Rarer tcsts kes Pre kat eo emcees CMC eee calagutis — Postdiscal line positioned medially, submargin- al black spots on all wings of uneven size (Figs. 1E, F) an Wig Bel > ng ie aa nad a ocellata Biology.—Seco species are distributed allopatrically in certain lowland xeric re- gions around the periphery of South Amer- ica. They occur in western Ecuador and Peru, northern Colombia and Venezuela, and southern Brazil, eastern Argentina and Uruguay. All Seco species are rare in col- lections and hence little is known of their biology. However, at the right time and place they appear to be relatively common. Males are encountered perching in small groups along streamsides or shaded forest paths within several meters of the ground and have been recorded nectaring from as- teraceous flowers (Hall and Willmott, un- published data). The food plants and early stages are unknown. Seco aphanis (Stichel 1910) (Pigs TA By ZAG 3As4) Charmona aphanis Stichel 1910: 19-20. Type locality: Uruguay. Syntypes, 2 ¢d and | 2, ZMHU [examined]. 944 Fig. 2. Male genitalia of Seco species in lateral view unless otherwise stated. A, S. aphanis, also tran- stilla tip in ventral view. B, S. calagutis, also pedicel and valve complex in ventral view. C, S. ocellata. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Identification and taxonomy.—Typical forewing length of both sexes 11 mm. Seco aphanis is readily distinguished from. its congeners by its possession of an outer sil- ver submarginal line on all wings that con- sists of rays along veins instead of a con- tinuous line paralleling the distal wing mar- gin, and sparsely setose instead of bare eyes. The male genitalia have a more prom- inent “‘vogelkopf’’ covered with more prominent acanthae, a short triangular up- per valve process, an asymmetrical laterally tilted transtilla, and lateral transtilla pro- cesses that are broad and dorso-ventrally in- stead of laterally compressed and inwardly as well as upwardly curved. This is the most poorly known of all Seco species and, perhaps because it is not sim- ilar in appearance to other Charis species, the genus in which it used to be treated (St- chel 1930, 1931; Bridges 1994; Callaghan and Lamas 2002), specimens in museums are curated under several different genera or, more typically, in “‘protem”’ drawers of unsorted material. Even in the BMNH (and hence in d’Abrera 1994), it is incorrectly identified zachaeus (Fabricius 1798): Biology.—Biezanko et al. (1979) report- ed that S. aphanis is ““frequent in the south- east region [of Rio Grande do Sul, Brazil], flying among flowering bushes in April.” The known months of capture are October and April, the earliest and latest months of summer. Distribution.—Seco aphanis is currently known from the southeastern Brazilian state of Rio Grande do Sul, eastern Argentina, and Uruguay (see Fig. 4), but it should also occur in southeastern Paraguay and at least as far north as the Brazilian state of Parana. Material examined.—ARGENTINA: Mi- siones, Iguazi (Oct) | ¢ BMNH; Entre Rios, La Soledad (Apr) 4 6, 1 2 BMNH. BRAZIL: Rio Grande do Sul, Pelotas 1 6, 1 2 SME URUGUAY: “Buschantal” [= forested valley] 1 ¢ SMF; No locality data 222 Ses ZMAWs as Audre VOLUME 104, NUMBER 4 0.5 mm A lester Se view. A, S. aphanis. B, S. calagutis. Female genitalia of Seco species in dorsal Seco calagutis (Hewitson 1871) (Bisse. D267 3B, :4) Charis calagutis Hewitson 1871: pl. 46, figs. 11-12. Type locality: La Chima, W. Ecuador. Syntype 6 BMNH [examined]. Identification and taxonomy.—Typical forewing length of both sexes 12 mm. Seco calagutis is most similar to S. ocellata but has a markedly distally displaced postdiscal line on all wings, submarginal black spots on all wings that are of an even size (in ocellata they are expanded into “‘eyespots”’ in forewing cells Cul and R4+5 and hindwing cell Cul), no additional silver tri- angle at the base of ventral forewing cell R4+5, and yellow-brown speckling across much of the ventral surface. The male gen- italia of S. calagutis have a vertically elon- gate and slightly recurved tip to the pedicel which is not weakly bifurcate as in S. ocel- lata, a more elongate and well sclerotized lower valve process, a considerably shorter and more angular upper valve process, a more posteriorly elongate transtilla, and 945 more elongate and prominently asymmet- rical lateral transtilla projections. Biology.—Seco calagutis is the most commonly represented species of the genus in collections. In Ecuador, it becomes in- creasingly rare towards the northern wetter end of its range, but can be locally common in the dry semi-deciduous woodlands of the southwest (Hall and Willmott, unpublished data). It has been recorded from sea-level to 1,100 m. In Ecuador, it has been collect- ed in March, May, and September, and in Peru during May, June, and October, all months around the beginning and end of the rainy season. Males are encountered perch- ing in small groups along forest edges, trails and streamsides, and more rarely on hilltops, between | and 3 m above the ground. They rest with their wings out- spread under the tips of leaves and _ their flight is somewhat weak. Males have been observed visiting flowers of Asteraceae (Hall and Willmott, unpublished data). Distribution.—Seco calagutis is restrict- ed to central and southwestern Ecuador and northwestern Peru (see Fig. 4). The follow- ing additional localities were reported by G. Lamas (personal communication, in coll. MUSM) for PERU: Tumbes, Matapalo, La Totora; Quebrada Angostura, Hualtaco; Puesto [de Vigilancia] Campo Verde; Ca- jJamarca, Hacienda La Vina. Material examined.—ECUADOR: Pi- chincha, Rio Tanti, Tinalandia | ¢ JHKW; Salto de Napac 1 2 USNM; Manabi, Pal- marcito, nr. Pedernales 1 6 DA; Ayampe | 6 JHKW; Los Rios, Rio Palenque 1 6 AME; Bolivar, La Chima 1 3, 1 2 BMNH; Chimborazo, Dos Puentes 3 6 AMNH; Chimbo 2 6 BMNH; Guayas, Duran [= Eloy Alfaro] 1 ¢d AMNH; Loja, Quebrada Curichanga, Rio Sabiango, nr. Macara 4 d JHKW. PERU: Piura, 3 km. W. of Canc- hague 1 6, 1 2 USNM; Ulunche 3.4 USNM;; La Tina 2 6 USNM. Seco ocellata (Hewitson 1867) (Fires Es sEeZ Ee 4) Symmachia ocellata Hewitson 1867: pl. 45, fig. 8. Type locality: Venezuela. Syn- types, 2 6, BMNH [examined]. 946 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Seco aphanis @ Seco calagutis A Seco occelata Fig. 4. Identification and taxonomy.—Typical forewing length of male 12 mm. Seco ocellata is most similar to its sister spe- cies §. calagutis and is distinguished in that species account. The female is un- known to us. Biology.—This is the most poorly rep- resented Seco species in collections, per- haps only because it is endemic to an infre- quently collected region. The label data of a Colombian specimen indicate the habitat 500 1000 Map of South America illustrating distributions of Seco species. of the species to be “dry hills with thick scrub.” Distribution.— Seco ocellata is restricted to northern Colombia and presumably the dry regions of northwestern Venezuela (the only Venezuelan specimens examined have no precise data). Material examined.mCOLOMBIA: At- ldntico, Puerto Colombia (Jul) 1 d CMNH; Magdalena, Atanques 3 6 BMNH. VE- NEZUELA: No locality data 2 6 BMNH. VOLUME 104, NUMBER 4 ACKNOWLEDGMENTS We thank the following for giving us ac- cess to the riodinid collections in their care and for the loan of specimens: P. Ackery (BMNH), W. Mey (ZMHU), L. and J. Mill- er (AME), J. Miller & EF Rindge (AMNH), W. Nassig (SMB), J. Rawlins (CMNH); and G. Lamas for providing locality data for S. calagutis from the MUSM. JPWH. thanks the Museo Nacional de Ciencias Naturales and INEFAN, in Quito, for arranging the necessary permits for research in Ecuador, and the following for the financial assis- tance of field and museum research: Oxford University (Poulton Fund), Cambridge Uni- versity (Christ’s College), The Royal En- tomological Society, The American Muse- um of Natural History (Collections Studies Grant), Sigma Xi, Equafor, The National Geographic Society (Research and Explo- ration Grant # 5751-96), The Smithsonian Institution (two postdoctoral fellowships) and The National Science Foundation (DEB # 0103746). We thank Keith Will- mott and an anonymous reviewer for com- ments on the manuscript. LITERATURE CITED Biezanko, C. M., O. H. H. Mielke, and A. Wedderhoff. 1979. Contribuigao ao estudo faunistico dos Riod- inidae do Rio Grande do Sul, Brasil (Lepidoptera). Acta Biologica Paranaense 7: 7—22. Bridges, C. A. 1994. Catalogue of the Family-Group, Genus-Group and Species-Group Names of the Riodinidae and Lycaenidae (Lepidoptera) of the World. C. Bridges, Urbana. 1113 pp. Callaghan, C. J. and G. Lamas. 2002. Riodinidae. /n Lamas, G., ed. A Checklist of the Neotropical Butterflies and Skippers (Lepidoptera: Papiliono- idea and Hesperioidea). Atlas of Neotropical Lep- idoptera (ed. by J. B. Heppner). Scientific Pub- lishers, Gainesville. In press. 947 Comstock, J. H. and J. G. Needham. 1918. The wings of Insects. American Naturalist 32: 231-257. d’Abrera, B. 1994. Butterflies of the Neotropical Re- gion, Part VI. Riodinidae. Hill House, Victoria, Australia. Pp. 880-1096. Eliot, J. N. 1973. The higher classification of the Ly- caenidae (Lepidoptera): A tentative arrangement. Bulletin of the British Museum of Natural History (Entomology) 28: 373-506. Hall, J. P. W. 1999. A Revision of the Genus Theope: Its Systematics and Biology (Lepidoptera: Riodi- nidae). Scientific Publishers, Gainesville. 127 pp. . 2002. A review of Chalodeta with a revision of the Chalodeta chelonis group (Lepidoptera: Riodinidae). Proceedings of the Entomological Society of Washington 104: 376-389. Hall, J. PB. W. and D. J. Harvey. 2001. A phylogenetic revision of the Charis gynaea group (Lepidoptera: Riodinidae) with comments on historical relation- ships among Neotropical areas of endemism. An- nals of the Entomological Society of America 94(5): 631-647. . 2002. A phylogenetic review of Charis and Calephelis (Lepidoptera: Riodinidae). Annals of the Entomological Society of America 95: 407— 421. Harvey, D. J. 1987. The Higher Classification of the Riodinidae (Lepidoptera). Ph.D Dissertation, Uni- versity of Texas, Austin. 216 pp. Harvey, D. J. and J. P. W. Hall. 2002. Phylogenetic revision of the Charis cleonus complex (Lepidop- tera: Riodinidae). Systematic Entomology 27: 265-300. Hewitson, W. C. 1867-71. Illustrations of New Species of Exotic Butterflies, Selected Chiefly from the Collections of W. Wilson Saunders and William C. Hewitson, Vol. 4. J. Van Voorst, London. Klots, A. B. 1956. Lepidoptera, pp. 97-110. Jn Tuxen, S. L., ed. Taxonomists’ Glossary of Genitalia in Insects. Munksgaard, Copenhagen, Denmark. Stichel, H. EK E. J. 1910. Vorarbeiten zu einer Revision der Riodinidae Grote (Erycinidae Swains.) (Lep. Rhop.). Berliner Entomologische Zeitschrift 55(1/ 2): 9-103. 1910-11. Lepidoptera Rhopalocera. Fam. Riodinidae, pp. 1—238. Jn Wytsman, J., ed. Genera Insectorum 112. J. Wytsman, Brussels. . 1930-31. Riodinidae, pp. 1-795. Jn Strand, E., ed. Lepidopterorum Catalogus 38-41. W. Junk, Berlin. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 948-954 REDISCOVERY OF THE GENUS PLATYSCELIDRIS SZABO (HYMENOPTERA: SCELIONIDAE) AND DESCRIPTION OF A NEW SPECIES NORMAN E JOHNSON AND LUCIANA MUSETTI Department of Entomology, The Ohio State University, 1315 Kinnear Road, Columbus, Ohio 43212-1192, U.S.A. (e-mail: johnson.2 @osu.edu) Abstract.—Platyscelidris kittenbergeri Szabo is redescribed, and a new species, P. fos- sorius, is described from Madagascar. The genus is remarkably similar in superficial characters to the Australian Mirobaeoides Dodd, including the possession of a pair of apical femoral spines in P. fossorius. The structure of the antenna and ovipositor place the genus within the Scelionini sensu Austin and Field. Key Words: egg- Many species of parasitoid wasps of the family Scelionidae display degrees of wing reduction. Some have wings that are merely shortened; others are practically apterous, with the wings so reduced as to be barely visible. This phenomenon is not restricted to any clade within the family, and it may be related, in part, to the need for females to search for host eggs within the litter and soil. Szab6 (1959) described Platyscelidris as a new genus of Scelionidae on the basis of a single brachypterous female specimen collected in East Africa in 1905 by the Hungarian big-game hunter Kalman Kitten- berger. Since then, no further specimens have been reported. The original descrip- tion contains two highly unusual features for a scelionid. The dorsal surface of the mesosoma was said to be made up of only two sclerites, the mesoscutum and the me- tanotum. Szabo asserted that the scutellum is lacking. Brachyptery is commonly asso- ciated with modifications in mesosomatic structure, but the absence of a scutellum is unprecedented. Second, the first metaso- matic, tergite is reduced: ~“Petiole very small, present as a semicircular field in the arasitoid, Afrotropical, Madagascar middle of the anterior margin of the second tergite”’ (our translation). The genus Baeus Haliday has T1 hidden in dorsal view, but most species have this sclerite well devel- oped. Szab6 considered the genus to be an in- termediate form between the subfamilies Baeinae and Scelioninae, but its precise po- sition within the family was uncertain. Mas- ner (1976) included a brief treatment of Platyscelidris in his review of the world Scelionidae and provisionally placed it in the tribe Gryonini (Scelioninae). In his key to genera, Platyscelidris falls in the section in which the genera of Baeini (including Id- rini) and Gryonini are distinguished, with the main diagnostic character being the re- duction in size of the first metasomatic ter- gite. We recently sorted a litter sample from Madagascar and discovered specimens of an unusual micropterous species. Our initial impression was that it was a species of Gryon Haliday or Mirobaeoides Dodd, but closer examination and a review of the lit- erature led us to conclude that we had found a new species of Platyscelidris. We here redescribe the genus and its type spe- VOLUME 104, NUMBER 4 cies, describe a new species, and discuss relationships of the genus within the Sce- lionidae. MATERIALS AND METHODS Specimens for this study are deposited in the collections of the Ohio State University (OSUC), Canadian National Collection of Insects (CNCI), National Museum of Nat- ural History, Smithsonian Institution, Washington, DC (USNM), and the Hungar- ian Natural History Museum (HNHM). The figures were prepared using a Spot Insight Color® camera (Diagnostic Instruments, Inc.), Auto-Montage version 3.05(3) (Syn- croscopy, Inc.), and Adobe® Photoshop® 6.0.1 software. The OSUC numbers refer to unique identifying bar code labels attached to the specimens. Platyscelidris Szabo Platyscelidris Szab6 1959: 45, original de- scription. Type species: Platyscelidris kit- tenbergeri Szab6, by monotypy and orig- inal designation. Masner 1976: 60, de- scription; Johnson 1992: 461, catalog of world species. Description.—Female. Head and com- pound eyes large, prominent; frons convex throughout, without deep scrobe; lateral ocelli contiguous with inner orbits; vertex separated from occiput by sharp angle or broadly rounded; palpal formula apparently 3-2 (palpi very small and largely hidden); antenna 12-merous, clava 6-merous, claval formula A1l2-A8/1-2-2-2-1; dorsum of me- sosoma flattened; skaphion absent, but me- soscutum strongly angled anteriorly; notauli absent; scutellum unarmed; metanotum, where visible, unarmed, appearing as a sim- ple transverse strip; propodeum hidden mesally; tibial spur formula 1-1-1; wings highly reduced or absent; metasoma with 7 tergites, 6 sternites, T7 extruded with ovi- positor; laterotergites incised into sternites, submarginal ridge developed; Tl trans- verse; T2 distinctly longer than T3. ¢ un- known. 949 Diagnosis.—The 12-merous antenna will distinguish Platyscelidris trom the superfi- cially similar Australian genus Mirobaeo- ides and all other Baeini (see below). The hidden propodeum and the narrowing of T1 laterally will adequately separate it from species in the diverse genus Gryon. Discussion.—Masner’s (1976: 7) key to world genera of Scelioninae must be adapt- ed to include the new species described be- low. 24(23). Metasoma apparently sessile, Tl much narrower than T2, appearing as a fusiform or small nail-like sclerite situated medi- ally on front margin of T2; scutellum ap- pearing as a narrow transverse sclerite; propodeum entirely hidden medially; mi- cropterous (male unknown) ........ Sh shearer ae Platyscelidris Szab6 p. 60 - Metasoma variable, Tl normal, almost as wide as T2, transverse; scutellum variable in shape, usually semicircular; propo- deum variable, usually entirely visible: wings variable KEY TO SPECIES OF PLATYSCELIDRIS 1. Mesosoma in dorsal view with large mesos- cutum followed by single transverse sclerite (scutellum) (Fig. 2); T1 in form of small hemi- spherical sclerite on anterior margin of T2 (Fig. 2); hind femur without apical spines; hyper- P. kittenbergeri Szabo — Mesosoma with mesoscutum followed by two occipital carina absent .. . transverse sclerites (scutellum and metanotum) (Fig. 4); Tl forming relatively wide transverse fusiform sclerite (Fig. 4); hind femur with pair of strong apical spines; hyperoccipital carina present P. fossorius, new species Platyscelidris kittenbergeri Szab6 (Figs. 1-2) Platyscelidris Kittenbergeri Szab6 1959: 46, original description; Masner 1976: 60, type information. Female.—Mesosoma, metasoma, A8- A12 brown; head, legs, Al—7 light brown; A1-A7 brown to yellowish brown; length: 0.81 mm. Head transverse in dorsal view, width 0.41 mm, wider than mesosoma, ratio of head width to mesosoma width 1.15: com- pound eyes hairless; hyperoccipital carina 950 Figs. 1—4. fossorius, holotype °, absent, vertex broadly rounded onto occi- put, aciculate; gena narrowly visible dor- sally, with sharp carina parallel to longitu- dinal axis of body arising from posterior or- bit and intersecting occipital carina; com- pound eyes not bulging, in dorsal view margin in continuous line with frons; oc- cipital carina largely obscured from view; occiput obscured medially by mesoscutum; head in frontal view oval, wider than long, vertex gently arched; ocelli minute; com- pound eyes relatively small, frons width 0.53 times head width, without distinct frontal depression or central keel; gena ap- pearing convex above anterior mandibular articulation, no surface sculpture visible; lower frons without distinct fanlike striae, with fine aciculate microsculpture, other ir- regular, asymmetical ridges present, espe- cially ventrally, probably remnants of glue; apex of mandible not visible. 1, Platyscelidris kittenbergeri, holotype &, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4 lateral view. 2, Same, dorsal view. 3, Platyscelidris lateral view. 4, Same, dorsal view. Scale line = 0.50 mm. Antenna |2-segmented, clava 6-segment- ed; claval formula not visible. Mesosoma slightly narrower than the me- tasoma, ratio of mesosoma width to meta- soma width 0.82; pronotum not visible in dorsal view; mesoscutum semicircular, dis- tinctly overhanging head anteriorly, surface aciculate, with evenly spaced setigerous punctures, associated with fine minute hairs, lateral margins gently raised, punc- tures absent along posterior margin, notauli absent; absent; mesoscutellum strongly transverse, 0.27 times length of mesoscutum, sculptured as mesoscutum; without distinct axillary pits; metanotum, propodeum not visible dorsally; lateral sur- face of pronotum with deep femoral de- pression, longitudinally carinate ventrally, with fine aciculate microsculpture above; fore and mid coxae closely approximated, ventral portion of mesepisternum obscured; wings VOLUME 104, NUMBER 4 mesopleural carina formed by raised ridge, crenulate posteriorly; mesepisternum ante- rior to mesopleural carina aciculate; meso- pleural scrobe very finely sculptured, me- sopleural pit present, suture between meso- and metapleuron distinct; antero-ventral corner of metapleuron obscured; metapleu- ron glabrous, metapleural pit not visible, metapleuron wrinkled, distinctly separated from lateral propodeum by ridge: lateral surface of propodeum smooth, posterior edge of metapleuron and propodeum nearly straight, without lamella; apex of hind fe- mur without apical spines. Metasoma 1.22 times longer than wide, entirely without longitudinal sculpture, strongly and evenly convex anteriorly; T] short, length 0.04 mm, longest medially, narrow, not reaching lateral margins of me- tasoma, aciculate, with very few setigerous punctures; T2 longest segment, length 0.24 mm, transverse, aciculate; T3 length 0.12 mm, punctures absent on posterior border, aciculate to faintly aciculate; T6 without visible setae; T7 hidden; laterotergites smooth, glabrous; S2—S6, where visible, with similar surface sculpture as tergites. Male.—Unknown. Host.—Unknown. Material examined.—Holotype 2 (ex- amined, HNHM): ‘‘Mto-ja-Kifaru, Febr. 905 [on reverse]; Africa or., Katona: Baei- dae n. gen.; Platyscelidris g.n. Kittenbergeri sp.n. - det. J. B. Szab6; Holotypus! 1958 J. B. Szabo [red label]; Holotypus @ Platys- celidris kittenbergeri Szab6 1959 [red mar- gined label]; Hym. Typ. No. 9572 Mus. Bu- dapest.”’ The specimen is in only fair con- dition, somewhat dirty, and glued on a small card. The second metasomatic tergite is cracked. Much of the surface sculpture of the body is obscured, probably because Szab6 seems to have enhanced the magni- fication of his microscope by immersing the specimen in a drop of water, thus dissolving and spreading some of the adhesive (**Wan- gen im Wassertropf gesehen ...,’’ Szab6 1959: 46). According to Masner (1976), the type locality is in present-day Tanzania. 95] Wheeler (1922) also cited this locality, as ““Mto-ya-Kifaru,”” placed it in German East Africa, and equated it to Arusha-chini, 335'S387225E. Diagnosis.—Distinguished from the new species described below, P. fossorius, by the apparent lack of a metanotum, the lon- gitudinal carina on the gena, the rounded vertex without a hyperoccipital carina, and the narrow, semicircular T1. Platyscelidris fossorius Johnson and Musetti, new species (Figs. 3—4) Female.—General coloration dark brown to black; legs, mandible brown; A1l—A7 brown to yellowish brown, A8-A12 dark brown; length: 0.90—-1.16 mm (mean = 1.08 mm, SD = 0.08 mm). Head transverse in dorsal view, width 0.48—0.55 mm (mean = 0.52 mm, SD = 0.02 mm), slightly arched around mesoso- ma, wider than mesosoma, ratio of head width to mesosoma width 1.05—1.15 (mean = 1.10, SD = 0.03); compound eyes hair- less; surface sculpture aciculate, with even- ly spaced setigerous punctures associated with fine minute hairs; hyperoccipital carina along vertex sharp, angle between vertex and occiput acute; gena not visible dorsally; compound eyes not bulging, margin in con- tinuous line with frons; occipital carina ex- tended ventrally to posterior mandibular ar- ticulation, strongly developed laterally, in- terrupted medially; occiput distinctly concave, aciculate; head in frontal view oval, vertex gently arched; ocelli minute; compound eyes relatively small, frons width 0.46—0.52 times head width (mean = 0.50, SD = 0.02), without distinct frontal depression or central keel; frons and vertex covered with evenly spaced setigerous punctures, with fine minute hairs; gena dis- tinctly reflexed above anterior mandibular articulation, surface aciculate; lower frons finely striate on either side of malar suture, striae radiating from clypeal margin, frons otherwise finely aciculate; mandible triden- 952 tate, with lower tooth longest, lower and upper teeth longer than median tooth. Antenna |12-segmented, clava 6-segment- ed; claval formula A8-A12 1-2-2-2-1. Mesosoma slightly narrower than the me- tasoma, ratio of mesosoma width to meta- soma width 0.84—0.90 (mean = 0.87, SD = (0.02); pronotum not visible in dorsal view; mesoscutum distinctly transverse, with anterior corners rounded, surface acic- ulate, with evenly spaced setigerous punc- tures, associated with fine minute hairs, lat- eral margins gently raised, punctures absent and faintly aciculate along posterior mar- gin, notauli absent; mesoscutellum strongly transverse, 0.27—0.41 times length of me- soscutum (mean = 0.34, SD = 0.04), sculp- tured as mesoscutum; two axillary pits an- teriorly on either side of mesoscutellum along transscutal articulation; metanotum clearly visible, very narrow, length 0.46— 0.68 times length of mesoscutellum (mean = 0.55, SD = 0.07), length 0.62—1.85 times length T1 (mean = 0.99, SD = 0.41), sur- face aciculate with 1 row of evenly spaced setigerous punctures; propodeum visible dorsally as triangular sclerites on either side of metanotum, surface imbricate-aciculate; lateral surface of pronotum with femoral depression, with longitudinal wrinkles ven- trally, otherwise finely aciculate through- out; fore and mid coxae distinctly separated by mesepisternum ventrally; mesopleural carina formed by simple raised ridge with no crenulations; mesepisternum anterior to mesopleural carina aciculate; mesopleural scrobe very finely sculptured, mesopleural pit present, suture between meso- and me- tapleuron distinct; antero-ventral corner of metapleuron produced ventrally to narrow sharp point; metapleuron glabrous, with distinct round metapleural pit, longitudinal sulcus extending from pit to posterior edge of propodeum; lower *% of lateral surface of propodeum finely sculptured, smooth above, posterior edge of metapleuron and propodeum expanded into thin lamella; apex of hind femur with a pair of spines PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON flanking base of tibia, posterior spine dis- tinctly longer. Metasoma 1.25—1.44 times longer than wide (mean = 1.31, SD = 0.06), entirely without longitudinal sculpture, strongly and evenly convex anteriorly; T1 short, length 0.03—0.06 mm (mean = 0.05 mm, SD = 0.01 mm), longest medially, aciculate, with very few setigerous punctures; T2 the lon- gest segment, length 0.23—0.32 mm (mean = 0.29, sd = 0.03 mm) transverse, acicu- late, punctures absent on anterior third, punctures also absent posteriorly, sculpture faintly aciculate around border; T3 length 0.17—0.24 mm (mean = 0.21 mm, SD = 0.02 mm), punctures absent on posterior border, aciculate to faintly aciculate; T6 covered with very short setae; T7 with long recurved setae on each side; laterotergites smooth, glabrous; S2— S6 with similar sur- face sculpture as tergites. Male.—Unknown. Host.—Unknown. Material examined.—Holotype 2: MADAGASCAR: Tuléar Prov., 18 km NNW Betroka, 825 m, 23°09'48"S 48°58'O7’E, 24.xi-14.xii.1994, M. A. Ivie and D. A. Pollock, OSUC 60936 (deposited in OSUC). Paratypes—Two @ with same data as holotype: OSUC 60919, OSUC 60905 (OSUC). MADAGASCAR: Prov. Toliara; lfaty;<23°09", 43°37'E; 17-22) Sept: 1993; pitfall trap in sand, desert scrub forest; collrs. W. E. Steiner, R. Andriamassiman- ana, 1 2: OSUC 90316 (CNC); PT im de- sent scrub: ‘W. Stemer 3 2sOSUG@ 90817: OSUC 90318, OSUC 90319 (USNM). Diagnosis.—Distinguished from P. kit- tenbergeri by the presence of a third dorsal mesosomatic sclerite (the metanotum), the lack of a longitudinal genal carina, the pres- ence of the hyperoccipital carina, and the wide, fusiform T1. Discussion.—The specimens of P. fos- sorius strongly resemble the genus Miro- baeoides, a group of 14 species known only from mainland Australia, Tasmania, and Lord Howe Island (Austin 1986, Johnson VOLUME 104, NUMBER 4 1992). Most female baeines are easily rec- ognized by their 7-segmented antenna with a large, often globose, clava. Mirobaeoides is exceptional in this regard, because the fe- male antenna is | 1-merous with four clearly separated apical clavomeres. One of the de- fining features of Mirobaeoides, in both sexes, is the possession of a pair of spines at the apex of the hind femur (Austin 1986). The only similar structure described in sce- lionids are the genual spines of some spe- cies of Gryon. These spines, however, are more irregular in distribution among spe- cies and are found at the base of the hind tibia. The specimens of P. fossorius have well-developed apical femoral spines. De- spite the difference in the number of anten- nomeres, we were initially excited to dis- cover an apparent new species of Miro- baeoides on the opposite side of the Indian Ocean. One of the specimens forced us to re- evaluate this conclusion. The apex of the metasoma of this female has a short, cylin- drical, membranous tube that bears the sev- enth metasomatic tergite at its apex (Figs. 3—4). We have not made detailed dissec- tions of the limited material available, but this formation has been described by Field and Austin (1994) and Austin and Field (1997) as a Scelio-type ovipositor. This is a set of morphological specializations that produces a long telescoping tube bearing the skeletal elements of the ovipositor at its apex. The length of this tube may be several times the apparent length of the entire me- tasoma and presumably allows the female much greater range for reaching and para- sitizing host eggs. Austin and Field (1997) suggested that the complex of apomorphic features of this ovipositor-type may be used to define a monophyletic group within the subfamily Scelioninae, their Scelionini sen- su lato, a grouping that dismembers several of the tribes defined by Kozlov (1970) and Masner (1976). All known Baeini have the plesiomorphic appendicular Ceratobaeus- type ovipositor. We concluded that the Madagascan specimens could not be as- 953 signed to an expanded concept of Miro- baeoides, but must somehow fit within the concept of Scelionini outlined by Austin and Field (1997). The Scelio-type ovipositor is a highly complex and integrated set of morphologi- cal specializations. We agree with Austin and Field (1997) that joint possession of this set of characters is strong evidence of phylogenetic affinity and can be of help in understanding scelionid relationships. Some groups, e.g., Opisthacantha Ashmead, have some species with the Scelio-type oviposi- tor and others with the Ceratobaeus-type ovipositor. This may well hint that such a taxon needs to be re-evaluated. We would like to add a note of caution, however. The supposition that the elements of this char- acter set are integrated into an emergent morphological structure implies that the characters themselves may not be indepen- dent. Without information on the genetic and developmental mechanisms, it seems plausible to us that any viable character re- versal may require reversion of the entire structure to the plesiomorphic Ceratobaeus- type condition. The best evidence of the utility of a character set is its compatibility with the totality of other characters and not an a priori assumption of its importance. Szabo (1959) suggested that Platysceli- dris represents a link between his concepts of Scelioninae and Baeinae. Most authors today consider the Baeinae to be a tribe within Scelioninae, so that hypothesis es- sentially proposes little other than a vague similarity to baeines. The Scelio-type ovi- positor allies the genus with the Scelionini sensu lato, and not with the Baeini Gryonini (as suggested provisionally by Masner 1976). A large part of the difficulty in finding the proper context for this genus arises from the paucity of characters that circumscribe the subfamilies and tribes within Scelionidae. Resolution of the phy- logeny of this family would help greatly to resolve such issues as well as provide valu- able insights into the mechanisms of host- nor 954 finding and shifts from one host taxon to another. ACKNOWLEDGMENTS Thanks to our good friends Michael A. Ivie (Bozeman, Montana) for passing on to us the samples of specimens from Mada- gascar, Lubomir Masner (Ottawa, Ontario) for searching for and loaning to us the ad- ditional material, Warren Steiner and David Furth (Washington, DC) for materal from the Smithsonian Institution, and Andrew D. Austin (Glen Osmond, South Australia) for his insights. Thanks also to Istvan Miko (K6szeg, Hungary) for the loan of the ho- lotype from the Hungarian National Muse- um of Natural History. This material is based upon work supported by the National Science Foundation under Grant No. DEB- 9521648. LITERATURE CITED Austin, A. D. 1986. A taxonomic revision of the genus Mirobaeoides Dodd (Hymenoptera: Scelionidae). Australian Journal of Zoology 34: 315-337. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Austin, A. D. and S. A. Field. 1997. The ovipositor system of scelionid and platygastrid wasps (Hy- menoptera: Platygastroidea): Comparative mor- phology and phylogenetic implications. Inverte- brate Taxonomy 11: 1-87. Field, S. A. and A. D. Austin. 1994. Morphology and mechanics of the ovipositor system of Scelio La- treille (Hymenoptera: Scelionidae) and related genera. International Journal of Insect Morpholo- gy and Embryology 23: 135-158. Johnson, N. FE 1992. Catalog of world Proctotrupoidea excluding Platygastridae. Memoirs of the Ameri- can Entomological Institute No. 51. 825 pp. Kozlov, M. A. 1970. [Supergeneric groupings of Proc- totrupoidea (Hymenoptera).] Entomologicheskoye Obozreniye 49: 203-226. (English translation, 1970 Entomological Review 49: 115-127.) Masner, L. 1976. Revisionary notes and keys to world genera of Scelionidae (Hymenoptera: Proctotru- poidea). Memoirs of the Entomological Society of Canada 97: 1-87. Szabo, J. B. 1959. Platyscelidris gen. nov., eine neue exotische Gattung der Scelioniden aus Afrika. Opuscula Zoologica (Budapest) 3: 45—49. Wheeler, W. M. 1922. Ants of the American Museum Congo Expedition. A contribution to the myrme- cology of Africa. Bulletin of the American Mu- seum of Natural History 45: 1-1139. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 955-963 TWO NEW SPECIES OF ORSILLINE LYGAEIDAE (HETEROPTERA) FROM THE HAWAIIAN ISLANDS DAN A. POLHEMUS Department of Systematic Biology—Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560-0105, U.S.A. (e-mail: bugman@ bishopmuseum.org ) Abstract.—Two new species of orsilline Lygaeidae are described from the Hawaiian Islands: Oceanides gruneri, taken in canopy fogging samples from ohia trees (Metrosi- deros polymorpha) on eastern Molokai; and Nesius (Physonysius) poepoe, collected from understory vegetation in the upper elevations of the West Maui Mountains. Shaded habitus drawings and distribution maps are provided for both new species, the male paramere of N. (Physonysius) poepoe is illustrated, and a revised key to the species in the subgenus Physonysius is included. Key Words: Lygaeidae, Hawaii, Oceanides, Nesius, new species, illustrations, key As interpreted by Ashlock (1967), the tribe Metrargini in the Lygaeidae contains 6 genera endemic to the Hawaiian Islands, which have produced a diverse insular ra- diation of 80 endemic species and subspe- cies (Usinger 1942; Usinger and Ashlock 1958; Ashlock 1966, 1983; Nishida 1997). Although most of the species occurring in obvious or easily accessible habitats have now been described, additional new species are discovered on a regular basis in remote localities or unusual microhabitats. The pre- sent paper describes two such taxa, taken from upland wet forest habitats on the is- lands of Maui and Molokai. In the descriptions below, all measure- ments are given in millimeters, and repre- sent the dimensions of the holotype speci- men. CL numbers in the material examined sections refer to a coding system used to cross reference ecological data and habitat photographs. Collection depository codons are given in the acknowledgments section. English system equivalents of metric ele- vational data are given in brackets, to pro- vide compatability with USGS topographic maps. Holotypes of both new species described herein are deposited in the National Muse- um of Natural History, Smithsonian Insti- tution, Washington, DC (USNM). Paratypes are also deposited in that collection, and in the Bernice P. Bishop Museum in Honolulu (BPBM). Oceanides gruneri Polhemus, new species (Figs. 1—2) Description.—Macropterous male: Over- all length 4.70; maximum width (across ab- domen) 1.85. General coloration dark brown to black, with extensive pale coloration on pronotum and hemelytra. Head: Length 0.85, width 1.10, general coloration black, tip of tylus dark yellow, eyes and ocelli pale reddish; dorsal surface slightly elevated between eyes, coarsely ru- gulose, sparingly clothed with recumbent pale pubescence, this pubescence denser in areas behind eyes and laterad to ocelli; an- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 956 itus. dorsal habi > Oceanides gruneri, male Fig. 1. VOLUME 104, NUMBER 4 Kolekole Cabin MOLOKAI Fig. 2. teocular length 0.42, equal to 1.2 dorsal length of an eye; eye length 0.35, width 0.25; interocular space 1.12; ocelli small; buccula low, gradually tapering in height, without abrupt change in width to base of head; rostrum pale brown, apex dark, reach- ing beyond hind coxae, segment I nearly reaching base of head, segment lengths (from base) 0.75, 0.70, 0.75, 0.65; antennae long, slender, segments I and IV distinctly thicker than IT and III, all segments dark brown, extreme base of segment I and ex- treme distal ends of segments I-III creamy white, all segments bearing short erect pale setae, these setae slightly shorter than the diameters of these segments, segment I with 3 longer setae near base, lengths of seg- ments I-IV = 0.35, 0.65, 0.60, 0.60. Pronotum: Length 0.55, maximum width 1.55, dark brown to blackish, with yellowish-white markings narrowly along extreme anterior margin, broadly on central portion of posterior lobe, and on postero- lateral angles; calli black, shining, margins bearing pale recumbent pubescence, ante- rior collar and posterior lobe shining, with- out evident setae, surfaces sparsely set with O5i/ Oceanides gruneri Distribution of Oceanides gruneri on Molokai. deep, dark brown punctations; lateral pro- notal margins bulging outward at lateral margins of calli. Scutellum: Length 0.52, maximum width 0.80, black, bearing a narrow, raised, lon- gitudinal medial carina on posterior half, marked by a slender white longitudinal stripe; surface coarsely rugulose, thickly clothed with recumbent pale pubescence. Hemelytra: Macropterous, extending well beyond end of abdomen, weakly ru- gose, bearing scattered pale recumbent se- tae; general coloration yellowish white with scattered dark markings, clavus entirely pale except for narrow dark patch along commisure, and lines of dark brown punc- tations on either side of claval suture; co- rium bearing a single roughly circular blackish spot in basal half traversing the cu- bitus, two additional larger and more irreg- ular blackish patches on distal half adjacent to base of wing membrane, and a small dark mark at extreme posterior angle; membrane uniformly pale translucent brown; costal margins broadly and evenly arcuate. Legs: Yellowish white, dorsal sufaces of femora spotted with dark brown, dorsal fac- 958 es of tibiae and tarsi narrowly medium brown; all segments clothed with fine, pale, semi-recumbent setae intermixed with scat- tered longer erect pale setae; tibiae slightly expanded near distal ends. Lengths of leg segments as follows: fore femur, tibia, tarsal I, tarsal I, tarsal 1 = 1212; 1.005 0:20, 0.10, 0.20; middle femur, tibia, tarsal I, tar- Sal Wie MtarsalewiL => 12 AkOO! 025520510! 0.20; hind femur, tibia, tarsal I, tarsal II, tarsal We 1.252405, 035) O80. 0x12: Ventral surface: Black, acetabula and peritreme dark yellow, adbominal ventrites VI-VIII marked with medium brown; ven- trolateral portions of head and thorax and all of abdomen bearing numerous pale re- cumbent setae. Macropterous female: Length 5.15; maximum width 2.00. Similar to male in general structure and coloration with fol- lowing exceptions: posteromedial portion of pronotum not so broadly marked with yellowish white; ventromedial portion of abdomen basal to ovipositor sheath broadly marked with dark yellow. Etymology.—It is my pleasure to name this species in honor of Daniel S. Gruner, a friend and colleague whose canopy fogging study of Metrosideros polymorpha, under- taken during his tenure as a graduate stu- dent at the University of Hawai, has great- ly increased our understanding of speciation and diversification processes in the Hawai- ian insect biota. Material examined.—Holotype, macrop- terous 6: HAWAIIAN ISLANDS, Molokai Is., rain forest at Kolekole Cabin, Kamakou TNCH Preserve, 1,220 m [4,000 ft], PALOGISUN 5075347" We 23) October 1997, CL 8071, canopy fogging sample MO97-1+2-Sc, D. A. Polhemus and D. Gruner (USNM). PARATYPES (all mac- ropterous): HAWAHAN ISLANDS, Molo- kai Is.: | ¢, same data as holotype except canopy fogging sample MO9705-1-1 (USNM); 1 2, same data as holotype ex- cept canopy fogging sample MO9704-2-1 (USNM); 1 2, same data as holotype ex- cept taken from bundle of dead Cibotium PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tree fern fronds, 21—25 October 1997, D. A. Polhemus (USNM); 2 6, Kamakou Pre- serve, road between cabin and Puu Kole- kole,, 1,170)me*(3;840 ft], 22 October 1997: Coll. #85, C. P. Ewing, on Cheirodendron trigynum (BPBM); | @, lee side of Puu Lua, nr. rim of Wailau Valley, 2,750—3,000 ft., 15 June 1999, 21°06'28’N, 156°48'48"W, CL 8342, D. A. Polhemus, on Cheirodendron sp. (USNM). Discussion.—Oceanides gruneri belongs to a distinctive set of Oceanides species with contrasting black and white dorsal markings. It is similar in general appear- ance to O. humeralis Ashlock from Kauai, O. perkinsi Usinger and O. incognitus Usin- ger from the Koolau Mountains of Oahu, and O. oresitrophus (Kirkaldy) from East Maui, and it seems likely that all of these species will eventually prove to be mem- bers of a monophyletic segregate within the Oceanides lineage. Both O. gruneri and O. oresitrophus may be separated from O. hu- meralis, O. perkinsi, and O. incognitus by the presence of a black spot transgressing the cubital vein in the basal half of the hem- elytron. In turn, O. gruneri and O. oresitro- phus may be separated from each other by the length of the anteocular portion of the head, which is 1.2 the dorsal length of an eye in O. gruneri but equal to the eye length in O. oresitrophus; by the length of antennal segment II, which is 1.8 the length of antennal segment I in O. gruneri, versus 2.0 the length of segment I in O. oresitrophus; and by the presence of an ir- regular patch of pale coloration centrally on the posterior pronotum in O. gruneri, which is lacking in O. oresitrophus. The pale markings on the corium of O. gruneri also have a more yellowish overall hue than those of O. oresitrophus, which are a clear- er, brighter white. Most captures of O. gruneri to date have come from canopy fogging samples taken from ohia trees (Metrosideros polymorpha). It is uncertain whether this species occurs in the canopy itself, or on the pads of moss that cover many of the branches; the col- VOLUME 104, NUMBER 4 lection of several additional specimens on Cheirodendron trigynum trees suggests the latter. In any case, the lack of previous cap- tures at the relatively well-collected Kole- kole Cabin type locality on Molokai by en- tomologists operating near ground level strongly indicates an arboreal habit for this species. Nesius (Physonysius) poepoe Polhemus, new species (Figs. 3—5) Description.—Submacropterous male: Overall length 3.80; maximum width (across abdomen) 1.75. General coloration uniform shining reddish brown, with a few contrasting paler or darker markings on head and scutellum. Head: Length 0.55, width 1.00, general coloration reddish brown, tylus, vertex ad- joining inner margins of eyes, and postero- medial portion of vertex dark yellow, pos- terior margin of head dark brown, eyes sil- very black, ocelli pale orange brown; dorsal surface slightly elevated between eyes, smooth; dorsal head sparingly clothed with recumbent pale pubescence, this pubes- cence slightly denser in areas behind eyes and laterad to ocelli, and flanking tylus; an- teocular length 0.30, equal to dorsal length of an eye; eye length 0.30, width 0.22; in- terocular space 0.60; ocelli small; buccula low, barely raised, gradually tapering in height without abrupt change in width to base of head; rostrum pale brown, apex dark, reaching to hind coxae, segment I nearly reaching base of head, segment lengths (from base) 0.50, 0.50, 0.50, 0.45; antennae long, slender, segments I and IV distinctly thicker than segments II and III, segment I pale brown, suffused with dark brown centrally, segment II dark brown, suffused with pale brown centrally, segment III dark brown, segment IV medium brown, extreme base of segment I and extreme dis- tal ends of segments I-III creamy white, all segments bearing short erect pale setae, these setae slightly shorter than the diame- ters of these segments, segment I with 3 959 longer setae near base, lengths of segments I-IV = 0,27,,0.55, 0.52; 0.60. Keneth 0.70; 1.37, uniform reddish brown, shin- ing, lacking pubescence; calli weakly raised, margins lacking setae; anterior collar Pronotum: maximum width and posterior lobe sparsely set with deep, dark brown punctures; lateral pronotal mar- gins weakly sinuate, not bulging strongly outward at lateral margins of calli. Scutellum: Length 0.55, maximum width 0.70, shining, lacking pubescence; overall coloration reddish brown, posterior half with a pale posteromedial stripe running along a narrow, raised, longitudinal medial carina; basal section smooth, lateral sec- tions deeply and coarsely punctate. Hemelytra: Convex and submacropter- ous, nearly coleopteriform, venation of co- rium obscure, membrane highly reduced; dorsal surface weakly rugose, shining, bear- ing scattered short, pale, semi-erect setae; coloration uniformly translucent reddish brown to orange brown, without contrasting markings; membrane of each hemelytron not completely overlapping with that of other when wings are at rest. Legs: Uniformly orange brown, except for a few small dark brown spots or mottled areas on posterior faces of middle and hind tibiae; all segments clothed with fine, pale, semi-recumbent setae. Lengths of leg seg- ments as follows: fore femur, tibia, tarsal I, tarsal II, tarsal III = 0.90, 0.87, 0.15, 0.07, 0.12; middle femur, tibia, tarsal I, tarsal I, tarsal [il —*1200;0.90," O7;, 0:073.:0:12; hind femur, tibia, tarsal I, tarsal II, tarsal II =" 1.20; 1.30; :0:25;.0,92,°0:15. Ventral surface: Dark brown to black, acetabula and peritreme yellowish white, evaporative area surrounding peritreme dull matte grey, adbominal ventrites VI—VIII marked with medium brown, genital seg- ment orange brown; ventrolateral portions of head and thorax and all of abdomen bearing numerous pale recumbent setae. Male paramere: Stout and set with spine-like setae basally, distal section form- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 960 ius) poepoe, female, dorsal habitus. SONYVS Nesius (Phy VOLUME 104, NUMBER 4 4 \ Fig. 4. paramere, dorsal view. Nesius (Physonysius) poepoe, male right ing a slender, elongate shaft, tip hooked and rounded (Fig. 4). Brachypterous female: Slightly larger than male, length 4.20; maximum _ width 2.00. Similar to male in general structure and coloration, but with end of terminal ab- domen bright red. Etymology.—The name _ “‘‘poepoe”’ comes from the Hawaiian word for “‘round- ed,” and refers to the overall body shape of this species. Material examined.—HOLOTY PE, sub- macropterous 2: HAWAIIAN ISLANDS, Maui Is., West Maui Mountains, moss forest nr. Puu Kukui summit, 1,750 m [5,740 ft], 20753) 39N, 156-35 22° W, 11 May 1992.Clz 8109, D. A. Polhemus (USNM). PARA- TYPES (all submacropterous): HAWAIIAN 961 ISLANDS, Maui Is.: 1 ¢, 1 2, West Maui Mountains, valley and ridge S. of Mt. Eke, Eke Natural Area Reserve, 1,310 m [4,297 ft], 20°55’04"N, 156°34'22”"W, 23-25 May 1997, CL 8291, D. A. Polhemus (USNM); 1 2, West Maui Mountains, Puu Kukui trail between Nakalalua and Violet Lake, 1,340— 1,490 m [4,395—4,487 ft], 3 March 1998, R. Takumi (BPBM); | 3, West Maui Moun- tains, Puu Kukui trail at Nakalalua, 1,350 m [4,430 ft], 20°54’55’N, 156°35'32’W, 22-23 May 1997, CL 8114, D. A. Polhemus (USNM). Discussion.—Usinger (= (1942) proposed Physonysius as a subgenus of Nesius to hold two very distinctive species character- ized by their rounded overall form, sub- macropterous hemelytra, and smooth, shin- ing dorsal surfaces: Nesius (Physonysius) molokaiensis trom eastern Molokai, and Nesius (Physonysius) ampliatus from Ha- leakala, eastern Maui. Nesius (Physonysius) poepoe may be easily separated from these two taxa by its smaller body size, longer rostrum, and shorter hemelytra, as given in the supplemental key provided below (re- vised from Usinger 1942). REVISED KEY TO SPECIES OF SUBGENUS PHYSONYSIUS Based on Holotypes 1. Rostrum long, extending to hind coxae; overall body length 4.6 mm or less; ground color dis- tinctly meddish:s West Maul os see. 254 2a > Axe Be, gry See oc tor schpen he oeeeeneds N. (P.) poepoe, n. sp. Rostrum shorter, extending only to middle cox- ae; overall body length exceeding 5.0 mm; ground color dark brown or black; East Maui andi MIOloKall ee tw. Ehechs Seen creeds aac heat 2 . General coloration uniform dark brown, with- NO out extensive pale markings; head uniformly dark brown dorsally, without a pale stripe along midline; embolium translucent reddish brown along entire length; scutellum with basal angles brown, not contrasting with central por- tion: Bast Maui (Ealeakala)iis- ia. eirc). -f.. 3 Heese konsike hap eee N. (P.) ampliatus Usinger — General coloration dark brown to black with scattered paler markings; head dark brown dor- sally but with a dark yellowish longitudinal midline stripe on vertex; embolium pale trans- lucent yellowish on basal two thirds; scutellum 962 Puu Kukui Trail Fig. 5. with basal angles pale yellowish, contrasting with dark central portion; Molokai ra Pat ae oNC RES Toy See N. (P.) molokaiensis Usinger Certain strongly brachypterous individu- als of Nesius (Nesius) kirkaldyi (Usinger), endemic to eastern Maui, can also approach Nesius (Physonysius) poepoe in size and shape, but are orange brown rather than reddish brown in overall coloration, have the embolar margin of the hemelytra less strongly expanded and arcuate, and have the overall hemelytral surface finely rugu- lose, rather than smooth and shining. These brachypterous forms of N. (Nesius) kirkal- dyi also retain a more completely developed wing membrane, so that the membranes of both hemelytra completely overlap when the wings are closed, while in N. (Physon- ysius) poepoe the wing membrane is greatly reduced, creating only a narrow amount of overlap, if any, between the two wing mem- branes in the closed position (Fig. 3). To PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Nesius (Physonysius) poepoe Distribution of Nesius (Physonysius) poepoe on Maui. date, no fully macropterous individuals of N. (Physonysius) poepoe have been collect- ed, while many such individuals of N. (Nes- ius) kirkaldyi are present in the Bishop Mu- seum collection. Physonysius clearly forms a distinct clade within the Hawaiian Nesius radiation, and could potentially be raised to full ge- neric status. This decision is complicated, however, by the fact that certain other Ha- waiian Nesius species exhibit morphologies intermediate between Physonysius and oth- er less highly modified Nesius species. In particular, the above-mentioned WN. (Nesius) kirkaldyi (Usinger), endemic to Maui, and Nesius (Trachynysius) whitei (Blackburn), from Hawaii island, display a range of var- iation in their wing morphs which in the most extreme states of submacroptery ap- proximate the arcuate-margined, nearly co- leopteriform condition seen in Physonysius. As a result, Usinger’s (1942) placement of VOLUME 104, NUMBER 4 Physonysius as a subgenus of Nesius is re- tained for the present pending a phyloge- netic analysis of the entire Hawaiian Nesius radiation. As currently understood, the distribution of Physonysius is limited to islands on the Maui Nui platform near the southeastern end of the Hawaiian Archipelago. With the collection of N. peopoe, species are now known from three of the four high volca- noes in this island group: Molokai, West Maui and Haleakala. Given this distribu- tion, it is probable that a member of the subgenus will also eventually be discovered on Lanai, the only other volcano on the Maui Nui platform that rises to an elevation sufficient to support wet upland forests fa- vored by Physonysius. AKNOWLEDGMENTS This revision is based primarily on ma- terial collected by the author during field surveys supported by the Smithsonian In- stitution’s Drake Fund (USNM), or held in the collections of the Bernice P. Bishop Mu- seum, Honolulu, Hawaii (BPBM). It could not have been completed without the gen- erous cooperation of the staff in the De- partment of Natural Sciences at the latter institution, particularly Scott E. Miller (now with the Smithsonian Institution), David Preston, Francis G. Howarth, and Gordon Nishida. I also thank James K. Liebherr of Cornell University and Curtis Ewing, from 963 the University of Hawaii, for companion- ship and assistance in the field; the staff of Pacific Helicopter Tours, Inc., who provid- ed access to remote field sites on Molokai and in the West Maui Mountains; Randy Bartlett and Hank Oppenheimer of the Maui Land and Pineapple Company, who kindly allowed access to Puu Kukui and surrounding ridge trails; and the Nature Conservancy of Hawaii (TNCH), who per- mitted collections in their Kamakou Pre- serve on eastern Molokai. This paper rep- resents contribution 2002-002 to the Hawaii Biological Survey. LITERATURE CITED Ashlock, P. D. 1966. New Hawaiian Orsillinae (He- miptera-Heteroptera: Lygaeidae). Pacific Insects 8(4): 805-825. . 1967. A generic classification of the Orsillinae of the world (Hemiptera-Heteroptera: Lygaeidae). University of California Publications in Entomol- ogy 48: vi + 82 pp. 1983. A new species, nomenclatural notes, and new records for Hawaiian Orsillinae (Hemip- tera: Heteroptera: Lygaeidae). International Jour- nal of Entomology 25(1): 42—46. Nishida, G. M. 1997. Hawaiian terrestrial arthropod checklist. Third edition. Bishop Museum Techni- cal Report 12: iv + 263 pp. Usinger, R. L. 1942. The genus Nysius and its allies in the Hawaiian Islands (Hemiptera, Lygaeidae, Orisillini). Bernice P. Bishop Museum Bulletin L/saatse loevepps, 12pl: Usinger, R. L. and P. D. Ashlock. 1958. Revision of the Metrargini (Hemiptera: Lygaeidae). Proceed- ings of the Hawaiian Entomological Society 17(1): 93-116. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 964-967 A NEW SPECIES OF PSEPHENOPS GROUVELLE (COLEOPTERA: PSEPHENIDAE) FROM MEXICO ROBERTO ARCE-PEREZ Instituto de Ecologia, A. C. Km. 2.5 Antigua Carretera a Coatepec No. 351, Congre- gacion El Haya, Apartado Postal arcerob @ ecologia.edu.mx) 63, 91070 Xalapa, Veracruz, Mexico (e-mail: Abstract.—Psephenops lupita, new species, is described from specimens collected in a stream running through a cloud forest at Coatepec (1,200 m), Veracruz State, Mexico. The total number of described species assigned to the genus is now eight. Resumen.—Se describe e illustra a Psephenops lupita, n. sp., con base en especimenes colectados en un arroyo de agua dulce de un bosque nebuloso en Coatepec (1,200 m snm), Veracruz, México, aumentando el numero total de especies descritas de este genero a ocho. Key Words: The genus Psephenops Grouvelle (= Xexanchorinus Grouvelle) is comprised of seven known species, with an apparently fragmentary distribution: P. smithi Grou- velle 1898 (Antillean); P. grouvellei Cham- pion 1913 (Guatemala); P. haitianus Dar- lington 1936 (Haiti); P. maculicollis Dar- lington 1936 (Colombia and Costa Rica); P. argentinensis Deléve 1967 (Argentina); P. prestonae Spangler 1990 (Costa Rica), and P. mexicanus Arce-Pérez and Novelo-Gu- tiérrez 2000 (Mexico). In this paper a new species is reported from Mexico. Ten specimens, all males, were collected in a stream located at Coatepec Village (1,200 m altitude), Municipality of Coate- pec; an additional specimen was collected in a stream located at Xico Viejo Village (1,800 m altitude), Municipality of Xico, Veracruz State, Mexico. Individuals were initially preserved in ethanol (80%) and some were later pinned; all collected ma- terial was examined. Male genitalia were extracted in ethanol, and potassium hydrox- Psephenidae, new species, Veracruz State, Mexico ide (KOH 5%) was used to clear genitalia to make illustrations. Microphotographs were taken with a scanning electron micro- scope JEOL, model JSM T20, with mag- nification of 200. Psephenops lupita Arce-Pérez, new species (Figs. 1—4) Holotype male.—Body oval, depressed; integument black, with reddish-yellow re- flections, completely covered with irides- cent, reddish-yellow, short setae; head, an- tenna, pronotum and scutellum black; ely- tron dark reddish brown, slightly elevated at humeral region; antennal segments 1—2 yellow; legs with coxae, trochanters, and femora yellow, tibiae and tarsi black; ven- trally reddish black, vestiture short, fine, and dense, grayish, with iridescent, yellow reflections; total length 3.80 mm, maximum humeral width 1.65 mm. Head: Wider than long, transverse. Clypeus subrectangular, declivous at an an- VOLUME 104, NUMBER 4 gle of less than 90 degrees from plane of frons, distal margin widely emarginate; fronto-clypeal surface with long, robust and stiff setae; vertex minutely punctate, with minute setae and a very shallow longitudi- nal furrow at middle. Antenna filiform, | 1- segmented, scape larger than other seg- ments, pedicel 0.57 length of scape and subglobose; third 0.64 length of scape; 7 flagellar segments small, apical segment acuminate. Eyes lateral, rounded, very prominent, dark reddish brown; postocular area amber yellow (Fig. 1). Labrum similar to clypeus but shorter; maxillary palpus 4- segmented, covered with long setae; seg- ment | smallest; segments 2 and 4 largest; relative proportion of segments: 0.30, 0.90, 0.60, 1.0, basal segments I—3 subconical; segment 4 oval, slightly depressed and end- ing in an inner subapical ridge. Labial pal- pus very short, 3-segmented; segments |—2 rounded and wide, segment 3 smallest, sub- conical and dark, segment 2 largest, robust (Bie..2): Thorax: Pronotum (Fig. 1) subtrapezo- idal (posterior margin 1.5 mm, lateral mar- gin 0.95 mm); anterior margin arcuate; pos- terior margin bisinuate and almost as wide as elytral base; lateral margins almost straight along anterior half and expanded laterally on posterior half; posterolateral an- gles subacute; all margins slightly thick- ened; pronotal disc with middorsal longi- tudinal carina weakly developed at distal half, and a wide, shallow, long depression at each side of carina; in lateral view, pro- notal disc (except for posterolateral expan- sions) convex; pronotal surface with punc- tation finer than that of head, and covered with short, cineraceous-yellow setae, and long dark reddish setae, setae shorter than those of head; prosternum (Fig. 2) black, short and slightly carinate; with long, con- cave, lanceolate, and slightly carinate po- stcoxal process, which reaches anterior half of mesosternum. Mesonotum with scutel- lum short and black, with tip broadly round; elytron dark reddish brown, with short yel- low setae and long reddish-yellow setae; 965 sides parallel on basal 0.75, then gradually converging posteriorly; base depressed on each side within humeri; surface without striae; length (from base to apex of suture) 2.65 mm; width (across humerus) 1.65 mm. Mesosternum, with wide groove for recep- tion of prosternal process. Metasternum bulky with medium longitudinal groove deep between matacoxae. Legs: Procoxa globose, mesocoxa coni- cal, metacoxa transverse; femora robust with dark yellowish setae; tibiae slender; metatibia as long as femur; protibia with distinct apical posterolateral denticle; tarsi, 5-segmented, tarsomeres | and 2 lobed ven- trally, with thick spongelike vestiture with short setae; lobe of tarsomere | extending distally to 0.30 of tarsomere 2; tarsomere 2 longer, extending distally and completely covering area of tarsomeres 3, 4, and 0.20 of tarsomere 5; proportional length of tar- someres: 0.40, 1.0, 0.25, 0.20, 0.85; a pair of long, curved, and slender apical claws present, each with small basal tooth. Abdomen: Vestiture short, fine and dense. Seven visible sterna in relaxed spec- imens; usually segment 6 hidden under seg- ment 5. Sternum 2 largest. Posterior mar- gins of sterna | and 2 deeply concave at middle, sinuate at sides. Posterior margins of sterna 3 and 4 straight, that of sternum 5 with a wide and shallow concavity at each side of midline. Sternum 6 very short, pos- terior margin widely concave. Pygidial plate narrow, oval, as long as sternum 5, and densely covered with long and stiff red- dish-yellow setae. Genitalia (Figs. 3—4): Trilobate, total length 0.50 mm. Parameres very wide and subsquare, with apices truncated (Fig. 3a), in dorsoapical view notched apically (Fig. 3b), broadly emarginate and curved inward in ventrolateral view (Fig. 4b); length 0.26 mm. Penis longer than parameres, in ventral view fingerlike, widened in basal half (Fig. 4a), length 0.30 mm, bearing a longitudinal lance-shaped sclerite along base which measures 0.18 mm in length; dorsal area of phallobase concave (Fig. 3a); ventral por- 966 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Psephenops lupita. 1, Details of head and pronotum showing mid-dorsal carina, dorsal view. 2, Detail of venter showing prosternal process. 3—4, Male genitalia. 3a, Dorsal view, showing phallobase and parameres. 3b, Microphotograph, dorsoapical view. 4a, Ventral view, showing long fingerlike penis widened in basal half. 4b, Microphotograph, ventrolateral view. VOLUME 104, NUMBER 4 tion bulky, subcylindrical; length 0.20 mm, width 0.20 mm (Figs. 4a, b). Female.—Unknown. (Apparently, fe- males of this group larger and bulkier than males, but less frequently collected.) Variation in paratype series.—Some specimens have reddish-brown elytra and in some the posterior margin of abdominal sternum 5 is not clearly convex at the mid- dle. Measurements (in mm): total length of body, 3.75—3.85; maximum humeral width 1.65—1.80; length of pronotum 0.70—0.95, basal width 1.45—1.55; elytral length 2.60— 2.70. Discussion.—Males of Psephenops lupi- ta may be distinguished from males of other species of the genus by the following com- bination of characters: Antenna filiform, el- ytra without conspicuous striae (moniliform in P. mexicanus and P. macutlicollis, elytra with striae); pronotum and elytra without tubercles or protuberances (pronotum and elytra with tubercles and protuberances in P. grouvellei); tarsomeres 1—2 lobed, pro- tibiae with denticle (tarsomeres 1-2-3 lobed, protibiae without denticle in P. pres- tonae); tarsomeres 1—2 lobed, elytra with- out conspicuous striae (promesotarsomeres 1-2 lobed, metatarsomere | lobed, elytra with striae in P. haitianus); antenna filiform 11-segmented, elytra without conspicuous Striae, penis fingerlike (antenna 8-segment- ed, elytra with striae, penis lanceolate in P. argentinensis). Apparently, Psephenops lu- pita 1s most similar to Psephenops mexi- canus, but the two may be distinguished by the following combination of characteristics (those of P. mexicanus in parentheses): To- tal length 3.75-3.85 mm (3.15—3.55 mm), maximum humeral length 1.65—1.80 mm (1.45—1.70 mm), antennae filiform (monil- iform), segments 1—2 yellow (black); legs with coxae, trochanters and femora yellow (black), parameres very wide and subsquare in dorsal view (slender and subtriangular), penis in ventral view fingerlike, widened in basal half (apical half subtriangular, with a small lateral toothlike projection). 967 Type material.—Holotype 6d labeled: ““MEXICO, Veracruz, Coatepec, Munici- pality Coatepec, 1,200 m, bosque mes6filo de montana, rio “‘Huehueyapan’’, 2-IV- 1999, R. Arce col.’’; Paratypes: same data as holotype (2 ¢); same locality 19-IV- 2000 (3 3); 26-IV-01 (3 3); and Xico Vie- jo, Municipality of Xico 26-VII-1998 (14). Holotype and paratypes deposited at Colec- ci6n Entomologica, Instituto de Ecologia, A.C., Xalapa, Veracruz, México. Etymology.—This species is dedicated to my beloved wife Guadalupe Gomez Men- diola; from spanish Guadalupe, diminutive ““Guadalupita, ““lupita.”’ Habitat.—Adult males of Psephenops lu- pita were captured in a stream with rapid water flow, on stones covered with moss and aquatic phanerogams. ACKNOWLEDGMENT I thank Dr. Harley P. Brown (University of Oklahoma) for his invaluable criticism on the final manuscript as well as for En- glish corrections. LITERATURE CITED Arce-Pérez, R. and R. Novelo-Gutiérrez. 2000. First record of the genus Psephenops (Coleoptera:Pse- phenidae) from Mexico, with a description of a new species. Entomological News 111(3): 196— 200. Champion, G. C. 1913. II. Notes on various Central American Coleoptera, with descriptions of new genera and species. Transactions of the Entomo- logical Society of London. 1913: 62-169. Darlington, P. J. 1936. A list of the West Indian Dry- opidae (Coleoptera) with a new genus and eight species, including one from Colombia. Psyche 43: 65-83. Deleve, J. 1967. Contribution 4 1 étude des Dryopo- idea (Coleoptera). XIX. Notes diverse et descrip- tions d’ especes nouvelles. Bulletin et Annales de la Société Entomologique Belgique 103: 414— 445. Grouvelle, A. H. 1898. Clavicornes de Grenada et de St. Vincent (Antilles) recoltés par M.H.H. Smith, et appartenant au Musée de Cambridge. Notes from the Leyden Museum 20: 35-48. Spangler, P. J. 1990. A new species and new record of the water-penny genus Psephenops (Coleoptera: Psephenidae) from Costa Rica. Entomological News 101(3): 137-139. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 968-979 DYSMICOCCUS FERRIS AND SIMILAR GENERA (HEMIPTERA: COCCOIDEA: PSEUDOCOCCIDAE) OF THE GULF STATE REGION INCLUDING A DESCRIPTION OF A NEW SPECIES AND NEW UNITED STATES RECORDS GARY L. MILLER AND DOUGLASS R. MILLER Systematic Entomology Laboratory, PSI. Agricultural Research Service, U.S. Depart- ment of Agriculture, Bldg. 005, Room 137, BARC-W, Beltsville, MD 20705, USA. (GLM e-mail: gmiller@sel.barc.usda.gov; DRM e-mail: dmiller@sel.barc.usda.gov) Abstract.—A key to the species of Dysmicoccus, Oracella, and Paradoxococcus of the Gulf State Region is presented. Dysmicoccus radinovskyi Miller and Miller, n. sp., is described and illustrated from the adult female, third-instar female, and second-instar female. Dysmicoccus merrilli (Ferris) is a new synonym of Dysmicoccus boninsis (Ku- wana). Dysmicoccus grassti (Leonardi), Dysmicoccus mackenziei Beardsley, and Dysmi- coccus neobrevipes Beardsley represent new distribution records. Key Words: mealybugs, Coccoidea, Pseudococcidae, Gulf State Region, new species, Dysmicoccus, Oracella, Paradoxococcus Ferris (1950) described Dysmicoccus with the intention of making Pseudococcus a natural group. It appears that characters chosen by Ferris may have been less than successful in this endeavor since some au- thors (Beardsley 1966, Miller and Mc- Kenzie 1971) have questioned the mono- phyly of Dysmicoccus. Dysmicoccus, as currently understood, 1s characterized by having the following combination of char- acters: lack oral-rim tubular ducts; have 5 or fewer conical setae in the cerarii; have auxiliary setae in cerarii anterior of the anal lobe pair; and have 17 or fewer pairs of ceraril. The genus currently includes 110 species worldwide with 39 species occur- ring in the Nearctic Region (Ben-Dov and German 200la, Ben-Dov, et al. 2001). Woody and herbaceous plants serve as hosts, including many grasses (Kosztarab 1996). Keys to species referable to the Ne- arctic Region have been provided by sev- eral authors including: Ferris (1950, 1953), McKenzie (1962, 1964, 1967), Miller and McKenzie (1973),Williams and Granara de Willink (1992), and Kosztarab (1996). In the United States, five species are of economic concern: Dysmicoccus boninsis (Kuwana) on sugar cane, Dysmicoccus vac- cinitt Miller and Polavarapu on blueberries, Dysmicoccus wistariae (Green) on yews, and Dysmicoccus brevipes (Cockerell) and Dysmicoccus grassii (Leonardi) on many hosts including several economic plants. Oracella is a monotypic genus contain- ing Oracella acuta (Lobdell). As with Dys- micoccus, Ferris (1950) erected Oracella with the intent of making Pseudococcus a more natural group. Oracella, as currently understood, is characterized by having the following combination of characters: lack of oral-rim tubular ducts, with 2 conical se- tae in the cerarii, with auxiliary setae in cer- aril anterior of the anal lobe pair, with dor- sal and ventral multilocular pores, with few oral-collar tubular ducts, and with less than VOLUME 104, NUMBER 4 17 pairs of cerarii. Although, there is some evidence that Oracella is congeneric with Dysmicoccus, we do not think it wise to consider them synonyms at this time, pend- ing an analysis of the monophyly of the ge- nus. However, we include Oracella here be- cause of its similarities to Dysmicoccus. Occurring primarily in southeastern United States (Lobdell 1930, Ben-Dov 1994, Kosz- tarab 1996) on Pinus spp., Oracella acuta can reach pest levels following repeated pesticide applications (Clarke et al. 1992). Oracella acuta has been accidently intro- duced into China where it is a serious pest (Kosztarab 1996). Paradoxococcus McKenzie is also a monotypic genus similar to Dysmicoccus with Paradoxococcus mcdanieli McKenzie (1962) as the type species. Paradoxococcus is characterized by having the following combination of characters: lack of oral-rim tubular ducts, with 2 conical setae in the ceraril, with auxiliary setae in cerarii ante- rior of the anal lobe pair, lack of multiloc- ular pores, presence of large oral-collar tu- bular ducts, and with less than 10 pairs of ceraril. The pupose of this paper is to provide a key to species of Dysmicoccus, Oracella, and Paradoxococcus of the Gulf State Re- gion. In addition, a new species description and synonymies are also presented. MATERIALS AND METHODS The Gulf State Region, as defined in this paper, encompasses the U.S. states that con- tact the Gulf of Mexico including: Ala- bama, Florida, Louisiana, Mississippi, and Texas. Terminology follows that of Wil- liams and Granara de Willink (1992) and Gimpel and Miller (1996) for adult females and immatures. Measurements were made with an ocular micrometer using a Nikon E600 compound microscope. Numbers and measurements of adult females are for 10 specimens and are given as an average fol- lowed by the range in parentheses. Holo- type measurements and numbers are record- ed separately in parentheses. Descriptions 969 of third-instar females and the second-instar female are based on six and one specimens, respectively. Associated enlargements of various structures on the illustrations are not proportional. Depositories of specimens are: The Natural History Museum (BMNH), London, United Kingdom; Flor- ida State Collection of Arthropods (FSCA), Gainesville, Florida; Museum National d’Histoire Naturelle (MNHN), Paris, France; University of California, Davis (UCD); National Museum of Natural His- tory (USNM), Coccoidea Collection, Belts- ville, Maryland. Information listed in the Specimens Examined section is verbatim from information recorded on the micro- scope slides. The abbreviation “‘ad.”’ refers to adult specimens and “‘sl.”’ refers to slide(s). RESULTS New Synonymy While examining specimens and descrip- tions of species to be included in the key, we discovered that Dysmicoccus merrilli (Ferris) and D. boninsis (Kuwana) were re- markably similar. Each species has: (1) Cer- arii that are restricted to the abdomen and have 1 or 2 on the head; (2) translucent pores that are present and abundant on the hind coxa and are absent or few on the tib- ia; and (3) oral-collar tubular ducts that are present over most of the dorsal surface, but are absent or greatly reduced in number on the first and eighth abdominal segments and on the metathorax. The only difference that we could find or mentioned in the literature was the shape of the circulus which is small and oval in D. merrilli and generally larger and more rectangular in many specimens of D. boninsis. However, after examining a long series of specimens it became apparent that D. boninsis occasionally has a small, oval circulus similar to the one in D. mer- rilli. The variable shape of the circulus on D. boninsis was illustrated by Williams and Watson (1988). Therefore, we consider D. merrilli (original combination 7Trionymus 970 merrilli Ferris 1953) as a junior, subjective synonym of D. boninsis (original combi- nation Dactylopius (Pseudococcus) bonin- sis Kuwana 1909), new synonymy. New UNITED STATES RECORDS OF INVASIVE SPECIES Through the courtesy of Avas B. Hamon, FSCA, we examined all Dysmicoccus spec- imens in the FSCA collection. Our exami- nation confirmed Hamon’s identifications and revealed two species for the Gulf State Region which are new records of distribu- tion. Dysmicoccus mackenziei Beardsley is known from Mexico, Central America, Ja- maica, and Italy (Ben-Dov and German 2001c). In the United States, it was previ- ously known only from California on Til- landsia punctulata Schlecht. & Cham. (Beardsley 1965), an ornamental flowering plant. The Florida material represents a new eastern United States distribution record. Collection data for D. mackenziei is as fol- lows: UNITED STATES: FLORIDA: M1i- ami, 14-I-1985, D. Barger, on Tillandsia sp., (5 ad. 2 on 5 sl.) FSCA; Tampa, Hills- borough, 28-XI-1988, C. Kamelhair, on 77/- landsia utriculata, (3 ad. 2 on 3 sl.) FSCA. Dysmicoccus neobrevipes Beardsley has been recorded from many countries in the Neotropical Region and is known from southern Asia and the Pacific (Ben-Dov and German 2001d). Dysmicoccus neobrevipes feeds on many hosts of economic impor- tance including various ornamental plants and food crops (Williams and Granara de Willink 1992). Originally described from Hawaii (Beardsley 1959), it has not been recorded from the continental United States until now. Therefore, this represents the first continental U.S. record. Collection data for D. neobrevipes is as follows: UNITED STATES: FLORIDA: Miami-Dade Co., Miami, 2-X-1978, P Chobrda, on (?) Fur- craea sp. (nr. Agave sp.), (3 ad. 2 on 3 sl.) FSCA; 22-VIII-1985, L. D. Howerton, on Yucca elephantipes [= Yucca guatemalen- sis], (4 ad. 2 on 4 sl.) FSCA; 9-X-1986, D. Leone et al., on Agave desmettiana, (4 ad. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2? on 4 sl.) FSCA; Orange Co., Apopka, 15- XI-1983, P. Gibson, on Yucca elephantipes [= Yucca guatemalensis], (2 ad. 2, 4 third- instar 2 on 6 sl.) FSCA; 16-II-1984, C. Phelps, on Yucca sp., (3 ad. 2 on 3 sl.) FSCA; 9-IV-1990, A. Capitano, on Agave angustifolia, (2 ad. 2 on 2 sl.) FSCA. Additionally, specimens identified as Dysmicoccus grassti (Leonardi) (= Dysmi- coccus alazon Williams) were also present in the USNM collection. Previously known from Nigeria, much of the Neotropical Re- gion, Sicily, and the Canary Islands (Ben- Dov and German 2001b), records of D. grassli represent new distribution data for the United States. Collection data for D. grassii are as follows: UNITED STATES: FLORIDA: Brevard Co., Cocoa Beach, 22- V-1970, J. MacGowan, on Coccoloba sp.., (1 ad. 2) USNM; Collier Co., Marco, 30- IV-1975, R. EK Denno, J. A. Davidson, and D. R. Miller, on Coccoloba sp., (1 ad. @) USNM; Dade Co., Key Biscayne, 12-I- 1966, S. Nakahara, on Coccoloba uvifera, (1 ad. 2) USNM; Cape Florida State Park, Crandon Pk., Key Biscayne, 8-XII-1970, S. Nakahara, on Coccoloba uvifera, (7 ad. & on 2 sl.) USNM; Key Biscayne, 4-IV-1974, R. EF Denno and D. R. Miller, on Coccoloba sp., (6 ad. 2 on 3 sl.) USNM; Matheson Hammock Pk., 4-[V-1974, R. EK Denno and D. R. Miller, on Rhizophora sp., (1 ad. 2) USNM; Lake just N. of Mahogony Ham- mock, 9-IV-1974, D. R. Miller and R. E Denno, on Persea borbonia, (1 ad. @) USNM; Adams Key, 24-II-1977, D. R. Miller, on Coccoloba diversifolia, (1 ad. 2) USNM;; Miami Lakes, 26-IV-1977, M. Cor- man, on Callistemon citrinus, (1 ad. 2) USNM;; Indian River Co., nr. Orchid, 8-V- 1975, R. E Denno, J. A. Davidson, and D. R. Miller, on Coccoloba uvifera, (3 ad. 2 on 2 sl.) USNM; Monroe Co., 19-VI-1969, D. Simberloff, on Rhizophora mangle, (3 ad. 2 on 3 sl.) USNM; Key Largo, 3 mi. N. Jct. Hghy 1 and 905, 5-IV-1974, R. EF Denno and D. R. Miller, on Ficus sp., C1 ad. 2) USNM; Key Largo nr. N. end, 8-I'V- 1974, R. EF Denno and D. R. Miller, on Lys- VOLUME 104, NUMBER 4 iloma_ latisiliquum, (7 ad. 2 on 4 si.) USNM;; Sarasota Co., Osprey, 17-V-1971, C. J. Bickner, on Codiaeum sp., (1 ad. 2) USNM. Material determined as Dysmicoccus bis- pinosus Beardsley is also present in the col- lections of FSCA and USNM. We suspect that current concepts of this species are too broad and anticipate that Florida specimens will eventually be discovered to be one or more different species. Therefore, this tax- on is not treated here. KEY TO SPECIES OF DysMICOCCUS ORACELLA, AND PARADOXOCOCCUS OF THE GULF STATES REGION (ADULT FEMALES) | Cincullusworesent.= sets 2 | ea win ere 2 ~ Circulustabsent)*aisiet tah. es 11 2) eee Wath iy/spains of Ceram: S426... ..). 3 - With less than 17 pairs of cerarll...... 8 3(2) Body setae normally not more elongate on dorsomedial area of abdominal segment 8 than on other areas of dorsum ........ 4 - Body setae on dorsomedial area of ab- dominal 8 conspicuously longer than on Othermareas Of dorsum. J. se. Pert roe eee Dysmicoccus brevipes (Cockerell) 4(3) — Dorsal body setae short, longest seta on dorsum of abdominal segment 8 less than LewidthOtwanalemme Fs ies. en lets 2 =) - Dorsal body setae long, longest seta on dorsum of abdominal segment 8 more than % width of anal ring ......... Dysmicoccus morrisoni (Hollinger) (in part) 5(4) Abdominal cerarii predominantly each Withe2conicalisetae ais. 958 Wis ee a ees 6 = Abdominal cerarii predominantly each with more than 2 conical setae ....... 7 6(5) Cluster of oral-collar tubular ducts present laterad of anterior spiracles; ventral ab- dominal multilocular pores present on seg- ments 4—8 (occasionally absent from seg- ment 4) .... Dysmicoccus grassii (Leonardi) — Cluster of oral-collar tubular ducts absent laterad of anterior spiracles; ventral ab- dominal multilocular pores present on seg- ments 6-8 .. Dysmicoccus texensis (Tinsley) 7(5) Hind coxa with translucent pores; opening of largest oral-collar tubular ducts approx- imately equal to diameter of multilocular pores .. Dysmicoccus mackenziei Beardsley - Hind coxa without translucent pores: opening of largest oral-collar tubular ducts less than diameter of multilocular pores eect aceed © Dysmicoccus neobrevipes Beardsley 8(2) a8) 10(9) 13(12) 14(11) 15(14) 971 Dorsal oral-collar ducts absent from ab- domen and thorax; translucent pores ab- SentetromiDindicoOxase eeu. nese 9 Dorsal oral-collar ducts present on abdo- men and thorax; translucent pores present Oumbimdicoxa™ Say eo = oasnewmenein, woes als Mey cl esse eens Dysmicoccus boninsis (Kuwana) Abdominal cerarii each predominantly with more that 2 conical setae ...... UAT NCR Roar Dysmicoccus difficilis (Lobdell) Abdominal cerarit each with 2 or fewer COMICAL SClACY Base sector < coe ere hee 10 Multilocular pores present on dorsum; dis- coidal pores absent near eyes ...... We he hc, Sod Sted oheia as Oracella acuta (Lobdell) Multilocular pores absent from dorsum; discoidal pores present near eyes .... Dysmicoccus morrisoni (Hollinger) (in part) Withy Ly (pats, Of Ceram sisi .0 ease re eye 12 With fewer than 17 pairs of cerarii ... 14 Oral-collar tubular ducts forming contin- uous ventral row across medial area of ab- dominal’sesmentS: 2.422834 an. a6 8: 13 Oral-collar tubular ducts on medial area of ventral abdominal segment 5 either entire- ly absent or not forming continuous row MALTA athe Sore Dysmicoccus lasii (Cockerell) Oral-collar tubular ducts forming contin- uous ventral row across medial area of ab- dominal segment 6; apical antennal seg- ment either partially or completely divid- ed; oral-collar tubular duct orifices smaller than diameter of trilocular pore ..... RE cea ema eee Dysmicoccus radinovskyi Miller and Miller, n. sp. Oral-collar tubular ducts on medial area of abdominal segment 6 either entirely ab- sent or not forming continuous row; apical antennal segment undivided; oral-collar tubular duct orifices larger than diameter Otmtmllocimanwpore: (lP ees ahs nee cea taes Se. Sh Ae ee Dysmicoccus diodium (McConnell) Multilocular pores absent; with less than SapPalnssOlyCChallilyette Voltas isis eye ake Peete oe Paradoxococcus mcdanieli McKenzie Multilocular pores present; with more than S palrsiof-ceranil) 4.2 25 4 Sse. i) Abdominal cerarii each predominantly with more that 2 conical setae; with 13 or feEWerpalrs Ol ,Ceratily wo: se fee a er ie ech eS eaa Dysmicoccus obesus (Lobdell) Abdominal cerarii each predominantly with 2 conical setae; with 15 or more pairs of cerarii ... Dysmicoccus milleri Kosztarab 972 Dysmicoccus radinovskyi Miller and Miller, new species (Figs. 1-3) Suggested Common Name: Dr. Rad’s Mealybug Holotype.—An adult female holotype is hereby designated with left label ““HOLO- TYPE Dysmicoccus radinovskyi Miller & Miller’; right label “Dade Co., Fla. Ever- glades Nat. Park 18 X 1972 W.H. Pierce on Psychotria undata”’|= Psychotria nervosa] (USNM). All specimens listed in the Spec- imens Examined section are paratypes ex- cept for the holotype. Etymology.—This species is named in honor and memory of Millersville Univer- sity of Pennsylvania biology professor, Dr. Syd Radinovsky, better known to his stu- dents as “Dr. Rad.” In his 34 years at Mill- ersville University, Dr. Rad’s enthusiasm for the biological sciences was unparalleled. His support for students was without equal. A field biologist in the true sense, Dr. Rad loved to talk about and visit unique habi- tats. It is fitting that the type localities of D. radinovskyi are from some of Dr. Rad’s be- loved unique habitats: South Florida, the Everglades, and the Florida Keys. ADULT FEMALE ig: 1) Field features.—Body white, covered with mealy wax. Mealybugs adhere to roots of Psychotria plants when pulled up; usu- ally found on roots of Psychotria growing as understory in hammocks along barrier is- lands of east central Florida (K. Hibbard, personal communication). Slide-mounted characters.—Body 1.08 (1.00—1.21) mm long (holotype 1.02 mm), 0.75(0.64—0.92) mm _ wide (holotype 0.80 mim) Dorsum with 17 pairs of cerarii (Fig. 1O); cerarii 1-11 and 13-14 with 2 conical setae; cerarii 12, 15, with 3(2—3) conical se- tae (holotype 3); cerarii 16-17 with 3 (3- 4) conical setae (holotype 3—4); all with auxiliary setae and associated trilocular PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pores. Anal-lobe cerarius (Fig. 11) with 3(1—4) auxiliary setae (holotype 3) (Fig. 1J), 2 conical setae (holotype 2) (Fig. 1K), 20(18—24) trilocular pores (holotype 18) (Fig. 1L), 2(1—4) discoidal pores (holotype 3) (Fig. 1M). Multilocular pores absent; tri- locular pores (Fig. IN) scattered nearly evenly over surface; discoidal pores pre- sent, about diameter of trilocular pore. Oral-collar tubular ducts absent. Longest submedial seta on segment VII 18(13—25) w long (holotype 25 2); 4 submedial setae on segment VIII, longest seta 19(13—28) w long (holotype 28 w). Anal-ring seta 100(88—113) w long (ho- lotype 100 w); 1.5(1.2—1.6) (holotype 1.6) times as long as width of anal ring. Venter with multilocular pores (Fig. 1G) usually in posterior and anterior bands on segments VI-VII and restricted to posterior band on segment VIII, occasionally with | or 2 pores on posterior margin of segments V: occasionally with | multilocular pore near base of pro- and mesocoxa. Trilocular pores scattered throughout. Discoidal pores common, of same size as on dorsum. Oral- collar tubular ducts of | size (Fig. IF), or- ifice smaller than diameter of trilocular pore, present in medial and mediolateral ar- eas of abdominal segments II-VII, few ducts near coxae. Setae as follows: 4 cisanal (Fig. 1H), longest 39(30—45) pw long (ho- lotype 45 wp); longest anal-lobe seta 119(105—130) pw long (holotype 113 wp); longest seta on hind trochanter 82(63—88) w long (holotype 78 wp). Circulus absent. Labium 129(113—143) wp long (holotype 133 p). Antennae 7—8 seg- mented with apical segment partially or ful- ly subdivided (Fig. 1A), 273(263—288) pw long (holotype 263 p). Eye with associated discoidal pore (Fig. 1B). Legs with trans- lucent pores restricted to hind femur and tibia, ventral surface (when leg is lying flat as shown in illustration) without pores, dor- sal surface of femur (Fig. 1D) with 18(12— 22) pores (holotype 12), dorsal surface of tibia (Fig. 1D) with 15(12—18) pores (ho- lotype 12). Hind femur 137(130—-142) p VOLUME 104, NUMBER 4 973 Milly ©2007 % Fig. 1. Adult female of Dysmicoccus radinovskyi. A, Antenna and detail of terminal segment; B, Discoidal pore near eye; C, Tarsal claw and digitules of fore leg; D, Dorsal surface translucent pores of femur and tibia; E, Detail of tarsal claw and digitules of hind leg; EK Oral-collar tubular duct; G, Multilocular pore; H, Cisanal seta; I, Anal-lobe cerarius; J, Auxiliary seta; K, Cerarian seta; L, Trilocular pore; M,. Discoidal pore; N, Trilocular pore; O, Cerarius. 974 long (holotype 130 w); hind tibia 121(113— 138) p long (holotype 115 w); hind tarsus 81(75—88) pw long (holotype 80 w). Tibia/ tarsus 1.5(1.3-1.7) (holotype 1.4); femur/ tarsus 1.7(1.6—1.8) (holotype 1.6). Length of hind femur divided by greatest width of hind femur 2.4(2.1—2.9) (holotype 2.9). Hind tibia with 14(12—16) setae (holotype 14). Claw digitules on all legs clubbed, ap- proximately same size. Tarsal digitules on hind 2 pairs of legs clubbed, each tarsus with | digitule noticeably longer and with club slightly larger than other (Fig. 1E); tar- sal digitules on front pair of legs of 2 dif- ferent sizes and shapes, | digitule clubbed and robust, other digitule without club, slender (Figyl@): Notes.—The above description is based on 124 specimens from 17 localities. Dys- micoccus radinovskyi is unusual with the terminal antennal segment being either en- tirely or partially subdivided, the circulus is absent, and the oral-collar tubular ducts are of 1 size. This species is close to D. diod- ium but differs by the diameter of the oral- collar tubular ducts which are smaller than the diameter of trilocular pores (D. diodium has oral-collar tubular ducts larger than the diameter of trilocular pores), the oral-collar tubular ducts forming a continuous row across the medial area of ventral abdominal segment 6 (D. diodium either lacks oral-col- lar tubular ducts or only has a few scattered oral-collar tubular ducts in the medial area of ventral abdominal segment 6), and the subdivided terminal antennal segment (D. diodium does not have a subdivided termi- nal antennal segment). The adult female can be distinguished from the third and second instars by having multilocular pores, translucent pores on hind femur and tibia, and a vulva. THIRD-INSTAR FEMALE (Fig: 2) Slide-mounted characters —Body 0.74 (0.69—0.80) mm long, 0.43(0.40—0.48) mm wide. Dorsum with 17 pairs of cerarii; cerari PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1-11, 13-14, and 16 with 2 conical setae; cerarii 12, 15, 17 with 3(2—3) conical setae; all with auxiliary setae. Anal-lobe cerarius with 1(1—3) auxiliary setae, 2 conical setae, 15(13—17) trilocular pores, | discoidal pore. Multilocular pores absent; trilocular pores scattered nearly evenly over surface; dis- coidal pores present, about % diameter of trilocular pore. Oral-collar tubular ducts ab- sent. Longest submedial seta on segment VIT 13(10—-15) pw long; 2 submedial setae on segment VIII, longest seta 13(10—15) p long. Anal-ring seta 87(78—100) wp long; 1.6(1.5—1.8) times as long as width of anal ring. Venter without multilocular pores; triloc- ular pores scattered throughout. Discoidal pores scarce, of same size as on dorsum. Oral-collar tubular ducts uncommon, one size, present anterior to clypeus and lateral mesothorax. Setae as follows: 4 cisanal, longest 25(15—35) pw long; longest anal-lobe seta 95(92—98) pw long; longest seta on hind trochanter 67(66—70) w long. Circulus absent. Labium 108(98—113) w long. Antennae 6-segmented, 227(213—237) w long. Legs without translucent pores. Hind femur 111(105—118) p long; hind tib- ia 88(83—93) » long; hind tarsus 77(75—80) w long. Tibia/tarsus 1.1(1.1—1.2); femur/tar- sus 1.4(1.4—1.5). Length of hind femur di- vided by greatest width of hind femur 2.8(2.6—3.0). Hind tibia with 9(9—10) setae. Claw and tarsal digitules same as adult fe- male. Notes.—The above description is based on 5 specimens from 3 localities. The third- instar female can be distinguished from all other instars by its 6-segmented antennae, absence of mutilocular pores, and 13-17 tri- locular pores on anal-lobe cerarii. SECOND-INSTAR FEMALE (Fig. 3) Slide-mounted characters.—Body 0.56 mm long, 0.31 mm wide. Dorsum with 17 pairs of cerarii although some cerariian setae are not distinct from 975 VOLUME 104, NUMBER 4 —e spe ere a co \ ee | ae aOR eee Third-instar female of Dysmicoccus radinovskyi. Fig. 2. 976 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Second-instar female of Dysmicoccus radinovskyi. VOLUME 104, NUMBER 4 auxiliary setae; cerarii 1-7, with 2 distinct conical setae; cerarius 17 with 3 conical se- tae; all with auxiliary setae. Anal-lobe cer- arius with | auxiliary seta, 2 conical setae, 7 trilocular pores, | discoidal pore. Multi- locular pores absent; trilocular pores scat- tered nearly evenly over surface; discoidal pores about diameter of trilocular pore. Oral-collar tubular ducts absent. Longest submedial seta on segment VII 5 p long; submedial setae absent on segment VIII. Anal-ring seta 63 w long; 1.6 times as long as width of anal ring. Venter without multilocular pores; triloc- ular pores scattered throughout. Discoidal pores scarce, of same size as on dorsum. Oral-collar tubular ducts uncommon, one size, present lateral to clypeus and lateral mesothorax. Setae as follows: 4 cisanal, longest 23 p long; longest anal-lobe seta 80 w. long; longest seta on hind trochanter 50 ww long. Circulus absent. Labium 75 pw long. An- tennae 6-segmented, 175 wp long. Legs with- out translucent pores. Hind femur 80 w long; hind tibia 63 p long; hind tarsus 63 w long. Tibia/tarsus 1.0; femur/tarsus 1.3. Length of hind femur divided by greatest width of hind femur 2.6. Hind tibia with 9 setae. Claw and tarsal digitules same as adult female. Notes.—The above description is based on | specimen from | locality. The second- instar female can be distinguished from all other instars by its 6-segmented antennae, absence of multilocular pores, and 5 triloc- ular pores on anal-lobe ceraril. SPECIMENS EXAMINED Paratypes—UNITED STATES: FLORI- DA: Brevard Co.: Florida Beach, S. side of Mullett Creek Rd., 26-II-2001, K. Hibbard, on Psychotria nervosa (28 ad. 2, 1 second- instar 2 on 6 sl.) BMNH, MNHN, UCD, USNM;; Broward Co.: Ft. Lauderdale, 4-X- 1982, M. McDonald, on Psychotria nervosa (3 ad. 2 on 3 sl.) FSCA; 3-XI-1987, T. Phillips, on Psychotria sp. (2 ad. @ on 2 sl) MES CA; -Charlotte"Co::-3: im. EE Punta 977 Gorda, 21-XI-1980, Z. Smith, on Psycho- tria sp.[?] (3 ad. 2 on 3 sl.) FSCA; Collier Co.: Collier Seminole State Pk., Juct. of 92 and 41, 11-IV-1974, D. R. Miller and R. FE Denno, on Psychotria sp. (6 ad. 2, 2 3 9 on 4 sl.) FSCA, USNM; Miami-Dade Co.: Miami, 31-VIII-1972, P. Pullara, on Psy- chotria nervosa (1 ad. 2) FSCA; Ever- glades Nat. Park, 18-X-1972, W. H. Pierce, on Psychotria nervosa (3 ad. 2, 1 3% 2 on 4 sl.) FSCA, USNM; Miami, 29-IX-1972, W. H. Pierce, on Psychotria nervosa (2 ad. 2 on 2 sl.) FSCA,USNM; 31-[X-1972, W. H. Pierce, on Psychotria nervosa (1 ad. 2) FSCA; on coast east of Perrina, 4-I[V-1974, R. FE Denno and D. R. Miller, on Psychotria nervosa (9 ad. 2 on 4 sl.) FSCA,USNM;; lake just N. of Mahogany Hammock, 9-IV- 1974, D. R. Miller and R. E Denno, on crinkle leaf—Psychotria sp. (7 ad. 2 on 4 sl.) FSCA, USNM; N. Miami Beach, 5-VI- 1986, D. Gruber and LL. Davis, (ad: 2on 3 sl.) FSCA; Indian River Co.: nr. Orchid, 8-V-1975, R. EF Denno, J. A. Davidson, and D. R. Miller, on Psychotria sp. (4 ad. 2 on 2 Sl.) "USNM=: Orchid, 137 mi. N.. of \S.R. 510 on N. Jungle Tr., 23-I-2001, K. Hib- bard, on Psychotria nervosa (42 ad. @, 2 3 2 on 7 sl.) USMN; Manatee Co.: Ellen- ton, 15-I-1980, J. Felty and Z. Smith, on Psychotria nervosa (3 ad. “2 ‘on 3 sl.) FSCA, USNM; Monroe Co.: Key Largo, 8- IV-1974, D. R. Miller and R. E Denno, on Psychotrias-sp- "(ss tads 2° on 4e'sis) FSCA,USNM;; 25 mi. west of 41 intersec- tion of 94, 10-IV-1974, R. E Denno and D. R. Miller, on Psychotria sp. (3 ad. 2 on 3 sl.) FSCA,USNM; Palm Beach Co.: West Palm Beach, 16-I-1967, N. Bezona, on Psy- chotria nervosa (2 ad. 2 on 2 sl.) FSCA; St Eucite. Co Port st. Joucie, 25=X-1982, K. Hibbard, on Psychotria nervosa (2 ad. 2 on 2 sl.) FSCA; White City, 7-X-1982, K. Hibbard, on Psychotria nervosa (2 ad. eon 2 sh) ESCA: CONCLUSIONS With the description of D. radinovskyi, a total of 12 species of the genus are known 978 from the Gulf State Region of the United States. Newly published records of D. gras- sii, D. mackenzii and D. neobrevipes alert economic entomologists to the presence of three new pest species either in Florida or the continental United States. ACKNOWLEDGMENTS We are especially grateful to K. Hibbard, Florida Dept. of Agriculture, for collecting additional specimens of D. radinovskyi and his observations regarding the field features of this insect. We appreciate manuscript suggestions and comments of A. Hamon (Florida Dept. of Agriculture and Consumer Services), M. B. Stoetzel (USDA, ARS, Systematic Entomology Laboratory, Belts- ville, MD), M. Williams (Dept. of Ento- mology and Plant Pathology), Auburn Uni- versity, Auburn, AL), and N. Woodley (USDA, ARS, Systematic Entomology Laboratory, Washington, DC). LITERATURE CITED Beardsley, J. W. 1959 (1958). On taxonomy of pine- apple mealybugs in Hawaii, with a description of a previously unnamed species (Homoptera: Pseu- dococcidae). Proceedings of the Hawaiian Ento- mological Society 17: 29-37. . 1965. Notes on the pineapple mealybug com- plex, with descriptions of two new species (Ho- moptera: Pseudococcidae). Proceedings of the Ha- waiian Entomological Society 19: 55—68. . 1966. Insects of Micronesia. Homoptera: Coc- coidea. Insects of Micronesia 6: 377-562. Ben-Dov, Y. 1994. A systematic catalogue of the mealybugs of the world (Insecta: Homoptera: Coccoidea: Pseudococcidae and Putoidae) with data on geographical distribution, host plants, bi- ology and economic importance. Intercept Limit- ed, Andover, UK. 686 pp. Ben-Dovy, Y. and V. German. 2001a. ScaleNet. Scales in a Family/Genus Query [Web Page] http://www. sel.barc.usda.gov/scalecgi/chklist.exe?Family = Pseudococcidae& genus=dysmicoccus [accessed 23 October 2001). . 2001b. ScaleNet. Valid Name and Catalogue Query Results. Dysmicoccus grassii. [Web Page] http://www.sel.barc.usda.gov/catalogs/pseudoco/ Dysmicoccusgrassii.htm [accessed 1 November 2001]. . 2001c. ScaleNet. Valid Name and Catalogue Query Results. Dysmicoccus mackenziei. [Web Page] http://www.sel.barc.usda.gov/catalogs/ PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pseudoco/Dysmicoccusmackenziei.htm [accessed 1 November 2001]. . 2001d. ScaleNet. Valid Name and Catalogue Query Results. Dysmicoccus neobrevipes. [Web Page] http://www.sel.barc.usda.gov/catalogs/ pseudoco/Dysmicoccusneobrevipes.htm [accessed 1 November 2001]. Ben-Dov, Y., Miller, D. R., and G. A. P. Gibson. 2001. ScaleNet. Scales in a Region/ Country/ Subunit Que- ry [Web Page] http://www.sel.barc.usda. gov/scalecgi/ region.exe?region= N&tamily = Pseudococcidae& intro= A & genus=dysmicoccus&country = &subunit = ®name= &ctryname= &action=Submit + Query&querytype=Region+ Query [accessed 23 October 2001]. Clarke, S. R., J. E Negron, and G. L. Debarr. 1992. Effects of 4 pyrethroids on scale insect (Homop- tera) populations and their natural enemies in lob- lolly and shortleaf pine seed orchards. Journal of Economic Entomology 85(4): 1246-1252. Ferris, G. F 1950. Atlas of the Scale Insects of North America. (ser. 5) [v. 5]. The Pseudococcidae (Part I). Stanford University Press, Palo Alto, Califor- nia. 278 pp. 1953. Atlas of the Scale Insects of North America, v. 6, The Pseudococcidae (Part IL). Stan- ford University Press, Palo Alto, California. 506 PP- Gimpel, W. F and Miller, D. R. 1996. Systematic anal- ysis of the mealybugs in the Pseuwdococcus mari- timus complex (Homoptera: Pseudococcidae). Contributions on Entomology, International 2: 1— 163. Kosztarab, M. 1996. Scale Insects of Northeastern North America. Identification, Biology, and Dis- tribution. Virginia Museum of Natural History, Martinsburg, Virginia. 650 pp Kuwana, S. I. 1909. Coccidae of Japan (IV). A list of Coccidae from the Bonin Islands (Ogasawaraji- ma), Japan. Journal of the New York Entomolog- ical Society 17: 158-164. Lobdell, G. H. 1930. Twelve new mealybugs from Mississippi. (Homoptera: Coccoidea). Annals of the Entomological Society of America 23: 209— 2306. McKenzie, H. L. 1962. Third taxonomic study of Cal- ifornia mealybugs, including additional species from North and South America (Homoptera: Coc- coidea: Pseudococcidae). Hilgardia 32: 637-688. . 1964. Fourth taxonomic study of California mealybugs, with additional species from North America, South America and Japan (Homoptera: Coccoidea: Pseudococcidae). Hilgardia 35: 211— DY Xe . 1967. Mealybugs of California with taxono- my, biology, and control of North America species (Homoptera: Cooccoidea: Pseudococcidae). Uni- versity of California Press, Berkeley. 526 pp. VOLUME 104, NUMBER 4 Miller, D. R. and McKenzie, H. L. 1971. Sixth taxo- nomic study of North American mealybugs, with additional species from South America (Homop- tera: Coccoidea: Pseudococcidae). Hilgardia 40: 565-602. 1973. Seventh taxonomic study of North American mealybugs (Homoptera: Coccoidea: Pseudococcidae). Hilgardia 41: 489-542. 979 Williams, D. J. and Watson, G. W. 1988. The scale insects of the tropical South Pacific region. Pt. 2: The mealybugs (Pseudococcidae). CAB In- ternational Institute of Entomology, London. 260 pp. Williams, D. J. and Granara de Willink, M. C. 1992. Mealybugs of Central and South America. CAB International, London, England. 635 pp. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 980-992 THE CERACANTHIA COMPLEX (LEPIDOPTERA: PYRALIDAE: PHYCITINAE) IN COSTA RICA. II. MEGARTHRIA RAGONOT, DRESCOMA DYAR, AND LASCELINA HEINRICH H. H. NEUNZIG AND M. A. SOLIS (HHN) Department of Entomology, North Carolina State University, Raleigh, NC 27695-7613, U.S.A.; (MAS) Systematic Entomology Laboratory, PSI, Agricultural Re- search Service, U.S. Department of Agriculture, % National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: asolis@ sel.barc.usda.gov) Abstract.—The genera Megarthria Ragonot, Drescoma Dyar, and Lascelina Heinrich are redescribed. Amegarthria Neunzig and Dow and Megacerdresa Neunzig are newly synonymized with Megarthria and Lascelina, respectively. One species of Megarthria, two species of Drescoma, and three species of Lascelina are recognized in Costa Rica. Two species are described as new: Lascelina papillina and Lascelina pitilla. Megacer- dresa cordobensis Neunzig is given the new combination Lascelina cordobensis (Neun- zig). Megarthria cervicalis Dyar is a new junior synonym of Megarthria peterseni (Zell- er). Keys are provided for males of Drescoma and Lascelina. Habitus photographs of male adults, line drawings of most male antennae and male genitalia of all species, are included. Also figured are the female genitalia of four of the six species, the forewing and hindwing of Drescoma cyrdipsa, and the costa of the forewing of Megarthria petersent. Key Words: Phycitinae, taxonomy, Neotropical This second paper treating the Ceracan- thia complex, a common, mostly Neotrop- ical group within the subfamily Phycitinae in Costa Rica, deals with the genera Me- garthria Ragonot, Dyar, and Lascelina Heinrich. In Part I (Neunzig and Solis 2002), we revised Ceracanthia Ra- gonot, the largest genus in the complex. In- formation relative to the contributions of Drescoma early workers, an overview of the salient diagnostic features of members of the com- plex, and a key to genera can be found in Part I. Specimens studied for Part If came from the following sources: Instituto Na- cional de Biodiversidad, Santo Domingo, Costa Rica (INBio); North Carolina State University Entomology Museum, Raleigh, N. C., U.S.A. (NCSU); National Museum of Natural History, Smithsonian Institution, Washington, D. C., U.S.A. (USNM); The Natural History Museum, London, England (BMNH); and the collection of Vitor O. Becker, Brasilia, Brazil (VOB). Megarthria Ragonot Megarthria Ragonot 1893:156. Type spe- cies: Myelois peterseni Zeller 1881. Orig- inal designation. Amegarthria Neunzig and Dow 1993:42. New synonymy. Type species: Megar- thria cervicalis Dyar 1919, here consid- ered a synonym of Myelois peterseni Zeller 1881. Original designation. Note.—The genus Amegarthria was pro- posed by Neunzig and Dow (1993) because of major differences they found between VOLUME 104, NUMBER 4 Megarthria cervicalis Dyar and other spe- cies placed by Heinrich (1956) in his con- cept of Megarthria. However, our present study, which included an examination of Zeller’s types in the BMNH and a more thorough review of the literature, has shown that Heinrich misidentified the type species of Ragonot’s Megarthria. Heinrich errone- ously selected a species belonging to the related genus Ceracanthia Ragonot (addi- tional information regarding this lapsus by Heinrich is given herein under “‘Remarks”’ following the treatment of Megarthria pe- terseni). The actual type species of Megar- thria is identical to the type species of Ame- garthria, and, therefore, Amegarthria is a junior synonym of Megarthria. Description.—Antenna of male with scape large (about 3 as wide as width of base of shaft); shaft strongly bent outward near its attachment to scape forming a short, strongly curved sinus; sinus without spines marking its basal and upper limits; inner surface of sinus covered with ap- pressed scales; very small triangular tuft of scales attached to posterior base of sinus; sensilla trichodea (cilia) of antenna abun- dant, about as long as width of shaft at mid- sinus. Antenna of female simple. Labial palpus of male upturned, extending above vertex; 3™ segment about as long as 2™ seg- ment. Maxillary palpus of both sexes small, short scaled. Haustellum well developed. Ocellus present. Forewing of male with pe- culiar, small, hood-shaped, basal, costal prominance (Fig. 25), and associated costal fold; underside of wing with subcostal streak of brownish-red scales; forewing of both sexes smooth, with I1 veins; R,,, and R; stalked for about % their lengths, M, straight; M, and M, separate; CuA, arises from lower angle of cell; CuA, arising well before lower angle of cell. Hindwing with eight veins (1A, 2A, 3A together treated as one vein); Sc+R, and Rs fused for less than Y% their lengths beyond cell; CuA, short stalked to M, and M,; CuA, arises from just before outer angle of cell; cell less than % length of wing. Male abdominal segment 8 981 with small tuft, consisting of broadly flat- tened and contorted scales (digitate pocket from sternite of segment 8 strongly bent and flattened). Male genitalia with uncus small relative to other genitalic structures; gnathos with weakly developed medial, hooklike element (fused with scaphium ex- cept for broadly swollen apex); transtilla present, consisting of a thin rectangular plate; juxta platelike with slender, seta-bear- ing lateral arms; valva broad, covered on inner surface with slender setae (without broadened spinelike setae on distal half); sacculus with apex a small, slender hooked, fingerlike process lying above a_ broad squared-off element, base of sacculus with three pair of ventral scale tufts (largest pair consisting of broad, tightly-coiled scales); aedoeagus long and slender, vesica of ae- doeagus with small, semi-oval, sclerotized plate and patch of microspines; vinculum broad (about as long as greatest width) and evenly rounded posteriorly. Female genita- lia with ductus bursae long, slender, twist- ed, with microspines within lower %; cor- pus bursae short, oval, slightly less than % as long as ductus bursae; microspines, sim- ilar to those in ductus bursae, extending for a short distance into corpus bursae; ductus seminalis attached to corpus bursae near junction of ductus bursae and corpus bur- sae. Megarthria peterseni (Zeller) (Figs. 1, 7, 12—13, 25-26) Myelois peterseni Zeller 1881:198. Megarthria cervicalis Dyar 1919:42. New synonymy. Megarthria beta Heinrich 1956:88. Identifying features can be seen easily in the unusual male antenna (Fig. 7) (in which the shaft is strongly bent outwardly near its attachment to a large, broad scape), the pe- culiar, small hood-shaped, costal promi- nence at the base of the forewing (Fig. 25), the coiled scales at the base of the sacculus (Fig. 12), and the twisted, long (about 2X length of corpus bursae) ductus bursae (Fig. Figs. 1-6. Lascelina cordobensis. 5, L. pitilla. 6, L. papillina. 26). Additional diagnostic features are in- cluded under the description of the genus. Costa Rican material examined.—San José, (no date on label), H. Schmidt (1 @); Rancho Quemado, 200 m., Peninsula de Osa, Provincia Puntarenas, October, 1992, FE Quesada, INBio CR 1000945014 (1 ¢). Other material examined.—Cuba, Tana- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 6 Males. dorsal view. 1, Megarthria peterseni. 2, Drescoma cyrdipsa. 3, Drescoma cinilixa. 4, mo, August (1 ¢; holotype of Megarthria cervicalis Dyar in USNM); Mexico, Jalapa, (no date on label), W. Schaus (1 @), Ori- zaba, March (no year on label), R. Muller (1 2: holotype of Megarthria beta Heinrich in USNM); Belize, 1000 Foot Falls, Moun- tain Pine Ridge, May, June, 1990, L. C. Dow (4 6, 3 &); Guatemala, February, VOLUME 104, NUMBER 4 Figs. 7-11. 9, Lascelina cordobensis. 10, L. pitilla. 11, L. papillina. May, October, W. Schaus & W. Barnes (1 6, 2 2); Panama, Porto Bello, May, 1912, A. Buseck: @ 2.) ILas Cumbres... 1971. M. Daykin (1 6); Columbia, Honda, (no date on label), Petersen (1 d; lectotype of Mye- lois peterseni Zeller in BMNH); Trinidad, Caparo, (no date on label), E Birch (1 @); Brazil, Rond6nia, Porto Velho, April, May, 1OSO. VO} Becker (@ ¢., 1-2): Remarks.—Heinrich (1956), apparently, did not look at the syntypes of Myelois pe- terseni. He stated “I have seen no speci- mens from Colombia” (the type locality). He, also, did not carefully read the descrip- tion of peterseni in Ragonot’s 1893 publi- cation. Heinrich’s description and figures purported to be Megarthria peterseni are of a species described as new in the genus Ceracanthia in our Part I (Neunzig and So- lis 2002). Particularly unfortunate was Heinrich’s erroneous statement in reference to the specimens upon which he based his description and figures, that “‘the males be- fore me ... agree in antennal characters with the type of peterseni as described by Ragonot.”” Heinrich described, and his fig- ure 279 illustrated, a male antenna with 983 11 ss a =P Os = I 10 Basal part of right male antenna, frontal view. 7, Megarthria peterseni. 8, Drescoma cyrdipsa. prominent spines associated with a sinus in the shaft. In actuality, no mention was made by Ragonot (1893) of a spined sinus. Ra- gonot stated that the distinctive feature of peterseni 1s the large scape that is swollen distally (“... l'article basilaire, qui est long, tres gros, dilaté au sommet.” (pp. 156—157)). Indeed, Ragonot based his ge- nus name, Megarthria (arge-joint) on this feature of the scape. It also should be pointed out that the original description of peterseni by Zeller (1881) did not mention spines on the anten- na, although he included similar features in his 1881 descriptions of the antennae of other new species having these structures. In addition, Zeller’s fig. 20 of the forewing of peterseni agrees more closely with Fig. 1 of the present paper than with fig. 2 in Part I of our study. HHN has examined Zeller’s syntypes of Myelois peterseni in the BMNH. There are two males and five females each with a green label on which is written ““Myelois peterseni Z. Honda pet.” and a white label with “‘Zell. Coll. 1884.” A male that agrees with Zeller’s and Ragonot’s description is 984 here selected as lectotype in order to fix the identity of Megarthria peterseni. A red la- bel with “LECTOTYPE peterseni Zeller, Neunzig and Solis” has been added to this specimen. Zeller’s other male syntype in the BMNH is a specimen of Drescoma cyrdip- sa Dyar. The holotype of Megarthria cervicalis Dyar in the USNM is the same species as the newly designated lectotype of Myelois peterseni Zeller, and, therefore, Megarthria cervicalis has been made a junior synonym of Megarthria peterseni. Megarthria beta Heinrich was synonymized with cervicalis by Neunzig and Dow in 1993. Drescoma Dyar Drescoma Dyar 1914:328. Type species: Drescoma cyrdipsa Dyar 1914. Original designation. Description.—Antenna of male simple (with slender scape and without sinus, spines, and/or scale tufts); sensilla trichodea (cilia) abundant, about % as long as width of shaft near base. Antenna of female sim- ple. Labial palpus of male upturned, reach- ing above vertex, 3rd segment almost as long as 2nd segment. Maxillary palpus small, short-scaled. Haustellum well devel- oped. Ocellus present. Forewing of male with costal concavity in basal half of wing: forewing of both sexes smooth, with 11 veins; R,;,, stalked for over % their length; M, straight; M, and M, separate; CuA, aris- es from lower angle of cell; CuA, arising before lower angle of cell. Hindwing of male with costal concavity in basal % of wing; underside with subcostal patches or streaks of contrastingly colored scales and with anal fold and associated scale pencil: hindwing of both sexes with eight veins (1A, 2A, and 3A together treated as one vein); Sc+R, and Rs fused for less than 2 their lengths beyond cell; M, slightly con- cave; stalk of Sc+R,+Rs+M, angularly bent just beyond base; M, and M, fused for over % their lengths beyond cell; CuA, short stalked to M, and M,; CuA, arises PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from lower angle of cell; cell about % length of wing. Male abdominal segment 8 without tuft (digitate pocket from sternite of segment 8 simple). Male genitalia with un- cus broadly rounded apically; gnathos with distomedial elements developed into a rod- like structure; transtilla present, consisting of wide, very shallow U-shaped structure; juxta small, platelike with short, setae-bear- ing arms; valva moderately broad, covered on inner surface with slender setae (cyrdip- sa), Or covered on inner surface, in part, with broadened, spinelike setae and, in part, with slender setae (cinilixa); outer surface of valva with long, black, weakly-attached scales that sometimes obscure some fea- tures of inner valva; sacculus with apex a pointed process; base of sacculus with an extended lateral arm bearing either a large, long scale tuft (cyrdipsa), or bearing a large, long scale tuft and a much smaller, short scale tuft (cinilixa); aedoeagus slen- der; vesica with elongate patch of micros- pines, and a folded, in part sclerotized, platelike element; vinculum slender, con- stricted medially, with well-developed, pos- teriorly directed, lateral arms at anterior end. Female genitalia (of cyrdipsa) (Fig. 28) with ductus bursae shorter than corpus bursae and with cluster of small spines at its distal end; corpus bursae with scobinate patch and large hooked spine (spine, in gen- eral, laterally directed and extending across about 4% width of corpus bursae). KEY TO SPECIES OF MALE DRESCOMA (A key including females is not possible because females of cinilixa remain unknown.) 1. Underside of hindwing with subcostal, rough patch of pale brown, semi-metallic scales, fol- lowed by group of yellow scales and a linear cluster of short, slender, black scales; sacculus of genitalia with apical part rounded (Fig. 14) Je tsh UEC ee aeonas Fey ee tett oh eth sa is heey eee a cyrdipsa Dyar — Underside of hindwing with subcostal, smooth patch of mostly whitish-yellow scales, fol- lowed by streak of black and streak of reddish VOLUME 104, NUMBER 4 brown scales; sacculus of genitalia with apical part slender and sharply pointed (Fig. 16) .. . cinilixa Dyar Drescoma cyrdipsa Dyar (Figs. 2, 8, 14-15, 24, 28) Drescoma cinilixa Dyar 1914:329 This 1s an easily recognized species. Both the forewing and hindwing of the male have obvious costal notches (the vein below the notch in the hindwing is, also, noticeably angled) (Fig. 24). Also, on the underside of the male hindwing there is a pale, rough, brownish, semi-metallic, subcostal patch of scales near the base of the wing, followed by a group of small, yellowish scales and a linear cluster of short, slender, black scales. In addition near these contrasting scales, between the stalk of Sc+R and the stalk of Cu+A is found a well-developed patch of black scales. Furthermore, a pale brown scale pencil, within a short yellowish-white fold, is present on the anal margin of the hindwing. The male genitalia (Figs. 14—15) have a well-developed transtilla with lateral arms that are distinctly broadened, and the apical part of the sacculus is rounded; the female genitalia (Fig. 28) have a large hooklike spine in the corpus bursae that extends about % the distance across the width of the corpus bursae. Costa Rican material examined.—Esta- cion Sirena, Parque Nacional Corcovado, Provincia Puntarenas, May—July, 1992, G. Fonseca, INBio CRI001741662, INBio CR 1000708220 (1 3, 1 ¢); Cerro Tortuguero, QO-—120 m. Parque Nacional Tortuguero, Provincia Limon, May, 1991, J. Solano, IN- Bio CRIO01398617 (1 <¢). Other material examined.—Mexico, Chiapas, May, 1915, R. Muller (1 2); Be- lize, San Ignacio, May, 1990, L. C. Dow (1 3d), 1000 Foot Falls, Mountain Pine Ridge, June 1990, L. C. Dow (1 ¢); Guatemala, Cayuga, January, May, June, August, (no year on label), W. Schaus & W. Barnes (2 3d, 2 2); Panama, Cabima, May, 1911, A. Busck (1 6), Corozal, February and No- 985 vember, 1911, A. Busck (1 6, 1 $3), La Chorrera, April and May, 1911, A. Busck (1 3d, 1 2; syntypes of Drescoma cyrdipsa in USNM), Rio Trinidad, March 1912, A. Busck (2 6, 1 2), Taberilla, (no date on label), (1 9); French Guiana, St. Jean Ma- roni, (no date on label), (1 2); Brazil, Pipa, March, 1994, V. O. Becker (1 ¢). Drescoma cinilixa Dyar (Figs. 3, 16—17) Drescoma cinilixa Dyar 1914:329 Drescoma cinilixa is closely related to Drescoma_ cyrdipsa. This relationship is particularly apparent in males of both spe- cies, each with the costa of the forewing and hindwing strongly notched. Neverthe- less, the two can be distinguished by the underside of the male hindwing. Drescoma cinilixa has the wing with a smooth, sub- costal, basal patch of mostly whitish-yellow scales (in some specimens the patch has a few brown scales), followed distally by a streak of black and a streak of reddish- brown scales. There is only a small black patch of scales, or no black patch, between the stalk of Sc+R and the stalk of Cu+A. A partially black fold on the anal margin of the hindwing encloses a short, black scale pencil. The male genitalia (Fig. 16) include a transtilla that is much thinner than that of Drescoma cyrdipsa. The sacculus at its api- cal end is slender and sharply pointed, and at its base bears both a large scaled tuft and a secondary, small tuft of scales. We have not been able to associate the sexes of Dres- coma cinilixa; consequently, the female genitalia are unknown. Costa Rican material examined.—Esta- cion Los Almendros, Provincia Guanacaste, 300 m., 4—12 September, 1994, E. Lopez, INBio CR 1002019761 (1 3); Acosa, Es- tacion Sirena, Provincia Puntarenas, 6—12 Aprils 1995; Av Picardo; ,INB10 "CR 1002196747 (1 c). 986 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON R ~ Wy =e Ne SS EY Figs. 12-17. Male genitalia, ventral view. 12, Megarthria peterseni (aedoeagus omitted). 13, M. peterseni, aedoeagus. 14, Drescoma cyrdipsa (aedoeagus omitted). 15, D. cyrdipsa, aedoeagus. 16, D. cinilixa (aedoeagus omitted). 17, D. cinilixa, aedoeagus. Other material examined.—Guatemala, Lascelina Heinrich Cayuga, May (no year on label), W. Shaus — Lascelina Heinrich 1956:264. Type species: & W. Barnes (1 6); Panama, La Chorrera, Lascelina canens Heinrich 1956. Original May, 1912, A. Busck (1 ¢; type in USNM). designation. VOLUME 104, NUMBER 4 987 Figs. 18-23. Megacerdresa Neunzig 1994:360. New synonymy. Type species: Megacerdresa cordobensis Neunzig 1994. Original des- ignation. Note.—Lascelina was rejected by Neun- zig in 1994 as a place to include cordob- Male genitalia, ventral view. 18, Lascelina cordobensis (aedoeagus omitted). 19, L. cordobensis, aedoeagus. 20, L. pitilla (aedoeagus omitted). 21, L. pitilla aedoeagus. 22, L. papillina (aedoeagus omitted). 23, L. papillina aedoeagus. ensis, and a new genus, Megacerdresa, was proposed, chiefly because Heinrich’s con- cept of the group did not contain species with more than seven veins in the hindwing (cordobensis has eight). The study of many recently available members of the complex 988 in Costa Rica and the description by Neun- zig (1996) of a new species of Lascelina in the Dominican Republic reveals significant similarities in the male genitalia. Thus we synonymize Megacerdresa and Lascelina despite differences in venation. Description.—Antenna of male with scape cylindrical, swollen apically; shaft with distinct sinus at base; basal and distal ends of sinus each with a small spine, or low, stout spinelike element; inner surface of sinus covered with appressed setae; sinus with or without brush or tuft of scales at- tached to posterior base; sensilla trichodea (cilia) of antenna about % as long as width of shaft at midsinus. Antenna of female simple. Labial palpus of male upturned, reaching above vertex, 3rd segment about as long as 2nd segment. Maxillary palpus of both sexes small, short scaled. Haustel- lum well developed. Ocellus present. Fore- wing of male simple, underside with dark subcostal patch or streak in some species. Forewing of both sexes smooth, with 1] veins; R,,, and R; stalked for about % or slightly over % their lengths; M, slightly bowed; M, and M, close, but separated at base, or shortly stalked; CuA, arises from lower angle of cell; CuA, arising well be- fore lower angle of cell. Hindwing under- side, in some species, with dark patch or streak; with seven to eight veins (1A, 2A, 3A together treated as one vein); Sc+R, and Rs fused for about % their lengths be- yond cell; M, straight; M, and M, fused for about % their lengths or completely fused; CuA, briefly fused with stalk of M, and M,, or stalked for about % its length with fused M,,;; CuA, arises from just before lower angle of cell, or from lower angle of cell; cell distinctly less than % length of wing. Male abdominal segment 8 with small tuft, consisting of rather straight, simple short scales (within pocket of sternite of abdom- inal segment 8). Male genitalia with uncus broadly rounded, truncated, slightly point- ed, or nipple-like distally; gnathos apically completely fused to wall of scaphium (without apical hook or rodlike element); PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON transtilla absent; juxta a plate with short, lateral, setiferous arms; valva moderately broad, covered on distal half of inner sur- face with broadened spinelike setae; sac- culus with apex usually with posteriorly projecting lobe; base of sacculus with or without moderately large, ventral scale tuft (all species without small secondary scale tuft); aedoeagus robust, with bent, supple- mental, basal appendage; vesica with patch of microspines and two sclerotized plates; vinculum broad, without constriction, about as long as greatest width. Female genitalia (cordobensis, canens, pitilla) with ductus bursae shorter than corpus bursae, slightly twisted, usually with a few scobinations near ostium bursae, and a dense, elongate, medial patch of many microspines and nu- merous sclerotized, irregular ridges near junction with corpus bursae; corpus bursae membranous, with curved denticulate spine, originating from near wall of corpus and directed mostly anteriorly. KEY TO SPECIES OF MALE LASCELINA (A key including both sexes is not possible because the female of papillina is unknown.) 1. Enlarged base of antennal shaft surrounding si- nus dark; apex of uncus broadly rounded (Fig. iis) es Sesame ee ers ert & — Enlarged base of antennal shaft surrounding si- nus pale; apex of uncus slightly pointed (Fig. 20) onmmipplelike (B1e22) her enn 2, 2. Antenna without scale tuft associated with si- nus (Fig. 10); apex of uncus slightly pointed (iese2.0) eee ee pitilla Neunzig and Solis, n. sp. — Antenna with scale tuft associated with sinus (Fig. 11); apex of uncus nipple-like (Fig. 22) cordobensis (Neunzig) papillina Neunzig and Solis. n. sp. Lascelina cordobensis (Neunzig), new combination (Figs. 4, 9, 18-19, 27) Megacerdresa cordobensis Neunzig 1994: 562: The male of Lascelina cordobensis can be identified using a combination of fea- tures. These are found mainly in the geni- talia and include a broadly rounded apex of VOLUME 104, NUMBER 4 989 Figs. 24-29. Venation of male right forewing and hindwing, dorsal view, hoodlike prominence on male right wing, dorsal view, and female genitalia, anterior part, ventral view, 24, Drescoma cyrdipsa. 25-26 Me- garthria peterseni. 27, Lascelina cordobensis. 28, Drescoma cyrdipsa. 29, L. pitilla. the uncus, a complete absence of a rodlike — the easily seen dark color of the enlarged median distal hook on the gnathos, and a_ base of the shaft of the male antenna that valva with its distal margin angulate (Fig. surrounds the sinus. 18). Useful for a tentative identification is Costa Rican material examined.—Esta- 990 cion La Casona, 1,520 m. Reserva Biolo- gica Monteverde, Provincia Puntarenas, September, 1992, N. Obando, INBio CR 1000947024 (1 3d); Cerro Tortuguero, Parque Nacional Tortuguero, 0-100 m., Provincia Limon, March, May, 1991, J. So- lano, INBio CR 1001399037, INBio CR 1000197451 (1 3, 1 &). Other material examined.—Mexico, Cor- doba, August—December, 1966, A. B. Lau G06 3913 2 holotype=o in USNMD): Lascelina pitilla Neunzig and Solis, new species (Figs. 5, 10, 20—21, 29) Diagnosis.—The external habitus of pi- tilla together with the general appearance of its male genitalia clearly show that pitilla belongs to the genus Lascelina (Figs. 20— 21). The male antenna will distinguish it from its congeners. Other species in the ge- nus have an obvious scale tuft associated with the sinus of the shaft; this feature is absent in pitilla (Fig. 10). Description.—Forewing length 6.0—7.5 mm. Head with frons white medially, pale reddish brown near eyes; vertex pale red- dish brown, darker anteriorly; labial palpus outwardly with basal segment mostly white, with some pale reddish brown, 2nd and 3rd segments mostly brown with reddish brown distally; maxillary palpus simple, mostly pale reddish brown with a few dark brown scales basally; antenna of male with sinus in base of shaft; distal and basal end of si- nus with small, sharply pointed, medially directed spine; inner surface of sinus cov- ered with appressed scales; no scale brush or tuft attached to posterior base of sinus. Forewing of male simple without costal fold, costal concavity or sex-scaling; wing above mostly white on costal half and pale brown, reddish brown, brown, and red on posterior half; base brown, suffused with red; a pair of mostly red to reddish-brown diagonally converging bands extend from costa to fuse with brown and red of poste- rior half of wing; bands divide costal half of wing into three contrasting, mostly white PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON patches (band that starts 4% from base of costa incomplete; distal band with weak margins); most distal white patch very small; middle white patch with small iso- lated, elongate red patch at about midcosta, and with pair of red and black discal spots; postmedial line vague, whitish. Hindwing simple, pale brown, darker along margins. Male genitalia (Figs. 20—21), with uncus slightly pointed distally; apical part of gna- thos fused to scaphium; transtilla absent; juxta a thin plate with rounded, setiferous, lateral arms; valva slightly pointed apically; inner base of valva with cluster of setae arising from protuberances; sacculus with ventral tuft of thin scales (tuft reaches to about apex of sacculus); aedoeagus robust, with angled basal supplement; vesica with patch of microspines, and two apically rounded, sclerotized plates; vinculum short, not medially constricted. Female genitalia (Fig. 29) with ductus bursae shorter than corpus bursae and with patch of microspi- nes distally; corpus bursae with curved, scobinate spine. Types.—Holotype: ¢. Est. Pitilla, 700 m., 9 km. S. Sta. Cecilia, P. N. Guanacaste, Prov. Guanacaste, Costa Rica, 27 jul. a 14 ago. 1992, C. Moraga, L-N 330200, 380200, INBio CR 1000393816, genitalia slide 393816 MC (INBio). Paratypes: Same collection label data as holotype except col- lector P. Rios, INBio CR 1000867296, gen- italia slide 4547 HHN (NCSU) (1 4d); Est. Pitilla, 9 km. S. Sta. Cecilia, Prov. Guana., Costa Rica, 700%m:, 18=—23° Jul 199352 Rios, L N 330200 380200, INBio CR 1001767419, genitalia slide 4546 HHN (IN- Bio) (1 o); Est. Pitilla, 700 my? 9 kins: Sta.. Cecilia; Prov. Guan., Costa Rica; €: Moraga, Apr. 1991, L N 330200 380200, INBio CR 1000617183, genitalia slide 617183 MC (INBio) (1 @); Est. Pitilla, 700 m., 9 km. S. Sta. Cecilia, Prov. Guan., Cos- ta Rica, P. Rios; Jul. 1991, LW 330200; 380200, INBio CR 1000346701, genitalia slide 4548 HHN (INBio) (1 &). Etymology.—The specific epithet is based on the type locality, Estacion Pitilla, VOLUME 104, NUMBER 4 located in the Guanacaste National Park, Costa Rica. Lascelina papillina Neunzig and Solis, new species (Figs. 6, 11, 22—23) Diagnosis.—The best diagnostic feature of Lascelina papillina is the triangularly- Shaped uncus with its distal, nipple-like protuberance (Fig. 22). Lascelina pederna- lensis, which occurs in the Dominican Re- public, has a similar uncus, but its distal protuberance is less strongly developed (see fig. 53 of Neunzig 1996). Description.—Forewing length 7.0 mm. Head with frons and vertex brownish white; labial palpus outwardly with basal segment white, whitish brown, and brown, 2nd and 3rd segments whitish brown, in part suf- fused with dark brown; maxillary palpus simple, whitish brown; antenna of male with well-developed sinus at base of shaft; basal and distal ends of sinus with low, stout, spinelike elements; inner surface of sinus covered with appressed scales; tuft of pale brown and brown scales attached to posterior base of sinus (tuft extends slightly beyond distal end of sinus). Forewing of male simple (without costal fold, costal notch, or contrastingly colored sex-scales); wing above mostly white on costal half and brown, reddish brown, and red on posterior half; base dark brown; a pair of black di- agonally converging bands extend from costa to fuse with brown, reddish brown, and red of posterior half of wing (distal band broad, somewhat diffuse with indis- tinct margins); black bands divide costal half of wing into three contrasting, mostly white, patches (most distal patch very small); middle patch with small, isolated, elongate black patch at about midcosta, and with two black distal spots; a sprinkling of red scales chiefly on white patches; post- medial line apparent, whitish, but weakly developed. Hindwing simple, pale brown, darker brown along margins. Male genitalia (Figs. 22—23) with uncus distinctly trian- gular, with a well-developed, nipple-like 99] apex; apical part of gnathos fused to sca- phium; transtilla absent; juxta a thin plate with rounded, setiferous, lateral arms; valva rounded apically; inner base of valva with broad, slightly elevated, moundlike, setifer- ous elements; sacculus with apex rather simple, with small, slender lobe; base of sacculus without ventral tuft of scales; ae- doeagus robust, with angled basal supple- ment; vesica with patch of microspines and two sclerotized plates; vinculum about as long as greatest width, truncated distally. Female genitalia unknown. Type.—Holotype: ¢. Rancho Quemado, Peninsula de Osa, 200 m., Prov. Puntarenas, Costa Rica, E Quesada, Dec. 1991, L-S- 292500, 511000, INBio CR 1000345866, genitalia slide 345866 MC (INBio). Etymology.—The specific epithet is de- rived from papilla, Latin for nipple, and re- fers to the appearance of the distal part of the uncus. ACKNOWLEDGMENTS In large measure this study was possible because of the collecting and curating ef- forts of numerous parataxonomists at the Instituto Nacional de Biodiversidad, Santo Domingo, Costa Rica. M. Comacho and E. Phillips, at the same institution, prepared genitalic slides and arranged for specimen loans, respectively. V. Becker, Brasilia, Bra- zil, generously made available additional study specimens. K. Tuck provided access to types in The Natural History Museum, London, England, and R. Blinn, North Car- olina State University, Raleigh, North Car- olina, U.S.A., made the habitus photo- graphs. We are also grateful to L. Deitz, North Carolina State University; T. Henry, Systematic Entomology Laboratory, U.S. Department of Agriculture, Washington, D. C., U.S.A.; and J. Shaffer, George Mason University, Fairfax, Virginia, U.S.A., for critical reviews of the manuscript. LITERATURE CITED Dyar, H. G. 1914. Report on the Lepidoptera of the Smithsonian Biological Survey of the Panama Ca- S92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nal Zone. Proceedings of the United States Na- tional Museum 47: 139-350. Heinrich, C. 1956. American moths of the subfamily Phycitinae. United States National Museum Bul- letin 207: 1-581. Neunzig, H. H. 1994. New genera and species of Mex- ican Phycitinae (Lepidoptera : Pyralidae). Proceed- ings of the Entomological Society of Washington 96: 357-366. . 1996. New species of Phycitinae (Lepidop- tera: Pyralidae) from the Dominican Republic. Proceedings of the Entomological Society of Washington 98: 774-801. Neunzig, H. H. and L. C. Dow. 1993. The Phycitinae of Belize (Lepidoptera : Pyralidae). North Carolina Agricultural Research Service Technical Bulletin 304: 1-133. Neunzig, H. H. and M. A. Solis. 2002. The Ceracan- thia complex (Lepidoptera : Pyralidae : Phycitinae) in Costa Rica. I. Ceracanthia Ragonot. Proceed- ings of the Entomological Society of Washington 104:837-855. Ragonot, E. 1893. Monographie des Phycitinae et des Galleriinae. /n Romanoff, N. M. Mémoires sur les Lépidopteres, Vol. 7, Imprimerie Générale Lahure, Paris, 658 pp. Zeller, P. C. 1881. Columbische, Chiloniden, Crambi- den und Phycideen. Horae Societatis Entomolo- gicae Rossicae 16: 154—256. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 993-996 THE CHEWING LOUSE GENUS KAYSIUS PRICE AND CLAYTON (PHTHIRAPTERA: AMBLYCERA: MENOPONIDAE) FROM THE PASSERIFORMES (AVES) ROGER D. PRICE AND ROBERT C. DALGLEISH (RDP) Department of Entomology and Plant Pathology, Oklahoma State University, Stillwater, OK 74078-0464, U.S.A.; (current address) 4202 Stanard Circle, Fort Smith, AR 72903-1906, U.S.A. (e-mail: rpricelice@aol.com); (RCD) 10601 Tierrasanta Boule- vard, San Diego, CA 92124-2692, U.S.A. (e-mail: rcdalgleish @san.rr.com) Abstract.—The menoponid genus Kaysius Price and Clayton, originally described as monotypic for K. emersoni Price and Clayton, is redefined and expanded to include two species previously placed in Machaerilaemus Harrison: Menopon tityrus Carriker and Machaerilaemus cotingae Carriker, both new combinations. Character states and illus- trations are provided for the separation of these three species. Key Words: When Price and Clayton (1989) de- scribed the menoponid chewing louse ge- the single new species, Kaysius emersoni, nus Kaysius, they did so basing it on whose type host is Glyphorynchus spirurus (Vieillot) from Peru. At that time, it was difficult to decide which features were of generic importance and which should be relegated only to the specific definition. This is virtually always the case in dealing with a monotypic genus such as this. How- ever, when a recent study of the genus Ma- chaerilaemus Harrison was undertaken, it was discovered that two of the 25 species placed in that genus to date actually are species belonging to Kaysius. With this finding that Kaysius is more widespread than initially thought, we have been able to re-examine the definition of Kaysius and contribute to the of these three species. We suspect that further col- lecting from additional passerines will broaden the known host range of Kaysius even further. identification chewing lice, Kaysius, Menoponidae, Phthiraptera, Passeriformes Kaysius Price and Clayton Kaysius Price and Clayton 1989:29. Type species: Kaysius emersoni Price and Clayton, by monotypy. The following features may be used to characterize species belonging to this ge- nus. Little sexual dimorphism, except that associated with smaller male dimensions, reduced male setal numbers, and terminalia. Head width approximately twice its length; without preocular notch or slit; all temple setae 21—27 present (Fig. 3, numbering lat- erad from midline), with 24 and 26 very short, 25 and 27 very long; alveoli of tem- ple setae 26 and 27 well separated; antennal fossa not deep, with antenna entirely or mostly exposed, but completely beneath head; gular plate pigmented only along se- tal bases; without ventral spinous processes; nodi and associated carinae weak; hypo- pharynx with weakly developed sclerites. Thorax with prosternal plate developed (Fig. 2), rounded posteriorly, usually with- out central setae, rarely with | central seta; 994 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Comparison of characters separating Kaysius from Machaerilaemus. Character Kaysius Machaerilaemus 1. Alveoli of temple setae 26 & 27. Well separated 2. Gular pigmentation Only along setal bases 3. Prosternal plate shape Rounded posteriorly 4. Prosternal plate central setae 0-1 5. Mesosternal plate setae Only 2 6. Sternite I Distinctly separate 7. Outermost seta on tergite I Very short 8. Male genitalia 9. Female subgenital plate margin Slight asymmetry with blunt par- ameres curved same direction Straight to gently rounded Adjacent More extensive Pointed posteriorly More than 3 More than 5 Fused with metasternum Very long postspiracular Symmetrical with long pointed outwardly curved parameres Distinct medioposterior con- vexity outer central pronotal seta longer than mi- nute inner seta; mesosternal plate with only 2 associated short setae; venter of femur III without ctenidia or well-developed brush; only 2 medioanterior mesonotal setae as- sociated with postnotum. Abdomen with no tergites enlarged, divided, or with anterior setae; tergite I with short seta laterad of postspiracular seta; all postspiracular setae on I-VIHI very long; sternite I present as distinct plate, with setae; lateral areas of sternites without ctenidia or brushes; female sternite VII partially fused with VIII; fe- male anus essentially oval, without inner setae; female subgenital plate margin straight to gently rounded, not convex me- dially; male genitalia with slender paramer- es flexed in same direction, rounded en- domeral plate, sac weakly spined with poor- ly to well-developed associated sclerite. Kaysius is separated from the closely relat- ed Machaerilaemus by the features given in Table 1. For brevity, generic characters will not be repeated under the species descrip- tions. Host classification follows that of Howard and Moore (1991). In the following descriptions, all mea- surements are in millimeters. Abbreviations for dimensions are TW, head width at tem- ples; HL, head length at midline; PW, pro- thorax width; MW, metathorax width; AWIV, abdomen width at level of segment IV; TL, total length; ANW, female anus width; GL, male genitalia length from tip of basal apodeme to end of parameres; GW, male genitalia width at paramere base. Kaysius emersoni Price and Clayton (Fig. 6) Kaysius emersoni Price and Clayton 1989: 31. Type host: Glyphorynchus spirurus (Vieillot). This species has been very adequately described and illustrated by Price and Clay- ton (1989) and details will not be repeated here, other than to provide an illustration of the male genitalia (Fig. 6) for comparative purposes with those of the other two species treated here. All material of K. emersoni known to date has been collected from G. spirurus |Dendrocolaptidae] in Peru. Kaysius tityrus (Carriker), n. comb. (Figs. 1—4) Menopon tityrus Carriker 1903:182. Type host: Tityra personata = Tityra semifas- ciata costaricensis Ridgway. Female.—As in Fig. 1. Metanotum with 10—12 marginal setae, 4 short medioanter- ior setae; metasternal plate with 12—18 se- tae. Tergal setae on I, 12—15; II-V, 16—20; VI, 17-20; VII, 14-19; VII, 12-15; [X, 9— 14. Prominent pleural thickenings on I- VIII. Sternal setae on I, 6-10; I, 22—26; III, 26-31; IV-VI, 26-35; VII, 19—27. Subgen- ital plate with 14-21 marginal, 12—17 an- terior setae. Anus with 30—39 dorsal, 36— VOLUME 104, NUMBER 4 a we FX a Ti ih Wy ¢ YY Including the Barnadesioidea, separated subsequently as a subfamily (Bremer 1994). *The only specimen of Lamproxynella separata we obtained was reared from one sample of Senecio brasi- liensis (Speng.) Less. capitula, and is tentatively considered exclusive for Senecioneae. * The total species for Brazil estimated by Barroso (1986) also includes the tribes Anthemideae, Calenduleae, and Cotuleae, from which we sampled no species. Nonetheless, there are important differences between the two major tribes of Brazilian Asteraceae, Vernonieae and Eupatorieae. The Vernonieae have greater tephritid spe- cies richness and a greater proportion of plant species with these insects, despite the similar numbers of species in Brazil, and of the sampled species in these tribes (Table 1). The proportion of tephritid species ex- clusive to each tribe is also markedly great- er among the Vernonieae, and very low in Eupatorieae (Table 1). Two possible non- exclusive explanations are that the Vernon- ieae are chemically and morphologically more distinct and/or were colonized by te- phritid lineages intrinsically more special- ized. The species from the main genera as- sociated with Vernonieae (i.e., Tomoplagia, Dictyotrypeta, and Tetreuaresta), are nor- mally restricted to related genera of host plants, in contrast to Xanthaciura, Cecido- chares and Trupanea species, the most im- portant genera associated with the tribe Eu- patorieae. Five of the seven Xanthaciura species primarily associated with Eupato- rieae occasionally use plants from the tribe Heliantheae, its sister group (Bremer 1994), and, more rarely, from the tribes Astereae and Vernonieae. Two of the three Cecido- chares species have as an occasional host Moquinia racemosa (Spreng.) DC., which has a controversial tribal status but is not an Eupatorieae (Robinson 1994). Five of seven sampled Trupanea species may occur on Eupatorieae, but only one 1s exclusive to this tribe. A notable exception to the general cor- relation of plant tribal diversity and tephri- tid diversity in southern Brazil is the virtual absence of endophagous tephritids on Se- necioneae. From capitula samples of 22 Se- necioneae species, we reared only one in- dividual each of Dioxyna chilensis (Mac- quart), 7rypanaresta coelestina (Hering), Lamproxynella separata (Malloch) (Appen- dix I), and 7Trupanea sp. The Senecioneae cannot be considered major hosts of any of these species, except perhaps L. separata, whose host range needs further investiga- tion (see next section). In the Nearctic Re- gion, capitula of Senecioneae are used by at least ten Tephritidae species in the genera 1012 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Number of obtained Tephritidae specimens and species (described, undescribed and total), by genus. Total species exceeds the sum of described and undescribed species because it includes morphospecies of unclear taxonomic status (especially Trupanea). Also shown, the number of described species for each genus for the Neotropical Region (according to Norrbom et al. 1999a), and the tribes of the main host plants for each genus. No. Descr. Undescr. Total Genus Specim.! Spp. Spp. Spp. Acinia Sy, 3 0) 8} Acrotaeniini gen. | 16 0) | | Acrotaeniini gen. 2 3] 0) l | Acrotaeniini gen. 3 401 0) | | Cecidochares 223) 2 | 3 Dictyotrypeta 35)7/ | 6 7 Dioxyna 155 3S: 0) 3 Dyseuaresta 387 O 3} Euarestoides 98 0) | | Lamproxynela l | 0) | Neomyopites 105 | | 2 Paracantha 16 l 0) l Plaumannimyia 29 | O l Tetreuaresta 347 0) 3 5 Tomoplagia 3,375 12 13 25 Trupanea HM) O 0) 8 Trypanaresta HD 3} 0) 5 Xanthaciura 3,294 6 3 9 TOTAL 9,697 35 3] 80 Neotrop. Spp. Main Host Tribes 9 Plucheeae Vernonieae Vernonieae Mutisieae 12 Eupatorieae 4 Vernonieae, Mutisieae, Heliantheae 5 Heliantheae, Helenieae 12 Heliantheae 3} Mutisieae 8 Senecioneae 28 Eupatorieae, Astereae 7 Heliantheae 2 Astereae 19 Vernonieae 45 Vernonieae* 80 Eupatorieae, Mutisieae 17 Astereae 17 Eupatorieae, Heliantheae 259 ' In order to show the relative importance of each genus, only specimens from extensive surveys (i.e., able to include all genera) are summed in this column. 2 Tomoplagia is primarily associated with the tribe Vernonieae, but three species are specialists on other tribes: T. costalimai and T. rivittata (on Mutisieae), and 7. biseriata (on Heliantheae). Aciurina, Stenopa, Tephritis, Trupanea, and Campiglossa (Foote et al. 1993). Since the latter two genera also occur in the Neotrop- ics, our finding of tephritid absence in Bra- zilian Senecioneae cannot be ascribed to the lack of colonizing groups. This absence might be attributed to the lower diversity of Senecioneae in Brazil compared to other important tribes (Table 1). However, in two tribes that are smaller and were less sam- pled than Senecioneae, tephritids are better represented (Table 1). In plants of the tribe Inuleae (genus Pluchea), we recorded three Acinia species (37 specimens) that are ex- clusive to this tribe (Table 2, and Appendix I). The only sampled genus of the tribe He- lenieae, Porophyllum, includes main hosts of Dioxyna chilensis (16 specimens in Po- rophyllum, Appendix I) and an unidentified species of 7rupanea (six specimens in Po- rophyllum). As already pointed out by Lew- insohn (1988), such scarcity of specialized tephritid species in Senecioneae is the first record of this nature for a whole Asteraceae tribe on a large geographical scale. In Appendix I we list the 35 identified species, their host plants, and sample local- ities. One third of these species and three of the genera (Acinia, Euarestoides, and Paracantha) were not previously recorded for Brazil (Norrbom et al. 1999b). The majority of the unidentified morpho- species are undescribed species, and they total a minimum of 31. The other species belong to genera whose species are difficult to identify because they are in need of sys- tematic revision, especially Trupanea, Try- panaresta and Dyseuaresta (Foote et al. 1993). We also obtained three new genera, all belonging to the Tribe Acrotaeniini (Ta- ble 2), associated with endemic hosts of highland meadows on rocky outcrops (the Campo Rupestre vegetation, Giulietti and Pirani 1988). Also, the genus Dictyotrypeta, VOLUME 104, NUMBER 4 as currently recognized, may be polyphy- letic (Norrbom and Prado, unpublished data). More than a third (31 of 80) of the spe- cies we obtained are undescribed, and this indicates how little known taxonomically neotropical tephritids still are. It also sug- gests that tephritid species richness is much higher than currently known. Extrapolating the ratio of 39% of new species we found to the total of tephritid species described for the Neotropical Region (716 spp., Norrbom et al. 1999a), we achieve an estimate of ap- proximately 990 species, similar to the number of species currently recognized in the Afrotropical and Oriental Regions, con- sidered today the areas with the greatest Te- phritidae richness (Norrbom et al. 1999a). Although the number of described species in these regions presumably will rise with new occurrence data and new descriptions (e.g., see Freidberg and Norrbom 1999), the Neotropical Region is perhaps the least studied (Foote et al. 1993), and may have a larger portion of its fauna unknown. The Tephritidae fauna of Asteraceae ca- pitula in Brazil is dominated by the genus Tomoplagia, which comprises 31% of the morphospecies and 35% of the reared spec- imens (Table 2). Xanthaciura and Trupanea are, respectively, the second and third most important genera, both in species and in specimen numbers (Table 2). The genera Dictyotrypeta, Tetreuaresta and Trypana- resta follow, with five to seven morphospe- cies and two to four percent of the speci- mens each (Table 2). For the other 12 gen- era there are a maximum of three morpho- species each, and together they represent 17% of total specimens (Table 2). This fau- na is predominantly Neotropical, except for the genera Trupanea, Dioxyna (with cos- mopolitan distribution), and Paracantha (Panamerican). The other genera are Neo- tropical in origin, totaling 80% of the spe- cies and 90% of the obtained specimens. The genus Acinia, as currently recognized, includes Palearctic and American species, but according to Korneyev (1999) the latter 1013 are not closely related to the former and should be removed, thus this group of spe- cies should also be considered of Neotrop- ical origin. Although our survey is restricted to part of Brazil, we obtained an average of 30% of the number of recorded species of the Neotropics for each genus. For Tomopla- gia, tor example, we obtained 25 morpho- species compared with 45 described neo- tropical species (55%). Other important genera in our sample show a similar trend, such as Xanthaciura (9 morphospecies compared with 17 described ones), 7rypan- aresta (5:17), and Tetreuaresta (5:19). For all of these examples, the proportions are high because we obtained not only a rep- resentative percentage of the described spe- cies, but also many undescribed ones. In the genus Dictyotrypeta, for example, we ob- tained more morphospecies than the num- ber of described ones (7:4). In contrast, the numbers of morphospecies of Trupanea, Paracantha, Lamproxynella, and Neomy- opites in this study were small in relation to the total species recorded in the Neo- tropics: (/—15%, see-Wable 2). It is. very doubtful that these genera are associated with plant families other than Asteraceae in Brazil, but at least some of the species might develop in plant organs other than the flowerheads. There are few species of these genera recorded for Brazil (Norrbom et al. 1999b), however, and they probably are more diversified in other areas in the Neo- tropical Region. The endophagous Tephritidae of capitula in southern Brazil are highly specialized. Sixty percent of the species were obtained from hosts belonging to a particular Aster- aceae tribe or lower taxonomic category (Fig. 2). The main hosts for 80% of the spe- cies are restricted to a subtribe or lower cat- egory, and only 4% of the species have main hosts in more than one tribe. Oli- gophagy may be a general trend in the Te- phritinae, since similar degrees of speciali- zation were observed in the Nearctic and Palearctic Regions (Zw6lfer 1982; Sobhian 1014 Species n=13 Natta A LA wy SA y| Genus ~~ n= 32 Fig. 2. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2 Tribes Pie diagram of main host plant range of the obtained species of Tephritidae. Wedges represent the percentages of tephritid species reared from single host species, a single host genus, subtribe, or tribe, or from 2 tribes. Species reared from a single sample were excluded, because the data were too scarce to ascertain their host ranges. and Zwolfer 1985; Goeden 1987, 1989, 1992, 1993, 1994, 1997; White and Elson- Harris 1992; Headrick and Goeden 1998). In the Tephritidae, capitula utilization de- pends on very specialized adaptations to the morphology, chemistry, and phenology of the host plant, which in general hinders the use of very different host plants (Zw6lfer 1982, 1987; Straw 1989a, b). The genera of endophagous Tephritidae of capitula also are fairly specialized. Only three of the ob- tained genera have main host plants in more than two tribes. Two of these, Tomoplagia and Dictyotrypeta, are associated with three Asteraceae tribes, but are composed mostly of specialized species. Tomoplagia is main- ly associated with one tribe (Vernonieae), with some species using plants from the tribes Mutisieae or Heliantheae. Among Dictyotrypeta species reared, two use plants of the related tribes Vernonieae and Mutis- ieae, two species are specialists on Helian- theae, and the remaining four species are associated with Vernonieae. Finally, 7ru- panea does in fact include many more gen- eralist species and has been recorded from at least six different tribes of Asteraceae in other regions of the world (Munro 1964, Goeden 1992, White and Elson-Harris 1992, Foote et al. 1993). However, the ge- nus is in dire need of revision, and host ranges of revised species may be smaller than they appear now. As will be detailed in the next section, VOLUME 104, NUMBER 4 host plant taxa for tephritid species and genera agree with the published records from other world regions and the scarce re- cords for the Neotropics (Foote et al. 1993, Norrbom et al. 1999a). The small variation in the host plants used, even in different biogeographic regions, suggests that the set of potential hosts for a particular Tephriti- dae taxon is under strong phylogenetic re- striction, as suggested by Zw6lfer (1982, 1987), Straw (1989b), and Lewinsohn and Prado (in press). NOTES ON THE OBTAINED GENERA In the following discussion, the genera are arranged alphabetically, except for Ew- arestoides and Plaumannimyia, which are discussed with 7rypanaresta. The genus Acinia has 13 described spe- cies, widely distributed in the Americas and Palearctic Region (Norrbom et al. 1999a,b). However, Korneyev (1999) has indicated that the nine American (mainly Neotropi- cal) species should be removed from the ge- nus. We obtained specimens of A. picturata (Snow) and A. ica Hering from Pluchea laxiflora Hook, and Arn. ex. Baker, and of A. aurata Aczél from Pluchea_ sagitalis (Lam.) Cabrera (Appendix I). According to these records and those of Aczél (1958), Frias (1992) and Foote et al. (1993), hosts of Acinia in the New World are restricted to the tribe Plucheeae (Inuleae s. 1.). No Acinia species was previously recorded for Brazil (Norrbom et al. 1999a, b), although Foote (1980, Table 1) indicated he had ex- amined Brazilian specimens in a geograph- ical distribution table of Neotropical gen- era. Cecidochares is a mainly Neotropical ge- nus, although C. caliginosa (Foote) (trans- ferred from Procecidocharoides by Norr- bom et al. 1999a, b) occurs in the south- western United States and Mexico. There are 13 described species (Norrbom et al. 1999a, b), and at least 10 undescribed (Norrbom, unpublished). Specimens iden- tified as C. connexa (Macquart) are referred to as ‘C. connexa complex’ (See Appendix 1015 I), because recent biological evidence sug- gest that they may be a complex of cryptic species (McFadyen et al. pers. com.). The species of Cecidochares of known biology are all associated with Eupatorieae, and most are gall makers (Lima 1934, Silva et al. 1968, Cruttwell 1974, Foote et al. 1993). We reared three species of Cecidochares from capitula of the Tribe Eupatorieae, most of them without signs of galling. In one sample of C. connexa complex (from Chromolaena laevigata (Lam.) R. King and H. Rob., Parana State), however, there were galls within the flower heads. Dictyotrypeta is an exclusively Neotrop- ical genus with numerous undescribed spe- cies (Foote 1980, Norrbom and Prado, un- publ.). Its limits are not well established, and it may be polyphyletic. The species reared in this study may not be congeneric with D. syssema Hendel, the type species (Norrbom and Prado, unpublished). The bi- ology of the genus is poorly known; Foote et al. (1993) stated that four species breed in Heliantheae flowers. Of the seven Dic- tyotrypeta morphospecies we recorded, three use Vernonieae hosts (including D. atacta (Hendel), see Appendix I), two use Heliantheae, and two species use hosts from the tribes Vernonieae and Mutisieae. Dioxyna is a broadly distributed genus with 13 described species. In the New World, their hosts are mainly species of the related tribes Heliantheae and Helenieae. The North American and Caribbean spe- cies, D. picciola (Bigot), is a relative gen- eralist, using plants from the tribes Aster- eae, Heliantheae and Helenieae, whereas D. thomae (Curran) is recorded only from Bi- dens bipinnata L. and Wedelia sp. (Helian- theae) (Foote et al. 1993). In our surveys, all of the main hosts for D. thomae and D. peregrina (Loew) belong to the genus Bi- dens. The third species we obtained, D. chi- lensis (Marcquart), was reared from /sostig- ma sp. (Heliantheae) and Porophyllum rud- erale (Jacq.) Cass. (Helenieae) (Appendix I). Dyseuaresta is a little known, American, 1016 mainly Neotropical genus (Foote 1980, Foote et al. 1993), with 12 described spe- cies (Norrbom et al. 1999a, b). Our data and records of D. mexicana (Wiedemann) from Melanthera spp. (Wasbauer 1972) suggest that this genus is primarily associated with the tribe Heliantheae. Frias (1992) reported D. impluviata (Blanchard) from Senecio, but this species may belong to Lamproxy- nella, a genus whose limits with Dyseu- aresta are not well defined. We obtained Dyseuaresta adelphica (Hendel) from ca- pitula of Aspilia spp. and from one species of Calea (Heliantheae). We reared two un- described species, one from flowerheads of Verbesina, Calea and Aspilia (all Helian- theae), and the other from Aspilia, although both species occasionally used plants of the tribe Eupatorieae. The genus Lamproxynella is exclusively Neotropical and is considered closely relat- ed to Dyseuaresta (Foote et al. 1993). Of eight described species, only L. separata (Malloch) is recorded from Brazil. We reared one specimen of this species from Senecio brasiliensis (Spreng.) Less. (Sene- cioneae) flowerheads (Appendix I). These data, and the rearing of two other species from Senecioneae in Chile (Norrbom, pers. obs.), are the only records of host plants of Lamproxynella, and suggest its association with the tribe Senecioneae. The genus Neomyopites, which was re- cently proposed to include many of the New World species previously placed in Urophora (Freidberg and Norrbom 1999), has at least 28 described species, all but two of which are restricted to the Neotropical Region. Their known hosts belong to the tribes Astereae, Eupatorieae, Heliantheae, and Liabeae (Freidberg and Norrbom 1999). Some species induce galls in the ca- pitula, as do many Urophora species in the Palearctic Region. We reared N. paulensis (Steyskal) from five Eupatorieae genera (Appendix I). This species is the main en- dophage of the genus Mikania, which is characterized by reduced capitula (with a maximum of four flowers), and has a poor PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON endophagous fauna. The other species we obtained, which is probably undescribed, has a completely hyaline wing and was reared from only one sample of Baccharis articulata (Lam.) Pers. (Astereae). Both species were reared from normal capitula, with no sign of galling. Paracantha is a genus restricted to the New World. Although five of its ten species occur in the Nearctic Region and can even reach southern Canada, the most closely re- lated genera are all Neotropical (Foote et al. 1993). Until now, no species had been recorded for Brazil, although Foote (1980) indicated he had examined Brazilian spec- imens. The previously known hosts for Paracantha are from the genus Cirsium (tribe Cardueae) and several Heliantheae genera and species (Foote et al. 1993). We obtained individuals of Paracantha tralis Malloch from five Heliantheae spe- cies from Minas Gerais, Rio Grande do Sul, and Santa Catarina (Appendix I). In Brazil, there are no native species of the tribe Car- dueae, but three introduced species were sampled in southern Brazil (Cirsium vul- gare (Savi) Ten., Arctium minus Bernh. and Carduus nutans L.). Although we collected a great volume of these plants in seven lo- calities from Santa Catarina and Rio Grande do Sul, no Paracantha emerged from these samples. Tetreuaresta is an exclusively Neotropi- cal genus, with a great number of unde- scribed species (Foote 1980, Norrbom and Prado, unpublished). Its limits are not well established, and it may be polyphyletic. Bi- ologically it is very poorly known, except for Tetreuaresta obscuriventris (Loew), 1n- troduced to Hawaii and other Pacific islands for control of its only known host plant, Elephantopus mollis H.B.K. (Vernonieae) (White and Elson-Harris 1992, Foote et al. 1993). The five Tetreuaresta species we ob- tained are all associated with Vernonieae hosts. The Neotropical genus Tomoplagia has 47 described species (Norrbom et al. 1999a, b) and is primarily associated with the tribe aus- VOLUME 104, NUMBER 4 Vernonieae. Of the two species that occur in the United States, 7. obliqua (Say) is re- corded from at least seven Vernonieae spe- cies, and 7. cressoni Aczél from Trixis cal- ifornica Kellog and Perezia microcephala (DC.) A. Gray (Mutisieae) (Goeden and Headrick 1991, Foote et al. 1993). In Bra- zil, the association with Vernonieae had al- ready been established (Lewinsohn 1988, Prado and Lewinsohn 1994), and 7. rudol- phi (Lutz and Lima) had been recorded making stem galls on Vernonanthura spp. (Lima 1934, Silva et al. 1968). The main hosts for all the species of Tomoplagia we obtained belong to this tribe, with three ex- ceptions, namely, 7. costalimai Aczél, which uses Trixis spp. (Tribe Mutisieae), 7. trivittata (Lutz and Lima), which uses Gochnatia spp. (Mutisieae), and 7. biser- lata (Loew), which uses Calea spp. (He- liantheae) (Appendix I). Plants of the tribes Mutisieae and Heliantheae are occasionally used by some other Tomoplagia species (Appendix I). Therefore, the occasional use of Heliantheae and Mutisieae by an ances- tor may have facilitated the evolution of the species specialized on these tribes. Most species of Tomoplagia are restrict- ed to only one tribe or to a lower taxonomic category of host plants. However, the most polyphagous species, 7. incompleta (Willis- ton), may occasionally use Mutisieae and Eupatorieae as hosts, apart from its Vernon- ieae main hosts (Appendix I, Prado and Lewinsohn 1994). Tomoplagia belongs to the mostly Neotropical tribe Acrotaeniini (Norrbom! 1987, Foote et al. 1993, but see also Korneyev 1999). The known _ host plants for other Acrotaeniini (genera Te- treuaresta and Acrotaenia) belong to the tribes Vernonieae and Heliantheae (Foote et al. 1993, Appendix I). Among the Vernon- ieae, the subtribe Lychnophorinae is virtu- ally endemic to montane rocky outcrops, with greatest diversity in the Serra do Es- pinhago in Minas Gerais (Robinson et al. 1980, Robinson 1992, Bremer 1994). In this study, we obtained six new Tomoplagia species specialized on this subtribe. These 1017 species had not been sampled before and are associated with plants with limited dis- tributions, hence they are also probably en- demic to rocky outcrops (Prado et al., un- published). We also obtained two species from two new genera of Acrotaentini. One of these species uses Lychnophora spp., a genus restricted to highland meadows on rocky outcrops (campos rupestres) and ad- jacent Brazilian savannas (cerrados) from Minas Gerais, Bahia and Goias (Coile and Jones 1981, Robinson et al. 1980, Robinson 1992). The other species was recorded from species of Lychnophora and Eremanthus, a genus found in campo rupestre and cerrado vegetation from central and southeastern Brazil (MacLeish 1987). About 400 speci- mens of a third Acrotaeniini genus were reared from capitula of Wunderlichia mir- abilis Riedel (Mutisieae), a primitive As- teraceae also restricted to rocky outcrops of central Brazil (Bremer 1994). Trupanea is one of the largest genera of Tephritinae, with more than 200 described species, including 80 in the Neotropical Re- gion (Norrbom et al. 1999a, b). Species identification in this genus is extremely dif- ficult, and the Neotropical fauna lacks an adequate taxonomic revision (Foote 1980, Foote et al. 1993). In a preliminary exam- ination, we separated our Trupanea speci- mens into eight morphospecies that are mainly associated with the tribes Eupato- rieae and Mutisieae. One morphospecies was also reared from Porophyllum (tribe Helenieae, formerly placed in the Helian- theae). The known hosts for 7rupanea in the Nearctic Region belong to these three tribes (Wasbauer 1972; Goeden 1985, 1992; Foote et al. 1993). In Chile, the genus is recorded from the tribes Astereae and He- liantheae (Frias 1985). In the present study, the majority of 7Trupanea morphospecies had broader host ranges than the species of the other tephritid genera. Four morphospe- cies were recorded from two to five Aster- aceae tribes. Trypanaresta is a genus of Neotropical origin, with 17 described species (Norrbom 1018 et al. 1999a, b). The synonymy of Euares- toides arnaudi Foote with E. flavus (Ad- ams) and its transfer to T7rypanaresta (Norr- bom et al. 1999b) expanded the known dis- tribution of the genus to the Nearctic Re- gion. Trypanaresta is closely related to Plaumannimyia (which has two described species, restricted to southern Brazil) and Euarestoides (three species in Central America, Mexico, and USA) (Norrbom et al. 1999a, b). Species of the tribe Astereae, mainly of the genus Baccharis, are recorded in the literature as hosts for the first two genera (Foote et al. 1993), which is in ac- cord with our data. The two more abundant species of Trypanaresta in our surveys, T. coelestina (Hering) and 7. imitatrix (Her- ing), were obtained from various Baccharis species (Appendix I), from either male or female flowers. The three remaining species were obtained in small numbers, and from few samples, also from Astereae. One in- dividual of 7. thomsoni (Hendel) was reared from capitula of Solidago chilensis Meyen (Appendix I), two individuals of an uniden- tified species were reared from Conyza spp. and S$. chilensis capitula, and two individ- uals of another unidentified species were reared from capitula of Conyza bonariensis (L.) Cronquist (all in Astereae). We also ob- tained ten individuals of Plaumannimyia pallens Hering from capitula of Baccharis spp. in southern Brazil (Appendix I). In the genus Euarestoides, only the hosts of E. acutangulus (Thomson) were previously known, and these include species of the tribes Heliantheae, Helenieae and Astereae (Foote et al. 1993). In the present study, we obtained one undescribed species of Eu- arestoides from capitula of four species of Gochnatia (Mutisieae) from Minas Gerais and Rio de Janeiro. Euarestoides was not previously recorded from Brazil (Norrbom etalkiyl999b): Xanthaciura is a Neotropical and Nearc- tic genus with 17 described species (Norr- bom et al. 1999a, b). It is known from the central, southeastern, southern and north- eastern regions of Brazil and is associated PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with the tribes Heliantheae and Eupato- rieae. The few published host plant data re- fer to three of the four species that reach the United States, X. connexionis Benjamin, X. insecta (Loew), and X. tetraspina (Phil- lips) (Foote et al. 1993), and their host plant tribes and genera are the same as presented in this study. The fourth species that reach- es the United States, X. chrysura (Thom- son), had no previously reported host plants (Foote et al. 1993). We recorded nine Xan- thaciura species from Asteraceae capitula, three of them undescribed (Table 2). The most abundant species in our samples were X. biocellata (Thomson) and X. chrysura, both primarily associated with various gen- era of Eupatorieae (Appendix I). Xantha- ciura quadrisetosa (Hendel), X. mallochi Aczél, and the three undescribed species are also primarily associated with Eupatorieae. Xanthaciura insecta (Loew) 1s primarily as- sociated with the subtribe Coreopsidinae of the tribe Heliantheae, as shown by our re- cords from Bidens spp. (Appendix I), and those of Wasbauer (1972) from Bidens spp. in the United States, and of Norrbom (in Foote et al. 1993) from Bidens squarrosa Kunth in Venezuela and Dahlia coccinea Cav. in Mexico. Xanthaciura unipuncta Malloch is also associated with Helian- theae, but also breeds in some species of Eupatorieae (Appendix I). ACKNOWLEDGMENTS This work was supported by Fapesp, CNPq, FAEP/Unicamp (Brazil), and the Smithsonian Institution and USDA/ARS (USA). Adalberto dos Santos, Flavia Batis- ta, Vinicius Motta, Gabrielle Althausen, and José Carlos da Silva (Unicamp) provided invaluable assistance in field and laboratory work. Our thanks also to Joao Semir, John Pruski, Nelson Matzenbacher, Harold Rob- inson, and the late Hermdgenes Leitao Fil- ho for plant identification; and to Angelo do Pires Prado for assistance in tephritid systematics and comments on drafts of this paper. John W. Brown, Amnon Freidberg, and Norman Woodley also kindly reviewed VOLUME 104, NUMBER 4 the manuscript. Field lodging at Diamantina was provided by the Centro Eschwege de Geologia, Universidade Federal de Minas Gerais. Special thanks are due to Nelson Matzenbacher, Rogério P. Martins, and Francisco Prado, and their families, who kindly provided housing during part of our field trips. LITERATURE CITED Aczél, M. L. 1958. Revision of the American species of the genus Acinia Robineau-Desvoidy (Diptera, Trypetidae). Revista Brasileira de Entomologia 8: 75-105. Almeida, A. M. insetos endofagos associados a capitulos de 77ri- chogoniopsis adenantha (DC) (Asteraceae). MSc Thesis, Universidade Estadual de Campinas, Cam- pinas. Barroso, G. M. 1986. Sistematica das Angiospermas do Brasil. Imprensa da Universidade Federal de Vicosa, Vicosa. 443 pp. Bremer, K. 1994. Asteraceae: Cladistics and Classifi- cation. Timber Press, Portland. 752 pp. Coile, N. C. and S. B. J. Jones. 1981. Lychnophora (Compositae: Vernonieae), a genus endemic to the Brazilian planalto. Brittonia 33: 528-542. Cruttwell, R. E. 1974. Insects and mites attacking Eu- patorium odoratum in the Neotropics. An anno- tated list of the insects and mites recorded from Eupatorium odoratum, with a key to the types of damage found in Trinidad. Technical Bulletin of the Commomwealth Institute for Biological Con- trol 17: 87-125. Foote, R. H. 1980. Fruit Fly Genera South of the Unit- ed States. Technical Bulletin 1600. United States Department of Agriculture, Washington. 79 pp. Foote, R. 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Fort- schrift fiir Zoologie 25: 331-353. . 1982. Life systems and strategies of resource exploitation in tephritids. CEC/IOBC Symposium, Athens, Nov. 1982. . 1988. Species richness, species packing, and evolution in insect-plant systems, pp. 301-319. In Schulze E. D. and H. Zwolfer, eds. Potentials and Limitations of Ecosystems Analysis. Springer- Verlag, Berlin. APPENDIX I Checklist of Described Tephritid Species Reared from Asteraceae Capitula in Southern and Southeastern Brazil This appendix lists the described species obtained in our survey, their hosts, and sample sites. We also in- VOLUME 104, NUMBER 4 dicate the number of individuals of each species we obtained and the number of samples from which they were reared. Nomenclature and geographical distribu- tions are in accord with Norrbom et al. (1999b). Lo- calities are grouped by states, which are indicated by their abbreviations. Host plants are grouped by tribe. Plant species from which fewer than five tephritid in- dividuals were reared from a single sample were con- servatively considered occasional hosts. For a map of localities see Fig. 1, and for a list of them see Appen- dix II. For full scientific names of host plants, see Ap- pendix III. Acinia aurata Aczél 2 samples, 2 individuals. Localities: SP: Bertioga, Campinas. Previously known distribution: Bolivia, Argentina. Main host: Inuleae: Pluchea sagitalis. Acinia ica Hering | sample, 30 individuals. Locality: MG: Ouro Branco. Previously known distribution: Peru. Main host: Inuleae: Pluchea laxiflora. Acinia picturata (Snow) | sample, 5 individuals. Locality: SC: Matos Costa. Previously known distribution: USA, S to Guatemala, West Indies, Hawaii (introd.). Main host: Inuleae: Pluchea laxiflora. Cecidochares connexa (Macquart) complex 15 samples, 45 individuals. Localities: MG: Camanducaia, Diamantina, Grao Mo- gol, Joaquim Felicio (Serra do Cabral), Parque Es- tadual de Ibitipoca, Serra do Cip6, RJ: Visconde de Maua. Previously known distribution: Venezuela, Argentina, Brazil. Main hosts: Chromolaena cylindrocephala, Chromolaena squal- Eupatorieae: Chromolaena_ chaseae, ida, Pseudobrickellia brasiliensis. Occasional hosts: Eupatorieae: Chromolaena ascen- dens, Chromolaena maximilianit, Chromolaena multiflosculosa, Chromolaena sp., Trichogonia sal- viaefolia, Trichogonia villosa, Moquinieae: Moqui- nia racemosa. Cecidochares fluminensis (Lima) 11 samples, 51 individuals. Localities: MG: Monte Verde, Parque Estadual de Ib- itipoca, Passa Quatro, RJ: Itatiaia, Visconde de Maua, SP: Campinas, Campos do Jordao. Previously known distribution: Central America, Trin- idad, Venezuela, Guyana, Brazil (AM, RO, RJ, SP). Main hosts: Eupatorieae: Chromolaena maximilianii, Chromolaena laevigata. Occasional hosts: Eupatorieae: Chromolaena hookeri- ana Dictyotrypeta atacta (Hendel) 1021 8 samples, 192 individuals. Localities: MG: Diamantina, Joaquim Felicio (Serra do Cabral), Serra do Cipo. Previously known distribution: Brazil, Paraguay. Main hosts: Vernonieae: Lessingianthus buddleiifolius, Lessingianthus coriaceus, Lessingianthus roseus, Lessingianthus vepretorum. Occasional hosts: Vernonieae: Proteopsis argentea, Eupatorieae: Trichogonia villosa. Dioxyna chilensis (Macquart) 7 samples, 77 individuals. Localities: MG: Diamantina, Joaquim Felicio (Serra do Cabral), RS: Guarba, SP: Campinas. Previously known distribution: Peru, Bolivia, Chile, Argentina. Main hosts: Heliantheae: /sostigma sp., Helenieae: Po- rophyllum ruderale. Occasional hosts: Heliantheae: Calea graminifolia, Helenieae: Porophyllum riedelii, Senecioneae: Se- necio oxyphyllus. Dioxyna peregrina (Loew) 7 samples, 26 individuals. Localities: MG: Passa Quatro, SP: Bertioga, Campi- nas, Campos do Jordao, Jundiaf (Serra do Japi). Previously known distribution: Brazil Main host: Heliantheae: Bidens pilosa. Occasional host: Eupatorieae: Praxelis clematidea. Dioxyna thomae (Curran) 3 samples, 39 individuals. Localities: MG: Diamantina, Grao Mogol, Joaquim Felicio (Serra do Cabral). Previously known distribution: U.S.A. (Florida), West Indies, Guyana. Main hosts: Heliantheae: Bidens spp. (two species). Dyseuaresta adelphica (Hendel) 23 samples, 202 individuals. Localities: MG: Diamantina, Grao Mogol, Joaquim Felicio (Serra do Cabral), Ouro Branco, Serra do Cipo. Previously known distribution: Brazil (MS, PR), Par- aguay. Main hosts: Heliantheae: Aspilia jolyana, Aspilia cy- lindrocephala, Aspilia laevissima, six other unde- temined species of Aspilia. Occasional hosts: Heliantheae: Aspilia foliacea, two other undetemined species of Aspilia spp., Calea sp. Lamproxynella separata (Malloch) | sample, | individual. Locality: MG: Pocgos de Caldas. Previously known distribution: Brazil (SP), Argentina. Host: Senecionieae: Senecio brasiliensis. Neomyopites paulensis (Steyskal) 37 samples, 130 individuals. Localities: MG: Camanducaia, Diamantina, Grao Mo- gol, Joaquim Felicio (Serra do Cabral), Ouro Bran- co, Parque Estadual de Ibitipoca, Passa Quatro, Ser- 1022 ra do Cip6, RJ: Marica, Visconde de Maua, SC: La- ges, Mafra, SP: Bertioga, Campinas, Campos do Jordao. Previously known distribution: Trinidad, Paraguay, Brazil (SP). Main hosts: Eupatorieae: Ayapana amigdalina, Chromolaena laevigata, Chromolaena maximilianii, Chromolaena squalida, Grazielia intermedia, Mt- kania cipoensis, Mikania micrantha, Mikania offi- cinalis, Mikania_sessilifolia, Praxelis clematidea, Stomathanthes polycephalus, Symphyopappus de- cussatus. Occasional hosts: Eupatorieae: Chromolaena chaseae, Chromolaena odorata, Mikania retifolia, Mikania sp., Pseudobrickellia brasiliensis. Paracantha australis Malloch 5 samples, 16 individuals. Localities: MG: Serra do Cipo, Diamantina, RS: Cam- bara do Sul, SC: Bom Jardim da Serra, Lages. Previously known distribution: Argentina, Uruguay. Main hosts: Heliantheae: Calea spp. (at least three spe- cies). Occasional hosts: Heliantheae: Aspilia sp., Verbesina subcordata. Plaumannimyia pallens Hering 4 samples, 10 individuals. Localities: RS: Guatba, SC: Lages, Matos Costa, Per- ico. Previously known distribution: Brazil (SC). Main hosts: Astereae: Baccharis dracunculifolia, Bac- charis punctulata, Baccharis uncinella, Baccharis spp. (at least two other species). Tomoplagia argentiniensis Aczél 39 samples, 300 individuals. Localities: ES: Linhares, MG: Serra do Cip6, Ouro Branco, RJ: Itatiaia, SP: Campinas, Ilha do Cardoso, Jundiat (Serra do Japi). Previously known distribution: Brazil (MG, RJ, SP), Argentina. Main host: Vernonieae: Cyrtocymura scorpioides. Tomoplagia biseriata (Loew) 4 samples, 29 individuals. Localities: MG: Diamantina, Grao Mogol, SC: Lages. Previously known distribution: Ecuador, Brazil. Main hosts: Heliantheae: Calea nitida, Calea oxyleps, Calea spp. (at least two other species). Tomoplagia costalimai Aczél 16 samples, 83 individuals. Localities: MG: Diamantina, Grao Mogol, Joaquim Felicio (Serra do Cabral), Serra do Cip6, RS: Guai- ba, SC: Lages, Matos Costa, SP: Jundiaf (Serra do Japi). Previously known distribution: Argentina. Main hosts: Mutisieae: Trixis mollissima, Trixis praes- tans, Trixis vauthieri, Trixis verbasciformis. Occasional hosts: Mutisieae: Jungia floribunda. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Tomoplagia fiebrigi Hendel 5 samples, 46 individuals. Localities: MG: Joaquim Felicio (Serra do Cabral), RS: Guaiba, SC: Lages. Previously known distribution: Paraguay, Argentina. Main hosts: Vernonieae: Vernonanthura aff. lucida, Vernonanthura nudiflora. Occasional host: Vernonieae: dieana. Vernonanthura twee- Tomoplagia formosa Aczél 13 samples, 45 individuals. Localities: MG: Caraga, Diamantina, Serra do Cip6o, PR: 20 km SE of Curitiba, RJ: Visconde de Maua, RS: Guatba, SC: Bom Jardim da Serra, Lages, Sao Joaquim. Previously known distribution: Brazil (MG), Argenti- na. Main hosts: Vernonieae: Chrysolaena flexuosa, Chry- solaena platensis, Lepidaploa salzmanii. Occasional hosts: Vernonieae: Chrysolaena herbacea, Lessingianthus (Oligocephalus) desertorum, Helian- theae: Verbesina sp. Tomoplagia incompleta (Williston) 132 samples, 512 individuals. Localities: MG: Diamantina, Grao Mogol, Joaquim Felicio (Serra do Cabral), Passa Quatro, Ouro Bran- co, Serra do Cipo, RJ: Itatiaia, RS: Capao Novo, Guaiba, SC: Bom Jardim da Serra, Lages, Matra, Perico. Previously known distribution: West Indies, Paraguay, Brazil (MG), Argentina. Main hosts: Vernonieae: Chresta sphaerocephala, Chrysolaena flexuosa, Chrysolaena herbacea, Chry- solaena platensis, Echinochoryne schwenkiaefolia, Lepidaploa lilacina, Lepidaploa spixiana, Lepida- ploa salzmanii, Lessingianthus (Oligocephalus) sim- plex, Lessingianthus carduoides, Lessingianthus li- nearis, Lessingianthus linearifolius, Lessingianthus psilophyllus, Lessingianthus pumillus, Lessingian- thus roseus, Lessingianthus stoechas, Lessingian- thus vepretorum. Occasional hosts: Eupatorieae: Chromolaena laeviga- ta, Heterocondylus alatus, Mikania sessilifolia, Mu- tisieae: Gochnatia amplexifolia, Trixis vauthieri, Vernonieae: Chresta sp., Eremanthus sp., Lepida- ploa sp., Lessingianthus (Oligocephalus) deserto- rum, Lessingianthus brevipetiolatus, Lessingianthus coriaceus, Lessingianthus glabratus, Lessingianthus laevigatus, Lessingianthus poliphyllus, Lessingian- thus rubricaulis, Lessingianthus sellowti, Lessin- gianthus warmingianus, Minasia sp., Vernonia in- cana, Vernonanthura glanduloso-dentata. Tomoplagia minuta Hering 48 samples, 648 individuals. Localities: MG: Diamantina, Formiga, Grao Mogol, Monte Verde, Ouro Branco, Parque Estadual de Ib- itipoca, Serra do Cip6, RJ: Itatiaia, Visconde de VOLUME 104, NUMBER 4 Maua, RS: Cambara do Sul, SC: Lages, Matos Cos- ta, Mafra. Previously known distribution: Brazil (MG, SP, SC). Main hosts: Vernonieae: Vernonanthura beyrichi, Ver- nonanthura catharinensis, Vernonanthura chamae- drys, Vernonanthura mariana, Vernonanthura mem- branacea, Vernonanthura montevidensis, Vernon- anthura phaeoneura, Vernonanthura phosphorica, Vernonanthura subverticillata, Vernonanthura wes- tiniana. Occasional host: Vernonieae: Vernonanthura petiolar- is, Vernonanthura tweedieana. Tomoplagia punctata Aczél 7 samples, 27 individuals. Localities: MG: Grao Mogol, Joaquim Felicio (Serra do Cabral), Ouro Branco, RJ: Itatiaia, Visconde de Maua, SP: Bertioga. Previously known distribution: Argentina. Main host: Vernonieae: Centratherum punctatum. Tomoplagia pseudopenicillata Aczél 14 samples, 55 individuals. Localities: MG: Ouro Branco, Serra do Cip6, SP: Ilha do Cardoso, Campinas. Previously known distribution: Brazil (MG, SP). Main host: Vernonieae: Cyrtocymura scorpioides. Tomoplagia reimoseri Hendel 57 samples, 1093 individuals. Localities: MG: Diamantina, Formiga, Grao Mogol, Joaquim Felicio (Serra do Cabral), Ouro Branco Serra do Cipo. Previously known distribution: Trinidad, Venezuela, Paraguay, Brazil, Argentina. Main hosts: Vernonieae: Vernonanthura glanduloso- Vernonanthura Ver- nonanthura phosphorica, Vernonanthura subverti- dentata, Vernonanthura mariana, membranacea, Vernonanthura mucronulata, cillata, Vernonanthura spp. (at least two other spe- cies). Occasional hosts: Astereae: Baccharis aphylla, Ver- nonieae: Cyrtocymura scorpioides, Minasia sp. Tomoplagia tripunctata Hendel 56 samples, 133 individuals. Localities: MG: Caraga, Diamantina, Ouro Branco, Parque Estadual de Ibitipoca, Serra do Cipo. Previously known distribution: Paraguay, Brazil. Main hosts: Vernonieae: Lessingianthus brevipetiola- tus, Lessingianthus carduoides, Lessingianthus li- nearifolius, Lessingianthus linearis, Lessingianthus psilophyllus, Lessingianthus roseus, Lessingianthus vepretorum. Occasional hosts: Mutisieae: Trixis vauthieri, Vernon- ieae: Lessingianthus pumillus, Lessingianthus ros- marinifolius, Lessingianthus stoechas, Lessingian- thus pycnostachyus. Tomoplagia trivittata (Lutz & Lima) 2 samples, 14 individuals. 1023 Localities: MG: Joaquim Felicio (Serra do Cabral), SP: Mogi Guacu. Previously known distribution: Brazil (MG, SP. MS), Argentina. Main hosts: Mutisieae: Gochnatia barrosii, Gochnatia floribunda. Trypanaresta coelestina (Hering) 14 samples, 76 individuals. Localities: MG: Diamantina, Ouro Branco, Passa Qua- tro, Serra do Cip6, PR: 25 km SE of Curitiba, RS: Cambara do Sul, Santo Antonio da Patrulha, SC: Lages. Previously known distribution: Brazil (PR, SC). Main hosts: Astereae: Baccharis dracunculifolia, Bac- charis helichrysoides, Baccharis leucopappa, Bac- charis brachylaenoides var. ligustrina, Baccharis trinervis. Occasional hosts: Astereae: Baccharis brachylaeno- ides, Baccharis ramosissima, Baccharis serrulata, Senecionieae: Senecio selloi. Trypanaresta imitatrix (Hering) 6 samples, 11 individuals. Localities: MG: Ouro Branco, Serra do Cipo6, PR: Parque Nacional de Foz do Iguacu, RS: Guaiba, SP: Campinas. Previously known distribution: Brazil (SP to SC), Ar- gentina. Main hosts: Astereae: Baccharis dracunculifolia, Bac- charis punctulata, Baccharis trinervis, Heterothal- amus psiadioides. Trypanaresta thomsoni (Hendel) 8 samples, 12 individuals. Localities: RJ: Marica, SC: Santa Cecilia, SP: Bertio- ga, Campinas. Previously known distribution: Bolivia, Argentina. Main hosts: Astereae: Conyza canadensis, Conyza bonariensis. Occasional hosts: Astereae: Solidago chilensis, Inu- leae: Pluchea sagitalis. Xanthaciura biocellata (Thomson) 165 samples, 1,506 individuals. Localities: MG: Diamantina, Grao Mogol, Joaquim Felicio (Serra do Cabral), Ouro Branco, Parque Es- tadual de Ibitipoca, Passa Quatro, Serra do Cipo, RJ: Engenheiro Passos, Itatiaia, Marica, Visconde de Maua, RS: Gravatai, SP: Bertioga, Campinas, Cam- pos do Jordao. Previously known distribution: Peru, Bolivia, Para- guay, Brazil, Uruguay, Argentina. Main hosts: Eupatorieae: Ageratum fastigiatum, Aya- pana amigdalina, Barrosoa_ betonicaeformis, Chromolaena_ barbacensis, Chromolaena chaseae, Chromolaena costatipes, Chromolaena cylindroce- phala, Chromolaena hookeriana, Chromo- Chromolaena decumbens, Chromolaena horminoides, laena laevigata, Chromolaena maximilianii, Chrom- olaena_ minasgeraesensis, Chromolaena pedalis, 1024 Chromolaena pungens, Chromolaena_ sagittifera, Chromolaena squalida, Heterocondylus alatus, Het- erocondylus amphidictyus, Heterocondylus jara- guensis, Koanophyllon adamantium, Praxelis cle- matidea, Symphyopappus cuneatus, Symphyopappus decussatus, Trichogonia hirtiflora, Trichogonia sal- viaefolia, Trichogonia villosa, Trichogoniopsis ad- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON three other species), Grazielia sp., Koanophyllon thysanolepis, Mikania cipoensis, Mikania cordifolia, Mikania decumbens, Mikania lasiandrae, Mikania micrantha, Mikania officinalis, Mikania sessilifolia, Symphyopappus decussatus, Symphyopappus ita- tiayensis, Vittetia orbiculata, Heliantheae: Aspilia pascalioides, Vernonieae: Eremanthus sp. enantha. Xanthaciura insecta (Loew) 3 samples, 31 individuals. Localities: MG: Diamantina, Joaquim Felicio (Serra do Cabral), Ouro Branco. Previously known distribution: USA, Mexico to Ve- nezuela, West Indies. Main hosts: Heliantheae: Bidens brasiliensis, Bidens sa, Chromolaena myriocephala, Chromolaena odor- sp. Occasional hosts: Eupatorieae: Acritopappus longifol- ius, Pseudobrickellia brasiliensis, Campuloclinium campuloclinoides, Campuloclinium chlorolepsis, Campuloclinium macrocephalum, Campuloclinium megacephalum, Chromolaena congesta, Chromo- laena matogrossensis, Chromolaena multiflosculo- ata, Chromolaena_ stachyophylla, Symphyopappus . . ee EE GRRE Xanthaciura mallochi Aczél| 3 samples, 10 individuals. Localities: MG: Parque Estadual de Ibitipoca. Previously known distribution: Costa Rica, Panama, Colobia, Ecuador, Peru, n. Argentina, s. Brazil. Main host: Eupatorieae: Chromolaena maximilianii. reticulatus, Symphyopappus att. reticulatus, Mutis- ieae: Inulopsis scaposa, Vernonieae: Lessingianthus (Oligocephalus) virgulatus, Lessingianthus hovae- folius, Lychnophora sp., Vernonanthura mariana. Xanthaciura chrysura (Thomson) 167 samples, 829 individuals. Localities: MG: Arceburgo, Camanducaia, Caraga, Di- amantina, between Formiga e Divinopolis, Grao Mogol, Joaquim Felicio (Serra do Cabral), Monte Verde, Ouro Branco, Parque Estadual de Ibitipoca, Passa Quatro, Serra do Cip6, RJ: Engenheiro Passos, Itatiaia, Visconde de Maua, RS: Cambara do Sul, Guaiba, Gravatat, SC: Bom Jardim da Serra, Lages, Matos Costa, Mafra, Perico, SP: Bertioga, Campi- tina. Main hosts: Eupatorieae: Ageratum conyzoides, Ager- Xanthaciura quadrisetosa (Hendel) 27 samples, 51 individuals. Localities: MG: Diamantina, Grao Mogol, Joaquim Felicio (Serra do Cabral), Ouro Branco, Parque Es- tadual de Ibitipoca, Passa Quatro, RS: Capao Novo, SC: Lages, RJ: Visconde de Maua, SP: Campos do Jordao, Piracicaba. Previously known distribution: Bolivia, Brazil, Argen- nas, Campos do Jordao, Ilha do Cardoso, Ilha Com- prida, Jundiai (Serra do Japi). Previously known distribution: USA (Florida), from atum fastigiatum, Campovassouria cruciata, Koan- ophyllon thysanolepis, Mikania purpurascens. Mexico to Argentina. Occasional hosts: Eupatorieae: Austroeupatorium sp., Grazielia intermedia, Symphyopappus casarettoi, Heliantheae: Bidens sp., Calea nitida. Main hosts: Eupatorieae: Adenostemma lavenia, Ag- eratum conyzoides, Ageratum fastigiatum, Ageratum sp., Austroeupatorium inulaefolium, Austroeupato- — Xanthaciura unipuncta Malloch rium paulinum, Austroeupatorium silphiifolium, 22 samples, 79 individuals. Barrosoa betonicaeformis, Campovassouria crucia- Localities: MG: Diamantina, Guaiba, Grao Mogol, Joaquim Felicio (Serra do Cabral), Ouro Branco, Serra do Cipo, RJ: Engenheiro Passos, Marica, SC: Mages, Perica, ‘S€; aces, Pericow Sb beriogas Campinas. Previously known distribution: Guatemala to Colom- bia and Trinidad, Paraguay, Brazil, Argentina. Main hosts: Eupatorieae: Campovassouria cruciata, Heliantheae: Aspilia cylindrocephala, Aspilia joly- ana, Aspilia spp. (at least four species), Wedelia pal- udosa. Occasional hosts: Astereae: Baccharis punctulata, Eu- patorieae: Barrosoa betonicaeformis, Chromolaena ta, Campuloclinium macrocephalum, Campulocli- nium purpurascens, Chromolaena chaseae, Chrom- olaena cylindrocephala, Chromolaena horminoides, Chromolaena laevigata, Chromolaena maximilianii, Chromolaena minasgeraesensis, Chromolaena pun- gens, Chromolaena squalida, Hatschbachiella twee- dieana, Heterocondylus alatus, Heterocondylus grandis, Heterocondylus jaraguensis, Grazielia gaudichaudeana, Grazilellia intermedia, Praxelis clematidea, Trichogonia hirtiflora, Trichogoniopsis adenantha. Occasional hosts: Austroeupatorium picturatum, Chromolaena barbacensis, Chromolaena hookeri- squalida, Mikania purpurascens, Trichogonia vil- losa, Heliantheae: Aspilia montevidensis, Calea elongata, Calea sp. Vernonieae: Cyrtocymura scor- ana, Chromolaena matogrossensis, Chromolaena multiflosculosa, | Chromolaena Chromolaena pedalis, Chromolaena spp. (at least myriocephala, pioides. VOLUME 104, NUMBER 4 1025 APPENDIX II Gazetteer This appendix lists the sites sampled in this study and their state (ES—Espirito Santo; MG—Minas Gerais: PR—Parana; RJ—Rio de Janeiro; RS—Rio Grande do Sul; SC—Santa Catarina; SP—-Sao Paulo), their code number in Fig. 1, and their geographical coordinates. Some localities have the same code number because they are not far enough apart to be represented as separate points in the map in Fig. 1. Code Locality State Coordinates l Guaiba RS 30sSSissi W. 2 Gravatat RS 29°55'S50°50'W 2) Santo Antonio da Patrulha RS 29°53'S50°50'W 3 Capao da Canoa RS 29°42'S50°20'W 4 Capao Novo RS 29°41'S49°49'W 5 Maquine RS 29°30'S50°50'’W 6 Torres RS 29°23'S49°49'W i Praia Grande SC 29°11'S49°49'W 8 Cambara do Sul RS 29°04'S50°38'W 9 Bom Jardim da Serra SG 28°18'S49°49'W 9 Lauro Miiller SE 28°26'S49°49'W 9 Perico KE 28°16'S49°49'W 9 Sao Joaquim Se 28°15'S49°49'W 10 Urupema SE 27°56'S49°49'W 11 Lages Xe 27°51'S50°36'W 12 Lebon Régis SC 26°55'S50°50'W 12 Santa Cecilia NE 26°47'S50°50'W 13 Cacador Se 26°46'S50°23'W 14 Matos Costa SC 26°28'S51°51'W 15 Matra Se 26°08'S49°49'W 16 Parque Nacional do Iguacu PR 25°30'S53°48'W 17, Curitiba PR 25°25'S49°15’W 18 Ilha do Cardoso SP 25°08'S47°58'’W 19 Bertioga SE 23°51'S46°09’W 20 Jundiat SP 23°11'S46°52'W 21 Campinas SP 22°59'S46°59'W 22 Marica RJ 22°55'S42°49'W 23 Camanducaia MG 22°46'S46°08'W 23 Monte Verde MG 22°52'S46°03’W 24 Piracicaba SP 22°42'S47°34'W 25 Campos do Jordao SP 22°41'S45°30'W 26 Engenheiro Passos RJ 22°30'S44°41'W 26 Itatiaia RJ 22°30'S44°34'W 26 Visconde de Maua RJ 22°19'S44°35'W De, Passa Quatro MG 22°23'S44°58'W 28 Mogi Guacu SP 22°15'S47°09'W 29 Pogos de Caldas MG 21°47'S46°34'W 30 Parque Estadual do Ibitipoca MG 21°43'S43°54'W 3 Arceburgo MG 21°22'S46°56’W 32 Ouro Branco MG 20°30'S43°43'W 33 Formiga MG 20°28'S45°26'W 34 Divinopolis MG 20°08'S44°53’W 35 Caraga MG 20°08'S43°30'W 36 Linhares ES 19°25'S40°04'W Bh) Serra do Cip6 MG 19°15'S43°43'W 38 Diamantina MG 18°13'S43°43'W 39 Joaquim Felicio MG 17°43'S44°44'W 40 Grao Mogol MG 16°34'S42°42'W 1026 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON APPENDIX III Full Scientific Names of Plants Cited in Appendix I This appendix lists the full scientific names (including authority) of the species of host plants cited in the Appendix I. Acritopappus longifolius (Gardner) R.M. King & H. Rob. Adenostemma lavenia (L.) Kuntze Ageratum conyzoides L. Ageratum fastigiatum (Gardner) R.M. King and H. Rob. Aspilia cylindrocephala H. Rob. Aspilia foliacea Baker Aspilia jolyana Barroso Aspilia laevissima Baker Aspilia pascalioides Griseb. Austroeupatorium inulaefolium (Kunth) R.M. King & H. Rob. Austroeupatorium paulinum (DC.) R.M. King & H. Rob. Austroeupatorium picturatum (Malme) R.M. King & H. Rob. Austroeupatorium silphiifolium (Mart.) R.M. King & H. Rob. Ayapana amygdalina (Lam.) R.M. King & H. Rob. Baccharis aphylla Sch. Bip. Baccharis brachylaenoides var. ligustrina (DC.) Maguire & Wurdack Baccharis dracunculifolia DC. Baccharis helichrysoides DC. Baccharis leucopappa DC. Baccharis punctulata DC. Baccharis ramosissima Gardner Baccharis serrulata DC. Baccharis trinervis Pers. Baccharis uncinella DC. Barrosoa betonicaeformis (DC.) R.M. King & H. Rob. Bidens brasiliensis Sherff Bidens pilosa L. Calea elongata Baker Calea graminifolia Sch.-Bip. ex Kraschen. Calea nitida Less. Calea oxyleps Baker Campovassouria cruciata (Vell.) R.M. King & H. Rob. Campuloclinium campuloclinoides (Baker) R.M. King & H. Rob. Campuloclinium chlorolepis (Baker) R.M. King & H. Rob. Campuloclinium macrocephalum (Less.) DC. Campuloclinium megacephalum (Mart. ex Baker) R.M. King & H. Rob. Campuloclinium purpurascens (Sch. Bip. ex Baker) R.M. King & H. Rob. Centratherum punctatum Cass. Chresta sphaerocephala DC. Chromolaena ascendens (Sch. Bip. ex Baker) R.M. King & H. Rob. Chromolaena barbacensis (Hieron.) R.M. King & H. Rob. Chromolaena chaseae (B.L. Rob.) R.M. King & H. Rob. Chromolaena congesta (Hook. & Arn.) R.M. King & H. Rob. Chromolaena costatipes (B.L. Rob.) R.M. King & H. Rob. Chromolaena cylindrocephala (Sch. Bip. ex Baker) R.M. King & H. Rob. Chromolaena decumbens Gardner Chromolaena hockeriana (Griseb.) R.M. King & H. Rob. Chromolaena horminoides DC. Chromolaena laevigata (Lam.) R.M. King & H. Rob. Chromolaena matogrossensis (Hieron.) R.M. King & H. Rob. Chromolaena maximilianii (Hieron.) R.M. King & H. Rob. Chromolaena minasgeraesensis (Hieron.) R.M. King & H. Rob. Chromolaena multiflosculosa (DC.) R.M. King & H. Rob. VOLUME 104, NUMBER 4 1027 APPENDIX III. Continued. Chromolaena myriocephala (Gardner) R.M. King & H. Rob. Chromolaena odorata (L.) R.M. King & H. Rob. Chromolaena pedalis (Sch. Bip. ex Baker) R.M. King & H. Rob. Chromolaena pungens (Gardner) R.M. King & H. Rob. Chromolaena sagittifera (B.L. Rob.) R.M. King & H. Rob. Chromolaena squalida (DC.) R.M. King & H. Rob. Chromolaena stachyophylla (Spreng.) R.M. King & H. Rob. Chrysolaena flexuosa (Sims) H. Rob. Chrysolaena herbacea (Vell.) H. Rob. Chrysolaena platensis (Spreng.) H. Rob. Conyza bonariensis (L.) Cronquist Conyza canadensis (L.) Cronquist Cyrtocymura scorpioides (Lam.) H. Rob. Echinochoryne schwenkiaefolia (Mart.) H. Rob. Gochnatia amplexifolia (Gardner) Cabrera Gochnatia barrosti Cabrera Gochnatia floribunda Cabrera Grazielia gaudichaudeana (DC.) R.M. King & H. Rob. Grazilelia intermedia (DC.) R.M. King & H. Rob. Hatschbachiella tweedieana (Hook. & Arn.) R.M. King & H. Rob. Heterocondylus alatus (Vell.) R.M. King & H. Rob. Heterocondylus amphidictyus (DC.) R.M. King & H. Rob. Heterocondylus grandis (Sch. Bip. ex Baker) R.M. King & H. Rob. Heterocondylus jaraguensis (B.L. Rob.) R.M. King & H. Rob. Heterothalamus psiadioides Less. Inulopsis scaposa (Remy) Hoffm. Jungia floribunda Less. Koanophyllon adamantium (Gardner) R.M. King & H. Rob. Koanophyllon thysanolepis (B.L. Rob) R.M. King & H. Rob. Lepidaploa lilacina (Mart. ex DC.) H. Rob. Lepidaploa salzmanii (DC.) H. Rob. Lepidaploa spixiana (Mart. ex DC.) H. Rob. Lessingianthus (Oligocephalus) desertorum (Mart. ex DC.) H. Rob. Lessingianthus (Oligocephalus) simplex (Less.) H. Rob. Lessingianthus (Oligocephalus) virgulatus (Mart. ex DC.) H. Rob. Lessingianthus brevipetiolatus (Sch. Bip. ex Baker) H. Rob. Lessingianthus buddleiifolius (Mart. ex DC.) H. Rob. Lessingianthus carduoides (Baker) H. Rob. Lessingianthus coriaceus (Less.) H. Rob. Lessingianthus glabratus (Less.) H. Rob. Lessingianthus hovaefolius (Gardn.) H. Rob. Lessingianthus laevigatus (Mart. ex DC.) H. Rob. Lessingianthus linearifolius (Less.) H. Rob. Lessingianthus linearis (Spreng.) H. Rob. Lessingianthus poliphyllus (Sch. Bip. ex Baker) H. Rob. Lessingianthus psilophyllus (DC.) H. Rob. Lessingianthus pumillus (Vell.) H. Rob. Lessingianthus pycnostachyus (DC.) H. Rob. Lessingianthus roseus (Mart. ex DC.) H. Rob. Lessingianthus rosmarinifolius (Less.) H. Rob. Lessingianthus rubricaulis (Humb. & Bonpl.) H. Rob. Lessingianthus sellowii (Less.) H. Rob. Lessingianthus stoechas (Mart. ex Baker) H. Rob. Lessingianthus vepretorum (Mart. ex DC.) H. Rob. Lessingianthus warmingianus (Baker) H. Rob. Mikania cipoensis Barroso 1028 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON APPENDIX III. Continued. Mikania cordifolia (L. f.) Willd. Mikania decumbens Malme Mikania lasiandrae DC. Mikansia micrantha H.B.K. Mikania officinalis Matt. Mikania purpurascens (Baker) R. King & H. Rob. Mikania retifolia Sch. Bip. ex Baker Mikania sessilifolia DC. Moquinia racemosa (Spreng.) DC. Pluchea laxiflora Hook. & Arn. ex Baker Pluchea sagitalis (Lam.) Cabrera Porophyllum riedelii Baker Porophyllum ruderale (Jacq.) Cass. Praxelis clematidea (Griseb.) R. King & H. Rob. Proteopsis argentea Mart. & Zucc. ex Sch. Bip. Pseudobrickellia brasiliensis (Spreng.) R. King & H. Rob. Senecio brasiliensis (Spreng.) Less. Senecio oxyphyllus DC. Senecio selloi (Spreng.) DC. Solidago chilensis Meyen Stomathanthes polycephalus (Sch. Bip. ex Robinson) H. Rob. Symphyopappus casarettoi B. Robinson Symphyopappus cuneatus (DC.) Sch. Bip. ex Baker Symphyopappus decussatus Turcz. Symphyopappus itatiayensis (Hieron.) R. King & H. Rob. Symphyopappus reticulatus Baker Trichogonia hirtiflora (DC.) Sch. Bip. ex Baker Trichogonia salviaefolia Gardner Trichogonia villosa Sch. Bip. ex Baker Trichogoniopsis adenantha (DC.) R. King & H. Rob. Trixis mollissima D. Don Trixis praestans (Vell.) Cabrera Trixis vauthieri DC. Trixis verbasciformis Less. Verbesina subcordata DC. Vernonanthura beyrichi (Less.) H. Rob. Vernonanthura catharinensis (Cabrera) H. Rob. Vernonanthura chamaedrys (Less.) H. Rob. Vernonanthura glandulosa-dentata (Hieron.) H. Rob. in litt. Vernonanthura mariana (Mart. ex Baker) H. Rob. Vernonanthura membranacea (Gardn.) H. Rob. Vernonanthura montevidensis (Spreng.) H. Rob. Vernonanthura mucronulata (Less.) H. Rob. Vernonanthura nudiflora (Less.) H. Rob. Vernonanthura petiolaris (DC.) H. Rob. Vernonanthura phaeoneura (Toledo) H. Rob. Vernonanthura phosphorica (Vell.) H. Rob. Vernonanthura subverticillata (Sch. Bip. ex Baker) H. Rob. Vernonanthura tweedieana (Baker) H. Rob. Vernonanthura westiniana (Less.) H. Rob. Vernonia incana Less. Vittetia orbiculata (DC.) R. King & H. Rob. Wedelia paludosa DC. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 1029-1035 A NEW SPECIES OF FRUMENTA BUSCK (LEPIDOPTERA: GELECHIIDAE: GNORIMOSCHEMINI) FROM MEXICO: A POTENTIAL BIOCONTROL AGENT AGAINST SOLANUM ELAEAGNIFOLIUM (SOLANACEAE) DAVID ADAMSKI AND JOHN W. BROWN Systematic Entomology Laboratory, PSI, Agriculture Research Service, U.S. Depart- ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0168, U.S.A. (e-mail: dadamski@sel.barc.usda.gov; jbrown @sel.barc.usda.gov) Abstract.—Frumenta solanophaga, new species, is described from San Luis Potosi, México. It was reared from Solanum elaeagnifolium Cavanilles (Solanaceae) as part of a survey for potential biocontrol agents against this introduced weed in southern Africa. Frumenta also includes two previously described species from southern United States; the group appears to feed exclusively on Solanum. Photographs and illustrations of F. solan- ophaga depict the imago, wing venation, intersegmental abdominal lobes of the female, and male and female genitalia. A key to the species of Frumenta is provided. Key Words: Frumenta was proposed by Busck (1939) to accommodate the single species, Gele- chia Zeller, 1873. Hodges (1983) synonymized Gelechia beneficentel- la Murtfeld, 1881, with Gelechia nundinel- la and transferred Asapharcha nephelom- icta Meyrick, 1930, to Frumenta. Since then, the genus has included only the two aforementioned species, which are known only from the United States. The genus be- longs to the tribe Gnorimoschemini. Since its description, Frumenta nundi- nella has been known as a herbivore of the nundinella noxious weed, horse-nettle (Solanum caro- linense Linnaeus; Solanaceae), and the moth has been cited repeatedly as a poten- tial biological control agent against this weed in the United States and Canada (e.g., Murtfeld 1881; Forbes 1923; Foott 1967; Bailey and Kok 1976, 1982; Bailey 1977). Solomon (1980, 1981, 1983) presented de- tails of the life history of F. nundinella, in- cluding hosts and herbivore relationships Gelechiidae, Solanum, biocontrol and phenology. Frumenta nephelomicta, which galls the stems and fruit of S. elaeag- nifolium Cavanilles, was released for the bi- ological control of this weed in South Af- rica (Neser et al. 1990). During recent efforts to discover new po- tential biological control agents against So- lanum elaeagnifolium, specimens of a un- described species of Frumenta where reared in San Luis Potosi, México, by personnel associated with the Plant Protection Re- search Institute, Pretoria, South Africa, and sent to the Systematic Entomology Labo- ratory, USDA, for identification. The pur- poses of this paper are to describe this new species and to summarize information on host plant usage in Frumenta in general. Adult vestiture was examined using an incandescent light source (reflected light). Kornerup and Wanscher (1978) was used as a color standard for the description of Fru- menta solanophaga. Genitalia were dissect- ed as described by Clarke (1941), except 1030 I Fig. |. Holotype of Frumenta solanophaga. the ventral part of the male genitalia was separated from the dorsal part on one side. The free part then was swung across the tegumen and mounted flat. Mercurochrome and chlorazol black were used as stains. Slide preparations were examined with dis- secting and compound microscopes. Ter- minology for genitalia and wing venation follows that of Polvolny (1991) and Powell and Polvolny (2001). KEY TO ADULTS OF FRUMENTA 1. Forewing white, intermixed with pale-yellow scales, with few small, pale-brown discal and MTEL NOUS 4 no acu sooo 00 nephelomicta — Forewing pale gray, intermixed with pale-yel- low scales, with many large gray discal and IMARSinal ¥SpPotseeseee che os ale ee eee eee 2 Be SIN EGS ates cock ost Belch puch cetibo ka Ree Te CREAT he cncy a ta 3 =e Remale see ea Scan a eet Oe acre atew ican: 4 3. Valva beyond basal arch long, gnathos about 12 width of base of uncus (Fig. 3) solanophaga — WValva beyond basal arch short, gnathos about Ya widthiot base Of WNCuS ya eee nundinella PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4. Ductus bursae about three times length of apophyses anteriores; apophyses anteriores narrow throughout length ......... nundinella — Ductus bursae about two times length of apophyses anteriores; apophyses anteriores di- latedibasallly.(Gio4e5)) eee eee solanophaga Frumenta solanophaga Adamski and Brown, new species (Figs. 1-5) Diagnosis.—Frumenta solanophaga is superficially most similar to F. nundinella, but it is genitally most similar to F. nephe- lomicta. Frumenta solanophaga shares with F. nephelomicta the following: valva elon- gate-rectangular beyond the basal arch; gnathos widened in distal junction of arms; and basal half of the apophyses anteriores dilated. Frumenta solanophaga differs from F. nephelomicta by having a darker fore- wing pattern, deeper basal arch of valva, distally wider valva, a longer aedeagus, and stouter apophyses anteriores. VOLUME 104, NUMBER 4 Description, adult.—Head: Vertex and frontoclypeus pale yellowish brown or pale gray; scape pale gray, intermixed with pale- brown scales, flagellum gray; outer surface of labial palpus pale brown, intermixed with white scales, inner surface white; pro- boscis white. Thorax: Tegula and mesonotum pale yel- lowish brown; legs with gray scales nar- rowly tipped with pale yellow, tarsomeres banded with pale yellowish-brown scales apically. Forewing (Figs. 1-2): Length 10.0-12.2 mm (mean = 11.6; n = 9). Ground color pale yellowish brown, inter- mixed with white and few pale-gray scales; darker gray and brown scales, each tipped with pale yellow form small costal spots, 2—3 larger, irregularly shaped discal spots, and large submarginal spots; in some spec- imens ground color as above with area pos- terior to CuP gray, or wing gray, intermixed with few pale yellow scales. Venation with R,— Rs; stalked about % distance from cell; anterodistal part of cell slightly attenuate; R, and medial veins divergent; CuA,—CuA, long, nearly parallel. Undersurface with gray scales, narrowly tipped with pale yel- low, intermixed with pale-yellow scales near apex. Hindwing (Figs. 1-2): Shiny gray. Venation with Sc + R,, Rs, and M, divergently arched from M, and M,; Sc + R, and radial branch connected by a short cross vein near % length of cell; distal mar- gin of cell concave inwardly; Rs and M, originating near anterodistal part of cell, slightly divergent, each juxtaposed to apex; M, closer to M, at base, arching closer to M, from % length to margin; CuA,—CuA, subparallel. Abdomen: Pale gray scales, intermixed with white scales; male terga 1-3 with brownish-orange scales; female interseg- mental membrane with large lateral lobe anterior to spiracle on seventh segment (Fig. 4). Male genitalia (Fig. 3): Uncus trapezoidal, lateral margins slightly curved ventrally, setose submarginally; gnathos elongate, flat, rounded apically; valva short, slightly narrowed submedially, distal lobe 1031 sparsely setose, blunt apically; vinculum elongate, acutely curved anterolaterally; ae- deagus elongate, bulbous at base, parallel- sided and narrow distally. Female genitalia (Fig. 5): Papillae anales with long setae in- termixed with short setae; apophyses pos- teriores slender; eighth sternum and tergum fused forming a sclerotized ring, extended anterolaterally into a pair of broad apoph- yses anteriores, and extended ventromedi- ally, forming a broadly rounded, shallowly invaginated antrum; ductus bursae elongate, swollen near inception of ductus seminalis; swollen area with a small, internal, semi- circular support; corpus bursae elongate, wider anteriorly than posteriorly, with an elongate signum on posterior end. Holotype.—¢, ‘““MEXICO, San Luis Po- tosi, San Luis de la Paz, 21.19N, 100.32W, 10.1x.1999, H. G. Zimmermann, AcSN 2109," “‘Emerged in quarantine from ber- ries of Solanum elaeagnifolium (Solanace- ae), Larvae eat seeds and flesh,’ ““National Collfection] of Insects, Pretoria, S[outh] Afrlica].”” Holotype not dissected. Depos- ited in the National Museum of Natural History Museum, Smithsonian Institution, USA [USNM]. Paratypes.—5 3,3 2, same label data as holotype, except, ““d genitalia slide by D Adamski, USNM 81218” [green label], “*d genitalia slide by D Adamski, USNM 82133” [green label], ““d wing slide by D Adamski, USNM 82134” [green label], **? genitalia slide by D Adamski, USNM 81219” [green label], 2 genitalia slide by D Adamski, USNM 81220” [green label], *“® genitalia slide by D Adamski, USNM 82132” [green label]. Five paratypes de- posited in USNM; one paratype deposited in the Transvaal Museum, Pretoria, South Africa; one paratype deposited in The South African Museum, Capetown, South Africa. Etymology.—The specific epithet is de- rived from the plant genus, Solanum, to which the host of F. solanophaga belongs, and from the Greek ‘“‘phagein,”’> meaning, “tor eat.” 1032 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Se RI VOLUME 104, NUMBER 4 1033 Fig. 4. Frumenta solanophaga. Lateral lobes of in- tersegmental membrane anterior to spiracle of seventh segment (female). DISCUSSION Association with Solanaceae.—The lar- val use of Solanaceae is common in the ge- lechiid tribe Gnorimoschemini, where many species are known to be pests of solana- ceousscrops~(Povolny 1973, 1975), ee. Scrobipalpa absoluta (Meyrick) on tomato (Lycopersicon esculentum Mill.) in northern South America; Scobipalpopsis solanivora Povolny on potato (Solanum tuberosum L.) in Central America; Phthorimaea isochlora Meyrick on “‘friegaplato” (Solanum sapon- aceum Duy) in Colombia; Phthorimaea op- erculella Zeller on potato throughout the New World; Keiferia lycopersicella (Wal- singham) on tomato throughout the New World; Keiferia colombiana Povolny on friegaplato in Colombia; and Symmetrische- ma tangolias (Gyen) (= S. plaesiosema (Turner)) on Solanum nigrum L. in the United States and presumably on Solanum in Australia. Members of Frumenta apparently are specialists on the plant genus Solanum. Frumenta nundinella, which occurs throughout southern and midwestern United Fig. 5. Frumenta solanophaga. Female genitalia. States (1.e., Georgia, Hlinois, Indiana, Lou- isiana, Missouri, Pennsylvania, Texas, and Virginia) is widely recognized as an impor- tant herbivore of Solanum carolinense. Fru- 1034 menta nephelomicta, which is known from Arizona and New Mexico, has been reared from Solanum elaeagnifolium and was in- troduced into South Africa for the biologi- cal control of this weed. An apparently un- described species of Frumenta from Texas and New Mexico also has been reared from S. elaeagnifolium (specimen data from USNM). The discovery of F. solanophaga as a herbivore of S. elaeagnifolium adds further support to the hypothesis that the genus is closely associated with Solanum. The biology of F. solanophaga 1s un- known. According to Bailey and Kok (1976), the larvae of Frumenta nundinella attack at least two stages of Solanum car- olinense. They feed on the tips of the young leaves before formation of fruits and pupate in characteristic leaf chambers in mid-July. After the plants fruit in late July, the larvae are fruit borers; one larva is capable of de- stroying all the seeds within a berry. The larvae pupate inside the berries, emerging as adults in August and September. In con- trast, the larvae of Frumenta nephelomicta have been described as gallers of stems and fruit (Neser et al. 1990). Potential use in biological control.—So- lanum elaeagnifolium goes by a variety of common names, including bull-nettle, silver horse-nettle, white-horse-nettle, silverleaf nightshade, silver-leaf-nettle, trompillo (Brako et al. 1995), and satansbos (the latter used primarily in South Africa). It is a per- sistent, prickly, perennial evergreen shrub native to the New World (Goeden 1972) that has been introduced inadvertently into southern Africa and Australia, where it has become an economically important weed (Siebert 1975, Wells et al. 1986). It may poison livestock where it occurs in areas used for grazing (Kingsbury 1964, Parker 1990). In South Africa, several studies have been conducted on potential biological con- trols for S. elaeagnifolium (Siebert 1975; Olckers and Hulley 1989a, b, 1991a, b). The status of the biological control of this weed in South Africa is summarized by Neser et al. (1990). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON In the mid 1980s Frumenta nephelomicta was introduced into South Africa for con- trol of S. elaeagnifolium. According to Ju- lien (1987), the insect failed to establish, apparently owing to severe drought and small releases; however, further releases are intended. The source of the specimens of F. nephelomicta released was not indicated, but if it was México (rather than south- western United States), then ““F. nephelom- icta’’ was most likely a misidentification of F. solanophaga. Because of possible dif- ferences in feeding habits among species of Frumenta, accurate identification of this po- tential biocontrol organism 1s critical. ACKNOWLEDGMENTS We thank V. M. Uys, Plant Protection Research Institute, Pretoria, South Africa, for sending the reared series of Frumenta solanophaga to the Systematic Entomology Laboratory (SEL), USDA, for identifica- tion; Ronald W. Hodges, retired SEL, for helping with the generic placement; and John Steiner, Office of Imaging, Printing and Photographic Services, Smithsonian In- stitution, for the photograph of the holo- type. We thank Ronald Hodges and Sonja Scheffer (SEL) for reviewing the manu- script. LITERATURE CITED Bailey, T. E. 1977. Recent development in the study of the gelechiid moth, Frumenta nundinella. Vir- ginia Journal of Science 28: 49 (abstract). Bailey, T. E. and L. T. Kok. 1976. Occurrence of Fru- menta nundinella (Lepidoptera: Gelechiidae) in southwest Virginia, and its potential as a biocon- trol agent of horsenettle. Virginia Journal of Sci- ence 27: 31 (abstract). . 1982. Biology of Frumenta nundinella (Lep- idoptera: Gelechiidae) on horsenettle in Virginia. Canadian Entomologist 114: 139-144. Brako, L., A. Y. Rossman, and D. F Farr. 1995. Sci- entific and Common Names of 7,000 Vascular Plants in the United States. APS Press, The Amer- ican Pytopathological Society, St. Paul, Minne- sota. 295 pp. Busck, A. 1939. Restriction of the genus Gelechia (Lepidoptera: Gelechiidae), with description of new genera. Proceedings of the United States Na- tional Museum 86: 563-593. VOLUME 104, NUMBER 4 Clarke, J. EF G. C. 1941. The preparation of the slides of the genitalia of Lepidoptera. Bulletin of the Brooklyn Entomological Society 36: 149-161. Foott, W. H. 1967. Occurence of Frumenta nundinella (Lepidoptera: Gelechiidae) in Canada. Canadian Entomologist 99: 443—444. Forbes, W. T. M. 1923. Lepidoptera of New York and neighboring states. Cornell University Agricultur- al Experiment Station, Memoir 68, 729 pp. Goeden, R. D. 1972. Insect Ecology of silverleaf night- shade. Weed Science 19: 45—51. Hodges, R.W. 1983. Gelechioidea, pp. 11-25. In Hodges, R. W. et al., eds. Check list of the Lepi- doptera of America North of Mexico. E. W. Clas- sey Ltd. and the Wedge Entomological Research Foundation, London, xxiv + 284 pp. Julien, M. H. 1987. Biological control of weeds: a world catalogue of agents and their target weeds, 2nd ed. C.A.B. International, Wallingford, 144 pp. Kingsbury, J. M. 1964. Poisonous Plants of the U.S. and Canada. Englewood Cliffs, N.J. Prentice-Hall Inc. + x11 + 626 pp. Kornerup, A. and J. H. Wanscher. 1978. Methuen Handbook of Colour. Third Ed. Methuen and Co., Ltd., London. 252 pp. Meyrick, E. 1930. Exotic Microlepidoptera 3(8): 545— 576. Murtfeldt, M. E. 1881. New species of Tineidae. Ca- nadian Entomologist 13: 242-246. Neser, S., H. G. Zimmerman, H. E. Erb, and J. H. Hoffman. 1989. Progress and prospects for the bi- ological control of two Solanum weeds in South Africa, pp. 371-381. In Delfosse, E. S., ed. Pro- ceedings of the 7th International Symposium on the Biological Control of Weeds, 1988, Rome. 1 XX ste Ol pp: Olckers, T. and P. E. Hulley. 1989a. Insect herbivore diversity on the exotic weed Solanum mauritian- um Scop. and three other Solanum species in the eastern Cape. Journal of the Entomological Soci- ety of South Africa 52: 81—93. . 1989b. Seasonality and biology of common insect herbivores attacking Solanum species in the Eastern Cape Province. Journal of the Entomolog- ical Society of Southern Africa 52: 109-118. . 1991a. Impoverished insect herbivore faunas on the exotic bugweed, Solanum mauritianum Scop. relative to indigenous Solanum species in Natal/KwaZulu and the Transkei. Journal of the Entomological Society of Southern Africa. 54: 39-50. 1035 . 1991b. Notes on some insect galls associated with Solanum plants in South Africa. South Af- rican Journal of Zoology 26: 55—61. Parker, K. E 1990. An Illustrated Guide to Arizona Weeds. University of Arizona Press, Tucson, Ar- izona. 1 + xii + 338 pp. Povolny, D. 1973. Scrobopalpopsis solanivora sp. 0.— A new pest of potato (Solanum tuberosum) from Central America. Acta Universitatis Agriculturae 21: 133-146. . 1975. On three neotropical species of Gnori- moschemini (Lepidoptera, Gelechiidae) mining Solanaceae. Acta Universitatis Agriculturae 23: 379-393. . 1991. Morphologie, systematik und phyloge- ny der tribus Gnorimoschemini (Lepidoptera, Ge- lechiidae). Acta Sci. Nat. Acad. Sci. Bohemoslov. (Brno) (new series) 25: 1—103. Polvolny, D. and J. Powell. 2001. Gnorimoschemine moths of coastal dune and scrub habitats in Cali- fornia (Lepidoptera: Gelechiidae). Holarctic Lep- idoptera 8, Supplement 1: 1—53 Siebert, M. W. 1975. Candidates for the biological con- trol of Solanum elaeagnifolium Cay. (Solanaceae) in South Africa. 1. Laboratory studies on the bi- ology of Gratiana lutescens (Boh.) and Gratiana pallidula (Boh.) (Coleoptera: Cassididae). Journal of the Entomological Society of Southern Africa 38: 297-304. Solomon, B. P. 1980. Frumenta nundinella (Lepidop- tera: Gelechiidae): Life history and induction of host parthenocarpy. Environmental Entomology 9: 821-825. . 1981. Response of a host-specific herbivore to resource density, relative abundance, and phe- nology. Ecology 62(5): 1205-1214. . 1983. Compensatory production in Solanum carolinense following attack by a_ host-specific herbivore. Journal of Ecology 71: 681—690. Wells, M. J., A. A. Balsinhas, H. Joffe, V. M. Engel- brecht, G. Harding, and C. H. Stirton. 1986. A Catalogue of Problem Plants in Southern Africa. Memortrs of the Botanical Survey of South Africa. No. 53. Botanical Research Institute, Department of Agriculture and Water Supply, South Africa. 658 pp. Zeller, PB. C. 1873. Beitrége zur Kenntiss der norda- mericanischen Nachtfalter besonders der Micro- lepidopteren. Zweite Abtheilung. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 23: 201-334, pl. I-IV. PROC. ENTOMOL. SOC. WASH. 104(4), 2002, pp. 1036—1054 DESCRIPTIONS OF THE IMMATURE STAGES OF FIVE MEXICAN SPECIES OF GYMNETINI (COLEOPTERA: SCARABAEIDAE: CETONIINAE) MIGUEL ANGEL MORON AND ROBERTO ARCE Departamento de Entomologia, Instituto de Ecologia, A.C. (SEP-CONACYT). Apdo. Postal 63, Xalapa, Veracruz 91000, México (e-mail: moron-_ma@ecologia.edu.mx) Abstract.—The third instar larva of Amithao haematopus (Burmeister) from Chiapas, México, is described as the first description of a larva in this genus. The third instar larvae and pupa of Marmarina maculosa (Olivier) from Veracruz, México are described. The third instar larvae and pupa of Hoplopyga liturata (Olivier) from Veracruz are described, including comments on first and second instar larvae. The third instar larva of Gymnetis hebraica difficilis Burmeister from Veracruz, is described. The larva of Gymnetis flavo- marginata sallei Schaum is redescribed with specimens from Veracruz, including differ- ences with second and first instar larvae and the description of the pupa. All of these species are included in a key to the larvae of New World Gymnetini, which now has 14 species in nine genera. Resumen.—Se describe la larva de tercer estadio de Amithao haematopus (Burmeister) de Chiapas, México, que representa la primera descripcion larvaria para el género. Se describen la larva de tercer estadio y la pupa de Marmarina maculosa (Olivier) de Ve- racruz, México. Se describen la larva y la pupa de Hoplopyga liturata (Olivier) con ejemplares de Veracruz, incluyendo comentarios sobre las larvas de primero y segundo estadio. Se describe la larva de tercer estadio de Gymnetis hebraica difficilis Burmeister de Veracruz. Se redescribe la larva de tercer estadio de Gymnetis flavomarginata sallei Schaum, con ejemplares de Veracruz, incluyendo las diferencias con las larvas de primero y segundo estadio, y la descripci6n de la pupa. Todas estas especies se incluyen en una clave para las larvas de los Gymnetini del Nuevo Mundo, que ahora contiene 14 especies de nueve géneros. Key Words: Amithao, Marmarina, Hoplopyga, Gymnetis, larvae, pupae, taxonomy, key, México Ritcher (1966) described the third instar (1984) described the larva of one species of larvae of four species of the genera Cotinis Burmeister, Gymnetis MacLeay and Gym- netina Casey from the United States. Mon- né (1969) described the larvae of two spe- cies of Blaesia Burmeister and Marmarina Kirby from Uruguay. Moron and Ratcliffe Argyripa Thomson from México, and a key to the seven then-known species of New Word Gymnetini. Vanin and Costa (1984) described the larva of one species of Ho- plopyga Thomson from Brazil. Mic6 et al. (2001) provided descriptions of the larva of VOLUME 104, NUMBER 4 one species of Hologymnetis Martinez from México and another species of Hoplopyga from Brazil, with a key to larvae of the 10 then-known species of Gymnetini from the Americas. In this paper, we describe for the first time the third instar larvae of one species of the genus Amithao; the third instar larvae of three species of Marmarina, Gymnetis and Hoplopyga:; the first and second instar larvae of two species of Gymnetis and Ho- plopyga; and the pupae of three species of Gymnetis, Marmarina and Hoplopyega. Also, we redescribe the larva of third instar of Gymnetis flavomarginata sallei Schaum, and present a new key to the third instar larvae of 14 known species and nine genera of Gymnetini. Technical terms are those of Ritcher (1966), Moron (1993), and Micé et al. (2001). Studied specimens are deposited in the collection of immature stages of the Departamento de Entomologia, Instituto de Ecologia, Xalapa, México (IEXA). Relative to the 3100 species of Cetoni- inae listed from the world, only 55 larvae representing 30 genera of Cetoniini, Gym- netini, Goliathini and Cremastoceilini are described at present. Consequently, it is dif- ficult to get a set of diagnostic characters that aid in distinguishing the larvae of the tribe Gymnetini from the larvae of other tribes. But as a preliminary introduction, based on Ritcher (1966), Mic6 et al. (2001), and our Own experience, we propose the following combination of characters as di- agnostic for larvae of Gymnetini: frons with 1—2 posterofrontal setae at each side; labrum symmetrical, anterior border tri- lobed; clithra present; plegmata absent; haptomerum with a transverse row of 7—19 heli; maxillary stridulatory area consisting of a row of 3—9 curved teeth with anteriorly projecting points; ocelli clearly defined, vague or absent; last antennal segment with 2—15 dorsal sensory spots; tarsungulus cy- lindrical, rounded apically, bearing 5—15 se- tae; raster usually with elliptical monosti- chous or polystichous palidia, each pali- dium with 12—38 pali. 1037 KEY TO THE KNOWN THIRD STAGE LARVAE OF GYMNETINI FROM THE AMERICAS. (MODIFIED FROM MICO ET AL. 2001.) i) 1. Palidia present — Palidia absent .. Gymnetina cretacea (LeConte) tN Raster with each palidium consisting of 2 or more irregular rows of pali. Last antennal seg- ment with 3—7 dorsal sensory spots. ...... 3 — Raster with palidia monostichous. Last anten- nal segment with 2—15 dorsal sensory spots 3. Tarsungulus with 7 setae. Maxillary stridula- tory area with 5 teeth. Last antennal segment with 3 ventral sensory spots ......... .. . Hologymnetis cinerea (Gory and Percheron) — Tarsungulus with 10-12 setae. Maxillary stridulatory area with 7—9 teeth. Last segment of antenna with 5—13 ventral sensory spots 4. Raster with inner row of each palidium hav- ing 7-10 pali much stouter and larger than those in outer row Cotinis mutabilis (Gory and Percheron) — Raster with inner row of each palidium hav- ing 9-10 pali slightly larger than those in out- EL TOW Verein is 5 ches meee Cotinis nitida (Linné) 5. Dorsum of abdominal segment VII with 3 an- nulets. Last antennal segment with 10-15 dorsalisensony Spots/as st se ae el oo 6 — Dorsum of abdominal segment VII with 2 an- nulets. Last antennal segment with 2—5 dorsal SENSOLY: SPOlUS "2a es Bee cht ie eS ee ee i, 6. Ocelli present, well-defined (Fig. 1). Hapto- meral region with 14—16 heli in a transverse row (Fig. 2). Each palidium consisting of a row of 17-20 pali (Fig. 12) cae toes, ome ee Amithao haematopus (Burmeister) — Ocelli absent. Haptomeral region with 10—15 heli in a transverse row. Each palidium con- sisting of a row of 23-26 pali Argyripa lansbergei (Sallé) 7. All tarsungulus bearing 5—7 setae ........ 8 — All tarsungulus bearing 8-15 setae ...... 10 8. With haptomeral cone-like process. Hapto- merum with a row of 10-16 short heli, and 7—14 short, stout, spine-like setae (Fig. 16) y) — Without haptomeral cone-like process. Hap- tomerum with a row of 16 medium size heli, and 16-17 medium size, stout, spine-like se- LAC Ae een a ere ae? Blaesia atra Burmeister 9. Last antennal segment with 5 dorsal sensory spots. Maxillary stridulatory area with row of 9 acute teeth. Each palidium consisting of an imecular row of 12—13 palit... .. 2.5. 5, (ones eis Marmarina tigrina (Gory and Percheron) — Last antennal segment with 2—3 dorsal sen- 1038 sory spots (Fig. 23). Maxillary stridulatory area with row of 6 acute teeth (Fig. 20). Each palidium consisting of an irregular row of 19— MO joyalli. (Ei, AS) 5 Ges Seats in". 3/4. Gia's & dod’ o WEE oe nee Ae Marmarina maculosa (Olivier) 10. Distance between the 2 lobes of respiratory plate of spiracles much less than the dorso- ventral diameter of the bulla (Fig. 48). Last antennal segment with 2—3 dorsal sensory spots — Distance between the 2 lobes of respiratory plate of spiracles slightly less than the dor- soventral diameter of the bulla, or as long as such diameter (Fig. 35). Last antennal seg- ment with 3—4 dorsal sensory spots 11. Metatarsungulus bearing 10—12 setae. Each palidium consisting of a row of 12-18 pali. Ocelli absent or vaguely defined. Spiracles with peritrema strongly sclerotized (Fig. 58) caer eh teat Gymnetis flavomarginata sallei Schaum — Metatarsungulus bearing 8 setae. Each palidium consisting of a row of 20-21 pali. Ocelli absent. Spiracles with peritrema weakly sclerotized Fvsneeh oy: Gymnetis hebraica difficilis Burmeister 12. Right mandible with 2 scissorial teeth. Hap- tomeral region with a transverse row of 8—11 heli ... Hoplopyga singularis (Gory and Percheron) — Right mandible with 3 scissorial teeth (Fig. 30). Haptomeral region with a transverse row of 12-19 heli 13. Metatarsungulus bearing 11—12 setae. Last antennal segment with 3 dorsal sensory spots. Each palidium consisting of a row of 14—18 pali .. Hoplopyga brasiliensis (Gory and Percheron) — Metatarsungulus bearing 9-10 setae. Last an- tennal segment with 4 dorsal sensory spots. Each palidium consisting of a row of 14—15 pali Hoplopyga liturata (Olivier) LARVAE OF AMITHAO THOMSON This larval description of Amithao hae- matopus (Burmeister) is the first for the ge- nus. Based on current knowledge of Gym- netini larvae, the larvae of Amithao are most similar morphologically to those of Argyripa species. The known larvae of Amithao have the ocelli well-defined, hap- tomeral region with 14—16 heli in a trans- verse row, last antennal segment with 12 dorsal sensory spots, tarsungulus bearing 7-8 setae, and each palidium consisting of a row of 17-20 pali. Larvae of Argyripa have the ocelli absent, haptomeral region PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with 10-15 heli in a transverse row, last antennal segment with 10-15 sensory spots, tarsungulus bearing 8—9 setae, and each palidium consisting of a row of 23-26 pali. Larvae of both genera have the dor- sum of abdominal segment VII with 3 an- nulets. The genus Amithao consist of 15 species that are found from the southeastern Mex- ico to Brazil, including Jamaica and His- paniola. The genus is being revised by Brett C. Ratcliffe. Adults of Amithao species have been collected with rotting fruit traps and are rarely attracted by lights (Moron et al. 1997). The known larvae feed in organic matter deposited in the axillary folds of leaves of epiphytes. Amithao haematopus (Burmeister) (Figs. 1-14) Third instar larva.—This description is based on four third instar larvae associated with dead adult females or their remnants collected from debris found in the axillary folds of leaves of epiphyte Acmaea sp. (Bromeliaceae). Locality data: México: State of Chiapas, Ocosingo municipality, Biosphere Reserve ““Montes Azules,’’ Boca del Chajul, 2-I-1983, 110 m elevation, C. Fragoso (1 larva) (IEXA); same data except 24-X-1984 (3 larvae) TEXA). Head (Fig. 1): Maximum width of head capsule 4.6—4.8 mm. Cranium smooth, or- ange yellowish. Frons with a median, lon- gitudinal depression extending anteriorly from the epicraneal stem, a single posterior frontal seta, single anterior angle seta on each side, and 4 anterior frontal setigerous punctures. Dorsoepicranium with 4 small setae and | long seta in a line diverging from center-base of head. Tentorial pits not defined. Clypeus: Shape subtrapezoidal with 2 posterior clypeal setae and 2 exterior clypeal setae on each side. Preclypeus weakly sclerotized, without setae. Labrum trilobed, clithra present. Epipharynx (Fig. 2): Corypha with 5 stout setae. Haptomeral region with cone-like process, without mac- roscopic sensilla, behind process a curved VOLUME 104, NUMBER 4 1039 Figs. 1-6. Amithao haematopus, third-instar larva. 1, Head, frontal view. 2, Epipharynx. 3, Right mandible, ventral view. 4, Right mandible, dorsal view. 5, Left mandible, dorsal view. 6, Left mandible, ventral view. Scale lines = 1 mm, except fig. 2 = 0.5 mm. row of 14—16 small heli, 6—7 stout spine- ma. Nesia with sensorial cone. Haptolachus like setae behind row. Acanthoparia with 7— with 2 sensilla below sensorial cone. Man- 9 short setae. Chaetoparia with 28—36 setae dibles: Right mandible (Figs. 3—4) with | on each side. Dexiotorma wide and long, — scissorial tooth anterior to scissorial notch with moderately developed pternotorma. and 2 scissorial teeth posterior to notch. Laeotorma elongated, with small pternotor- Stridulatory area elongate-oval, length 2 1040 Figs. 7-13. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ro 12 iS Amithao haematopus, third instar larva. 7, Hypopharynx and right maxilla, dorsal view. 8, Stridulatory area of maxilla. 9, Last antennal segment, ventral view. 10, Last antennal segment, dorsal view. 11, Abdominal spiracle. 12, Palidia. 13, Tarsungulus of posterior leg. Scale lines = 0.5 mm. times its width. Molar area with 3 lobes. Calx wide. Brustia formed by 6-8 setae. Lateral edge with 3—5 setae. Left mandible (Figs. 5—6) with | scissorial tooth anterior to scissorial notch, 1 tooth posterior to notch, and 2 teeth on premolar area. Strid- ulatory area elongate, length 3 times its width. Molar area with 2 lobes. Acia ab- sent. Brustia formed by 6—8 setae. Lateral edge with 3—4 setae. Maxilla: Galea and lacinia fused (Fig. 7), forming mala. Mala with large uncus at apex and 1| subterminal uncus vaguely bifid. Surface with 4 indis- tinct rows of setae. Stridulatory area (Fig. 8) with row of 5 curved, acute teeth and a distal, truncate process. Labium (Fig. 7): Dorsal surface with large, erect, truncate process. Hypopharyngeal sclerome with group of 7 setae on left side; both lateral lobes with 3—5 setae arranged in | row. Glossa with 8 setae at middle, and 2 lateral rows formed by 3—4 setae on each side. An- tenna: First segment as long as following 2 segments together. Surface (Figs. 9-10) of last segment with 12 dorsal and 8 ventral sensory spots. Ocelli clearly defined (Fig. 1). Thorax: Thoracic spiracles with C- shaped respiratory plate 0.58 mm high and 0.51 mm wide; plate with 23 holes across diameter at middle; holes irregularly oval. Dorsal surface of each segment with many short setae. Abdomen (Fig. 14): Spiracles of abdominal segments I—VII similar in size, those abdominal segment VIII slightly smaller. Distance between 2 lobes of respi- ratory plate slightly less than dorsoventral diameter of bulla (Fig. 11). Bulla irregular- ly oval, slightly convex. Dorsal areas of VOLUME 104, NUMBER 4 1041 Fig. 14. Amithao haematopus, third instar larva. Scale line = 5 mm. each segment with many short setae. Pres- cutum of abdominal segments IV, VI and VII with irregular, transverse rows of long setae. Scutum of abdominal segments II— VII with transverse row of long setae. Scu- tellum of abdominal segments II-VI with irregular, transverse rows of long. setae. Segments IX and X fused, covered with short setae and some sparse, long setae to- ward posterior borders. Spiracular area and pleural lobes of abdominal segments I—VIII with few, sparse, short setae. Raster with pair of palidia (Fig. 12) each consisting of an irregular row of 17—20 pali, rows joined anteriorly. Septula diffuse. Tegilla com- posed of scarce short, thick setae and some slender long setae. Lower anal lip with mix- ture of medium size setae and short setae. Legs: Tarsungulus (Fig. 13) cylindrical, apex rounded and bearing 7-8 setae. LARVAE OF MARMARINA KIRBY The larval description of Marmarina ma- culosa (Olivier) from Mexico is the second for the genus. Marmarina tigrina (Gory and Percheron), from Uruguay, was described by Monné (1969). Based on our current knowledge of Gymnetini larvae, the larvae of Marmarina are most similar morpholog- ically to those of Blaesia species. Larvae of Marmarina have a haptomeral cone-like process; with a row of 10—16 short heli, and 7—14 short, stout, spine-like setae behind row, last antennal segment with 2—5 dorsal sensory spots, maxillary stridulatory area with row of 6—9 acute teeth, and each pal- idium consisting of an irregular row of 12— 20 pali. Larvae of Blaesia have a slightly convex haptomeral process, with a row of 16 medium size heli, and 16-17 medium 1042 size, stout, spine-like setae behind row, maxillary stridulatory area with row of 7 acute teeth, last antennal segment with 4 dorsal sensory spots, and each palidium consisting of an irregular row of 14-16 pali. Larvae of both genera have the dor- sum of abdominal segment VII with 2 an- nulets. The genus Marmarina (= Maculinetis Schurhoff) consists of three species, three subspecies and three varieties that are found from the southeastern Mexico to Argentina (Blackwelder 1944). The genus is being re- vised by Brett C. Ratcliffe. Adults of Mar- marina species have been collected on ma- ture tropical fruits, rarely with rotting fruit traps, and sifting soil litter (Moron et al. 1997). The known larvae feed in organic matter deposited under rotten logs or in de- bris of ant nests of Acromyrmex sp. Marmarina maculosa (Olivier) (Figs. 15-27) Third instar larva.—This description is based on one exuvium of a third instar larva reared to an adult female, collected under a rotten log from tropical rain forest. Locality data: México: State of Veracruz, Monte Pio municipality, Estacion de Biologia Tropical ‘Los Tuxtlas,” 6-HI-1986, 150 m eleva- tion, M. L. Castillo (1 exuvium) (IEXA). Head (Fig. 15): Maximum width of head capsule 3.8 mm. Cranium nearly smooth, with sparse shallow punctures, reddish brown. Frons with median, longitudinal de- pression extending anteriorly from epicra- neal stem, a single posterior frontal seta and single anterior angle seta on each side, and 4 anterior frontal setigerous punctures. Dor- soepicranium with 4 small setae widely separated. Tentorial pits not defined. Clyp- eus: Shape subtrapezoidal, with 2 posterior clypeal setigerous punctures and 2 exterior clypeal setigerous punctures on each side. Preclypeus weakly sclerotized, without se- tae. Labrum trilobed, clithra present. Epi- pharynx (Fig. 16): Corypha with 6 stout, setae. Haptomeral region with cone-like process, without macroscopic sensillae, be- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON hind process a transverse row of 10 small heli, 7 stout spine-like setae behind row. Acanthoparia with 4—5 medium size or short setae. Chaetoparia with 36—47 setae on each side. Dexiotorma wide and long, with moderately developed pternotorma. Laeotorma short, with small pternotorma. Nesia with sensorial cone. Haptolachus without macroscopic sensilla below senso- rial cone. Mandibles: Right mandible (Fig. 17) with 1 scissorial tooth anterior to scis- sorial notch and 2 scissorial teeth posterior to notch. Stridulatory area elongate, sinu- ose, length 4 times its width. Molar area with 3 lobes. Calx short. Brustia absent. Lateral edge without setae. Left mandible (Fig. 18) with 1 scissorial tooth anterior to scissorial notch, | tooth posterior to notch, and | tooth on premolar area. Stridulatory area elongate, length 4 times its width. Mo- lar area with 2 lobes. Acia absent. Brustia absent. Lateral edge without setae. Maxilla: Galea and lacinia fused (Fig. 19), forming mala. Mala with large uncus at apex and | subterminal uncus vaguely bifid. Surface with 4—5 indistinct rows of setae. Stridula- tory area (Fig. 20) with row of 6 curved, acute teeth and a distal, truncate process. Labium (Fig. 21): Dorsal surface with large, curved, truncate process. Hypopha- ryngeal sclerome without setae on left side; both lateral lobes with 4—8 setae. Glossa with 2 setigerous punctures at middle, 2 transverse rows of setigerous punctures near basal margin, and 2 lateral irregular rows formed by 4—7 setiferous punctures on each side. Antenna: First segment slightly longer than the following 2 segments to- gether. Surface (Figs. 22—23) of last seg- ment with 2—3 dorsal and 3—4 ventral sen- sory spots. Ocelli not defined (Fig. 15). Thorax: Thoracic spiracles with C-shaped respiratory plate 0.54 mm high and 0.50 mm wide; plate with 20 holes across di- ameter at middle; holes irregularly oval. Dorsal surface of each segment with many short setae and some slender, long setae. Abdomen: Spiracles of abdominal seg- ments I-VIII similar in size. Distance be- VOLUME 104, NUMBER 4 1043 Figs. 15-24. Marmarina maculosa, third instar larva. 15, Head, frontal view. 16, Epipharynx. 17, Right mandible, ventral view. 18, Left mandible, ventral view. 19, Right maxilla, dorsal view. 20, Stridulatory area of maxilla. 21, Hypopharynx, dorsal view. 22, Last antennal segment, ventral view. 23, Last antennal segment, dorsal view. 24, Abdominal spiracle. Scale lines = 0.5 mm, except figs. 15, 17-18. tween 2 lobes of respiratory plate much less — short setae. Prescutum of abdominal seg- than dorsoventral diameter of bulla (Fig. ments I-VI with irregular, transverse rows 24). Bulla regularly oval, slightly convex. of long setae. Scutum of abdominal seg- Dorsal areas of each segment with many’ ments II—VII with transverse row of long 1044 ie ji i i : Figs. 25-27. 27, Abdomen, dorsolateral view. Scale lines = 0.5 mm, except fig. setae. Scutellum of abdominal segments I— VI with irregular, transverse rows of long setae. Segments IX and X fused, covered with short setae and some sparse, long setae toward posterior borders. Spiracular area and pleural lobes of abdominal segments I— VIII with many short setae. Raster with pair of palidia (Fig. 25), each consisting of a row of 19—20 pali, rows joined anteriorly. Septula elongate. Tegilla composed of many short, thick setae and some slender, long setae. Lower anal lip with many short PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON PS —mer y Ona nES Uoontn Anes AbN coe be aaranacoae 964 PATE AINIAIS Sa Mies Mie Sess CHRO DE RA IRA IER WS sactenertst tease aeeecistenie secre cetee a meee ncteae ae 554 BALXERASs JOAOUIN—See BROWIN-AIOEINGW) Weccrceecteeere eter ras-e seater ety eeeeen: 318 1090 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON BAL OGHS GEORGE see WILT ERDINGSJOFINGE SII esese sree eee eee ne ereeer eee eeeeeee 485 BEAULIEU, FREDERIC and TERRY A. WHEELER—Insects (Diptera, Coleoptera, Lepidop- tera) reared from wetland monocots (Cyperaceae, Poaceae, Typhaceae) in southern Quebec RE eS CEO ORO CC On ance nn ceeean oan Sooo orenannr cea rncanohodsetoe tana rncneccrsens 300 BLANK, STEPHAN M.—Taxonomic notes on Strongylogasterini (Hymenoptera: Tenthredini- (0 2: Ro ae rie COS Con Rene RE ota BAe eA SS - Qa B Era aC Mone AEE Seno HnY CAR SGT Baa nite ricn recat tiie 692 BOHN: 1.4 3:—See HARRISON} BE As. 0s Jeo ecaat seek sec esee clnete ies eee s atiaiat cee eaeres 655 BRAILOVSKY, HARRY—A new species of Maevius Stal from Australia and some notes on thevfanuly Eyocephalidae (Hemipteraleteroptera) nace eeeeeree ea eae eae eee 4] BRAILOVSKY, HARRY—A new genus and a new species of Daladerini (Hemiptera: Heter- optera:sCoreidae)* from#Madapascartseactsccscrecctseees eres, «ist ceislers ora renee Pare ae sate ioe ol otalevst a aielercisleletee itil BROWN, JOHN W., JORGE PENA, THERESA VASQUEZ, and JOAQUIN BAIXERAS— Description of a new tortricid pest (Lepidoptera: Tortricidae: Olethreutinae) of litchi (Litchi chinensis) in Florida, with a review of tortricid pests of litchi worldwide ...................... 318 BROWNS TOHNGW:=— Sees AD AMS Ku AWAD Wye Boer sagas Fe tose des ate cleric eiecre eee ten aerate 1029 BUFFINGTON, MATTHEW L.—Description of Aegeseucoela Buffington, new name, with notes on the status of Gronotoma Forster (Hymenoptera: Figitidae: Eucoilinae) .............. 589 BURGER, JOHN E—Description of five new species of Tabanidae (Diptera) from Costa Rica and revised keys to species for the genera Fidena Walker, Scione Walker, and Chrysops I Kesbeg Sat shown loys PAL SCE rere aa tnatc dccad don (aaa Met aenen ne Een otbos Aaep anon ananeSeaAnnoan SAR AAnnBonnee 928 BUYS) BRUNO: DD: See PRADO se ANWIEO) INA C@IOs. boo Spe tisccnaes yi ccseste anteascsinice tassel 1007 CANO, ENIO and MIGUEL ANGEL MORON—Additions to Phyllophaga suggenus Chlae- nobia (Coleoptera: Melolonthidae: Melolonthinae) from Guatemala ....................22...... 352 GARREE- JAMES Ee See BISINBREMFOMAS Aarese eee eee tees ascent eee cece eae ree 437 CARROLL, J. E—Notes on the responses of host-seeking nymphs and adults of the ticks Lxodes scapularis and Amblyomma americanum (Acari: Ixodidae) to canine, avian, and deer-pro- GUGCEAUSM STAM CSS aici seicccls,ciiciare crceaperetete state ete telet ces ai cveie clause aie aee aig evaie is eTorayte eveimyelekatete a auevnne inierane asta ere ete 73 CARVALHO, CLAUDIO JOSE BARROS de and MARCIA SOUTO COURI—Cladistic and biogeographic analyses of Apsil Malloch and Reynoldsia Malloch (Dipterea: Muscidae) of SoutherneS Outh*AmEentCal sce, «ae cere Poe are sia eo ne Sede reins se elaetee ea cle arsenite see Lanett ase 309 CHABOO, CAROLINE S.—Range extensions of New World tortoise beetles (Coleoptera: Ghrysomelidaem@assicimae) ia saxrsretetos ste atch aes eles: ase sicsstaleyniaicycraye clateioie seep asieve aiaihte oreyesetere aeolanicroteee ere ae ee 716 COPE SSE Sées HARRISON: BAG eae siss spete siete toretarenepstatom sites eyersiersrelere Mie ojetoreis aya wieteholeisiemjeielsio iss Oee 655 COSCARONEM -deliG=Sce DER DAP BMP IM c2-ta.g. gs eecee tic aasag char ene eee ote 168 COURI, MARCIA SOUTO—See CARVALHO, CLAUDIO JOSE BARROS de ............... 309 CRYAN, JASON R. and LEWIS L. DEITZ—Enigmatic treehopper genera (Hemiptera: Mem- bracidae): Deiroderes Ramos, Holdgatiella Evans, and Togotolania, new genus .............. 868 DAE GEEISHSROBERTs C= See RPRICEWROGERSD irreeneee cee eee eee oe eee eens 993 DeBENEDICRISSJOEIN See McEAB Es lINI© TEI Yes eeee sete sees eeeeeeereeerer cece 505 DEWMEZ. IE WIS: LE SeeiGRiVAN PIAS ON Re caiereteretevesc ste ere ele ovstesese siersieleleletonn ai sefare st ereietelecointe ierase|etelecorn= 868 DELLAPE, P. M., M. del C. COSCARON, and B. EF AMARAL FILHO—Immature stages of Montina confusa (Stal) (Heteroptera: Reduviidae: Harpactorinae) ...........................008- 168 DEONIERSD le SeCeu BIR ER As Be cer secec cins cate sinclar geome tae Gielas aise a RCE 458 DeWALT, R. EDWARD, DONALD W. WEBB, and AMY M. SOLI—The Neoperla clymene (Newman) complex (Plecoptera: Perlidae) in Illinois, new state records, distributions, and an IG eyaUATCEIKOVIN EN/« 5 Sqn odo pasavaqnaciesudnaan cdBoUosoGcads dd 0d005 0006 SeouERogg GB aodoaSHA pod b Hoses soNadsaS 126 DIKOW, TORSTEN and WAYNE N. MATHIS—A revision and phylogenetic study of Acto- celombecker: (DipterawE phy. dridae) messes hee eet oes laste eee teal orr eee cca ttre 249 EISNERGMARIAS See EISNER THOMAS Feseeererecese = oO-r een eee eer sate nee eee eee Eee 437 EISNER, THOMAS, JAMES E. CARREL, EILEEN VAN TASSELL, E. RICHARD HOE- BEKE, and MARIA EISNER—Construction of a defensive trash packet from sycamore leaf trichomes by ‘a shrysopid’ larva’ (Neuroptera: @hrysopidac) so 22 see. eee eee sisi steele eee 437 FERREIRA, PAULO SERGIO FIUZA and THOMAS J. HENRY—Descriptions of two new species of Fulvius Stal (Heteroptera: Miridae: Cylapinae) from Brazil, with biological and biogeorraphicmotesjontthessenusieeeses heer eeess heat een eee etree re ee rn ete eee eemee eerste 56 FIL HOSBaEVAMARAL “See, DEIIGA Plu Pi Mi &11)..245. cela nieeh Anaad 0 eee ee ee 168 ERING OGIVERSS JR See, HARRIS NSE MEN (Ci eio-eecceeeeet cree eeeeeeee eee eerie 195 FLORES, GUSTAVO E. and CHARLES A. TRIPLEHORN—Entomobalia, new genus, the first member of Nycteliini (Coleoptera: Tenebriondae) from Brazil ...........................5- 602 VOLUME 104, NUMBER 4 1091 . GOEDEN, RICHARD D.—Descriptions of Tephritis footei and T. headricki, new species (Dip- tera: Tephritidae), with notes on their life histories in southern California ...................0. 142 GOEDEN, RICHARD D.—Description of immature stages of Tephritis stigmatica (Coquillett) (Oipterarelepinritrdae): ketal sus Wee nl eva saan acs an Meanie cect neiaatle teeclnaecke ae ache aarti: 335 GOEDEN, RICHARD D.—Life history and description of immature stages of Oxyna aterrima (Doane) (Diptera: Tephritidae) on Artemisia tridentata Nuttall (Asteraceae) in southern Cal- BUTI Aaneeter eee orck ajchen Ae ast inch ois sue CREAM nian. Pence ahee Sassy SoaeER sen wea SEE AE Paanne 510 GOEDEN, RICHARD D.—Life history and description of immature stages of Oxyna palpalis (Coquillet) (Diptera: Tephritidae) on Artemisia tridentata Nuttall (Asteraceae) in southern CG AE RIN Ale 115 0 id Rigrena ck 2 SE OAS See a eee AERA OG E abl 5 an eA as EO SDSL ES ea 937 GOEDEN, RICHARD D.—Life history and description of immature stages of Goedenia rufipes (Curran) (Diptera: Tephritidae) on /socoma acradenia (E. Greene) in southern California ... 576 GOEDEN, RICHARD D.—Life history and description of immature states of Goedenia setosa (Foote) (Diptera: Tephritidae) on Ericameria brachylepis (A. Gray) H. M. Hall in southern co AMS sits statis wating erat aoe eee See ae eer CaaS on eee Mane eek LP Mis knee eee mr ey, 629 GOEDEN, RICHARD, D.—Life history and description of adults and immature stages of Goe- denia stenoparia (Steyskal) (Diptera: Tephritidae) on Gutierrezia californica (de Candolle) Torrey and A. Gray and Solidago californica Nuttall (Asteraceae) in southern California .... 702 GOEDEN, RICHARD D.—Life history and description of adults and immature stages of Goe- denia steyskali, n. sp. (Diptera: Tephritidae) on Grindelia hirsutula Hooker and Arnott var. halli (Steyermark) M. A. Lane (Asteraceae) in southern California ...................0000eeeee: 785 GRIER GS CSAP TRIB "eri Freese ets coe RSE ee ON ies ee ee Seer TT hc eT ae eee 646 HALL, JASON P. W.—A review of the new riodinid butterfly genus Panaropsis (Lepidoptera: Riodinidac:ys yiMIMAachiimi)) «ees sence earome ee te eee ree Ieee ROMEO eee ener re 63 HALL, JASON P. W.—A review of Chalodeta Stichel with a revision of the chelonis group (epidopleraRIOCIN dae) Fate cewss ntaa csi ieiocee ae seine Gita valeias stecten seen a etaaaeainereee seine ster 376 HALL, JASON P. W. and DONALD J. HARVEY—A revision of the Neotropical butterfly seenusiseco Halland Harvey (Lepidoptera: Riodinidae) ii -..--seesaaceea sts cecacee - seeemiae cere 94] HARBACH, RALPH E. and THERESA M. HOWARD—Sabethes (Peytonulus) paradoxus, a newsspeciessot Sabethini (Diptera: Culicidae)! from Panama 22.222. sc.e-cess. sos esecee seme eciene 363 HARRIS Sie VEN ‘©. Sees HOLZENTHAT RAL BED Wis teccrs cries cease co rsiones eceesmetnesit: spice 106 HIARRISS Ss 3Gi——See-K BIPERs JosBe 2 is ect cnet a aethrem tecising soe oe teeta. ater ache slo cietaneenesyasieiane 291 HARRIS, STEVEN C. and OLIVER S. FLINT, JR.—New Alisotrichia (Trichoptera: Hydrop- tilidae) from Central and South Ameerica and the Greater Antilles .....................-.00-+.- 195 HARRISON, B. A., P. B. WHITT, S. E. COPE, G. R. PAYNE, S. E. RANKIN, L. J. BROWN, E M. STELL, and C. J. NEELY—Mosquitoes (Diptera: Culicidae) collected near the Great Dismal Swamp: New state records, notes on certain species, and a revised checklist for INV 0.55 OUTTA pera 28a eters ofehosevasheieye dun rage Sito sare cise bepctcbansrevosa a oeke cyaleroteneictoge Sale ope WiorosS adel els ofersies fed ciccleelescierare aesiarere oes 655 HARVEY «DONALD J.—See- HALL, JAS ON BW: cassie ce ciessccisice otto 3 etote a oteicreini otro oie ctermtatsje esis aces 94) HASTRITER, MICHAEL W. and MICHAEL E WHITING—Macropsylla novaehollandiae (Si- phonaptera: Hystrichopsyllidae), a new species of flea from Tasmania ....................0005: 663 HIAVIBIEKAC IAN —S ce LEB, SBUNGEIWAIN aioe ore chistes cree ignee este eniccinee eye eiisseutisser 447 HENRY, THOMAS J.—See FERREIRA, PAULO SERGIO FIUZA ......2..2....00001seeeececeee 56 HENRY, THOMAS J.—New species of the plant bug genera Keltonia Knight and Pseudato- moscelsmreuter (Heteropteras Miarid aes Pinyility ac) iar tartrate sere toreretare stool sie ipa staxctarer crashes 97 HENRY, THOMAS J. and RANDALL T. SCHUH—Two new genera to accommodate two North American plant bugs (Heteroptera: Miridae: Phylinae)) ...:.....0...jc000cecsssess2008ee2e0: 211 HESPENHEIDE, HENRY A. and LOUIS M. LAPIERRE.—A review of Prous Champion (Co- leopteras Curculionidae: Cry ptorknyme li ae) ye ree eae ge sclelatarecctate ole 2 etelerets aca lates stele tareretniere etNctere eletere 856 HORBEKEE RICHARD—_See EISNER, DHOMAS eters: tens sete ene blac te see eee steer 437 HOEMAN: JAROSLAV—See LEE, SEUNGEIWAN) feice cece nineties stelas ste oe ae loinettelsteeryacietelaiste/s 447 HOLZENTHAL, RALPH W. and STEVEN C. HARRIS—New species of Nothotrichia Flint @irichoptera: My droptilidac) from Brazil andiCosta Rica) 22a. 2 n-gassec< te owls te cmanesee 24 ante 106 HOWARD) THERES AUMi==See HAR BA CE RIATE PEA vores. oid slarsaeleiare inate sieteleleie siaictaieieraislale 9 aclatAor 363 HUSBAND, ROBERT W.—A new species of Eutarsopolipus Berlese (Acari: Podapolipidae) from the Galapagos Islands, a parasite of Agonum chathami Van Dyke (Coleoptera: Carabi- (GE Te)) gs Se Pe As PP ed Se Bae oe A A ae eA errs REC RCC On crt 563 TTANNIN© Mii — See KEIPER: Ja Ban eeaiece seen ctras tonsa cmt. ote inemrcters « mae te eie hice t aise weietlz Maereiere 458 1092 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON JOHNSON, NORMAN E and LUCIANA MUSETTI—Rediscovery of the genus Platyscelidris Szabo (Hymenoptera: Scelionidae) and description of a mew species ..................00...0 00: 948 KEIPER, J. B. and S. C. HARRIS—Biology and immature stages of Ochrotrichia footei (Tri- choptera: Hydroptilidae), a new microcaddisfly from a torrential mountain stream ........... 291 KEIPER, J. B., D. L. DEONIER, J. JIANNINO, M. SANFORD, and W. E. WALTON—Biol- ogy, immature stages, and redescriptions of Hydrellia personata Deonier (Diptera: Ephydri- dae) fa ESN GDAIMUIN Cte cecaee cts solopeis Sewers is wine able sicie oilieishele uae ge coeis vo ereke See meee eel ee Rome 458 KIMANI-NJOGU, SUSAN W. and ROBERT A. WHARTON—Two new species of Opiinae (Hymenoptera: Braconidae) attacking fruit-infesting Tephritidae (Diptera) in western Kenya BF seaySpst Aeectoloay re tic ove orzc aU ios Rios ple Sia ee alors oR STAS tare Sb ns muoceis ore oraiv ees aseiateccke be Giclee Lace Ge ec ansiene rsterete ERO 79 JUUSXCTSINI SIRS IRS IR——Sewe DMO INIDIRVANIMUSIELS 18k (Co Se pcoccssonscauceonoeeenuooansaduaascsnasneaags60c08 51 KONDRATIEFE B. C. and R. EK KIRCHNER—Perlesta etnieri (Plecoptera: Perlidae), a new UAHA Oi Co mlishy TKO MSNNEISAS sna saoncnencdoceadcndgnsac ccd gaae se scouansHaTdgne a00dsonea000nIaE 51 EAPIERRE SE OUISSME— Scere SPEINDIE IDE SEE INR YGACr cae eeeee eet cece Bent Stree 856 LEE, SEUNGHWAN, JAROSLAV HOLMAN, and JAN HAVELKA—Taxonomic revision of the genus Megoura Buckton (Hemiptera: Aphididae) from the Korean peninsula with the descniptionyoteammewaspecies andvaskeyatonthe pworlduspecicspeeaeeee eee sere eee eee eee eee ee 447 EEWINSOHIN, THOMAS Mi—Sce:PRADO) PAULO INAGIO) 0... .4.,5-0--4.9-02008) ee: 1007 LOPES MARCELO) B= See /PRADOME AU ONIN AGIO us soca eee ae eee ee 1007 LOPEZ, ROLANDO — See IVINS Rt GRINS, «2 ck acres oe ae eRe aA Ree en eee 160 MACEDOSANTONIOICAREOS— Sea PeADO. PAULO MNACIO) 3.1 ea ee ee 1007 MANE EY] DONALD iG Sce- PIR SSU AN EMSaP So ssesepenemecce ect anen tee nae CEG eee ener 672 MARILUIS, JUAN C.—A new species and new records of Microcerella Macquart (Diptera: Sarcophagidae) iompAreentineanibatasonilageasssseeasa- > -eeeeeeee tees eee cee eeeeeeaeeeeeeeree 9] MATHESS WAWINE SN: Sees DIK OWAMOR STEIN ee ccnssossceeeetie accereene cee eee ase een 249 MAWDSLEY, JONATHAN R.—Comparative ecology of the genus Lecontella Wolcott and Chapin (Coleoptera: Cleridae: Tillinae), with notes on chemically defended species of the beetlestarmnailiy Seri el ae wy op-fa ia sts ote os SPs este te ako oes a sais orererke Sets A a eee « ae PS 164 McCABE, TIMOTHY L., WILLIAM D. PATTERSON, and JOHN DeBENETICTIS—Pseu- dobryomima fallax (Hampson) and P. muscosa (Hampson) (Lepidoptera: Noctuidae) leaf- IMMUN SgnOCtuid Sr OMEN Sie 1 chrsee ceo ae cat eee eee cles. ciate oie ae aaa eee Canoe P REE CEE co 505 MIPREDUR ANTE Mc P=—See NIETO NAPRIA- Wa ).2..- <6 sc ee ecco ones 918 MILLER, DOUGLASS R. and GARY L. MILLER—Redescription of Paracoccus marginatus Williams and Granara de Willink (Hemiptera: Coccoidea: Pseudocoddidae), including de- SGHpPWOUS Olsthesmmatunre stacesrandsadultamalemesrereerenesece cee once eer ccaenetere acaeeeaceree | MILLER, DOUGLASS R., GARY L. MILLER, and GILLIAN W. WATSON. —Invasive spe- cies of mealybugs (Hemiptera: Pseudococcidae) and their threat to U.S. Agriculture ......... 825 MIEEERY DOUGEASS R'—See WILEVAMS DOUGLAS E iran. setenceeee ects eee emer 896 MAO LETS SS IDXONOKGIIANSSSY IRE SY ete IMI EIR MGUANIKING IL Gocedosooccodsdusoonuocedono cans oa0o0bRb dona 986 MITETE BR GAYS Seer Mille ERD OUGIEAS SaRG ser ceerrie eo scias a-ciae see sees eeieeee erat 1 MILE ERIGARYS See MIEBER] ID OWGEAS SIRS erseereee racers a eerecee eee eaeeee ere ee 825 MILLER, GARY L. and DOUGLASS R. MILLER—Dysmicoccus Ferris and similar genera (Hemiptera: Coccoidea: Pseudococcidae) of the Gulf State Region including a description of AME wWASpEeclesuandene we UMiItedes tates meCOLldSy: ets eee eeeeeeee eee eee eee eee ererre 968 MILLER, GARY L., MANYA B. STOETZEL, ROLANDO LOPEZ, and DANIEL A. POT- TER—Geoica setulosa (Passerini) (Hemiptera: Aphididae): New distribution records for NorthvAmentcaljyayasceiicdsketinne ss cae s tage bo ouis sb ge sinats Aae ene sere das since neh aiagieh meet ee 160 MORON gMIGUEL“ ANGEL = See CANOMENIO@ -- ie peeenerere) | 2.) GU eee at 352 MORON, MIGUEL ANGEL and ROBERTO ARCE—Descriptions of the immature stages of five Mexican species of Gymnetini (Coleoptera: Scarabaeidae: Cetoniinae) ................... 1036 MUSES WeliAINA— See JOHNSONSNORIMAINGE peste seeeeree ree eae ee eee eee eeeeetee 948 INBELY,,.C.7J-—-ScesHIARRIS ON): BiigAs saeiejsiois «nae ae eee mote oreeors: tw oot ee oem mentee eter tel 655 NEUNZIG, H. H. and M. A. SOLIS—The Ceracanthia compex (Lepidoptera: Pyralidae: Phy- citinae) in Costa Rica. I. Ceracanthia Ragonot ..... Be SU OPH O RED ROO RECO con Oat Sane ce MOC an cHoee 837 NEUNZIG, H. H. and M. A. SOLIS—The Ceracanthia complex (Lepidoptera: Pyralidae: Phy- citinae) in Costa Rica. Il. Megarthria Ragonot, Drescoma Dyar, and Lascelina Heinrich ... 980 NIETO NAFRIA, J. M., M. PR MIER DURANTE, and J. ORTEGO—Pentamyzus Hille Ris Lambers, a Neotropical genus of the tribe Macrosiphini (Hemiptera: Aphididae: Aphidinae), with the description of a new species ................ « LAS ete has erniays sieicid ple aah ee ate Se ae Seeeeos eee 918 VOLUME 104, NUMBER 4 1093 NORRBOM, ALLEN L.—A revision of the Anastrepha serpentina species group (Diptera: ANSTO OVAIUGE TS): contd e OGRE OREEET EAR ANGE Cn otra ME oee sine ae ni toca: Meni oo Bean kee ence ee aanccnmnte 390 NORRBOM, ALLEN L.—A new species and key for the genus Zonostemata Benjamin (Dip- (HERES) INSSOLOVOINICIEYS)) -caepneenoe a aren aise cece nccbciadetc cree rear mince atemsecanos srt oducsoeancocededr 614 NORRBOM*s ALPEN L.—See PRADO; PAULO INAGIO, c2iicc ccd: cssnctce tyctouceecseussveessean 1007 OCAMPO, FEDERICO C. and FERNANDO Z. VAZ-DE-MELLO—A new species of Daim- othoracodes Petrovitz (Coleoptera: Scarabaeoidea: Hybosoridae) and a key to species of the 1 STIS rt Pe SPS oie Loe NIELS iene soS nse, SS ke Diuccie sisienans Ghiais acste|ninis oietale cies ots ah otal eetareaw eeomanoe aetna i cia 912 ORDEGO SM =See NINO NAPREA OSM) oo. soc2iis vances cue cata eee tL oeaaet areata hates 918 PATERSON, WILEIAM D—See McCABE; TIMOTEY Teves i5 ccc cees creeds seiecetcpisise clecices « 505 RPAYINE | GaRe——See HARRIS ONMBe A ss. csicle so -civ due diice on netden ck ceiis se ccitues om cee ela eeieeietreis 655 PENN IORGE— Seer BROWING JOEIN Wo oc xcees nosoveead oncag acadtce ae sana a Askin ciloaeealees Atsed 318 PEREZ-GELABERT, DANIEL E.—Further characterization of Paleomastacris ambarinus Perez et al. (Orthoptera: Eumastacidae) from Dominican Amber, .... 4/26... ssn. -e -ceee «cee 330 PIKE, K. W., P. STARY, G. GRAE and D. ALLISON—Pauesia columbiana, n. sp. (Hyme- noptera: Braconidae: Aphidiinae) on juniper aphids, and a key to related species ............ 646 PITTS, JAMES P. and DONALD G. MANLEY—Description of females of Stethophotopsis Pitts and Sphaerothalma (Photopsioides Schuster) (Hymenoptera: Mutillidae) ................ oy2 POLHEMUS, DAN A.—A new species of micropterous damsel bug (Heteroptera: Nabidae) HOmMMNUKUMIVax Marquesas SlamaSe: vic os icocte ones cncue dialer caer Chat fen orbiqeen toner eeasaeaan ase a= 640, POLHEMUS, DAN A.—Two new species of orsilline Lygaeidae (Heteroptera) from the Ha- WATTAMOUSTANGS 9% crecinoere sleaeepice eee Setar tie ae ciclo w eieci ciao Web wonreg Deby Sadan Sapmeeerie meant ne dened sagas 955 POTTERSDANIEILA:—see MIBIEBRY GARY (Ly hi. 5s 0003 Ge asi50051s's Su steinee eee Aeee teasers 160 PRADO, PAULO INACIO, THOMAS M. LEWINSOHN, ADRIANA M. ALMEIDA, ALLEN L. NORRBOM, BRUNO D. BUYS, ANTONIO CARLOS MACEDO, and MARCELO B. LOPES—tThe fauna of Tephritidae (Diptera) from capitula of Asteraceae in Brazil .......... 1007 PRICE, ROGER D. and ROBERT C. DALGLEISH—The chewing louse genus Kaysius Price and Clayton (Phthiraptera: Amblycera: Menoponidae) from the Passeriformes (Aves) ....... 993 PRICE, ROGER D. and ROBERT M. TIMM—Two new species of the chewing louse genus Gliricola Mjéberg (Phthiraptera: Gyropidae) from Peruvian rodents .....................eeee eee 863 RAINING S3E— SeesHARRIS ONE BRA. .cdcadansctcin vue nes oeeleetygebiisaeee dostidusemn dass sMeestecese 655 SAINI, MALKIAT S., DAVID R. SMITH, and V. VASU—The sawfly genus Kambaitina Mal- aiSes (Ehymenoptera ade mtiime dimmicl ae) ata yD ht choo feat ene ratte cose = oinjatato ohater state ots eye ainjelevoleveiele Mieetare nie alate ra ep stelers 884 SATNISNMIAT KATES SS CORVAS WG Noirs nis site de siectk «aed Sess ieateie atest slacss sbideem ase ee sen re eee seers 997 SANE ORID SIM. S ce dIRE IRE RE MB satiate tez bncite soins cielat nie sie io eels ee meen ntl aeee gerne Sie me = 458 SCARBROUGH, A. G.—Redescription of two species of Ommatius Wiedemann, with lectotype and paralectotype designations for Ommatius tenellus van der Wulp and range extension and a replacement name for Ommatius tibialis Ricardo (Diptera: Asilidae) ...............+.++2.005- 680 SCHEFFER, SONJA J.—New host record, new range information, and a new pattern of vol- tinism: Possible host races within the holly leafminer Phytomyza glabricola Kulp (Diptera: INET RONTINVAKEHTS) coe, Sorin Eda0 6 be qEae eee OED ane nacebaaccsnecandsecerdp anaes cue chudsancdoD adh posto gor 571 SCHIBEGNATETAINIM:—SeerS MiIMiEs DAVID) Re vice eset ssstales strlaid\-reisiaieloie ee = otis Se eeceiaaer'- 174 SCHRODER, R. E W. and M. M. ATHANAS—Biological observations of Centistes gasseni Shaw (Hymenoptera: Braconidae), a parasitoid of Diabrotica spp. (Coleoptera: Chrysomeli- GEO) Sracenbseen noun onic bt eo CRASS Dodie Soe aa NnE te aon oronc aGe itcnn Aba bndnanaods spacorcosnnaasarracdscosKd 554 SGHUOEBIRAND Albleul——Seev HENRY. THOMAS I tar cece ssc ects rite eicteieets teijaet slackers 2g SHINO HAR AS NKIETIK@==See SMI. IAW TID Ree aicrresterrfotsie srelsrale oe ois.cle cielelew miele sleek rare 479 SHINO HAR AR AKGEIK@=—SeerS METH sD A VAD BRo wero erstasetieacloeie eas nese oaiaere eines eee 624 SLATER, JAMES A.—The status of some plesiomorphic blissid bugs (Heteroptera: Lygaeo- idea sRlissidae) wiromiethes Orem tess. cere rca cre ae eo creas ie cc cieecorete stele orate to cyepeister-folevsloke Gisrarctavestats eae ater 348 SMITH, DAVID R. and NATHAN M. SCHIFF—A review of the siricid woodwasps and their ibaliid parasitoids (Hymenoptera: Siricidae, Ibaliidae) in the eastern United States, with em- phasis; on the mid-Atlantic region: 32... 5.2.2) 2 iets cece doce e ne ae cae nde vente scisie'ss vies spams sje sieainsas 174 SMITH, DAVID R. and AKIHIKO SHINOHARA—The stem-boring sawfly genus Cephus Latreille (Hymenoptera: Cephidae) im Japan ............002-secee eee e reset eee cce tee neteree senses cee’ 479 SMITH, DAVID R. and AKIHIKO SHINOHARA—A new genus and new species of Cephidae (Hymenoptera) from Sulawesi Utara, INGONESi as been cee re eee ee ee a Eee eee 624 SMUT DAVID YR: —See SATNEINEATE KATES 2) cerestccy aisle septa nctesiats alsie Selaclotwie elerelehelefelote elernicteleletatats 884 1094 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SMEPEE DAVID Ri= 2S Ce VAS WRI sas orcc cle ssa ane se eats STE Ie Oe ea ee nee 997 SOLI, AMY M.—See DeWALT, R. EDWARD ....... Siete eek rtere SO eee le cee Ci ort eee 126 SOLIS SIME AS eeu NE UINZ LGU aa aac iars von erele otter toes oretereh cones Ce etaranae Sees eer eee Eee 837 SOETSs MSA =—=SeesNEUIN ZIG aE cate ose alarore te erases ere SO IOS URES EC cE Ene 980 STAINES, C. L.—Hamlet Clark: Publications and proposed taxa ...................00..c0ecceeeee 468 STAINES, C. L.—The New World tribes and genera of Hispines (Coleoptera: Chrysomelidae: (EZ S V0 Uiot 12) lec anticta peneae aaeaeh op tn don SOC eR EM EERE NC Guat orion onc ce OCU TSA Ss aae SEHR RIAA Se Nita R SANS, 721 SIPAR Ya'1P——S ees PIIKE. IRE S sic: aprrer rest ars desis sie aivrsrete cia cee aya rietoes Bae SOS noe EOE EEE REET 646 SMELL, JF -Mi—See HARRIS ON Ba Aten iit: a scnaeenannnicttdrmsatemen oe woes ae ae eee eee ees 655 SHOETZERS MANYAGB—Sec) MIEIBERAGARY SIE esnacescese teens tae eee ee eee 160 TIMM, ROBERT M.—See PRICE, ROGER D. ....... = fatute s Giskesmiarateece tea ars eteoer oh ee ein ase eietate ae 863 TOGASHI, ICHIJI—Description of a new species of the genus Masaakia Takeuchi (Hyme- noptera-wlenthrecdinidae) sirommMapanwerreree-eeeereeeaeeeeteeacee eee Ceree te area eee eee 373 DRIPEBHORN: CHAREES AC See PBORESGUSPAVIOlLEeeeeeseseecetsae tee ee oe ee eee 602 VANDENBERG, NATALIA J.—The New World genus Cycloneda Crotch (Coleoptera: Coc- cinellidae: Coccinellini): Historical review, new diagnosis, new generic and specific syno- nymsandrankimproved Keyato North yAmentcan SPECleSmeeesa-ae- eerie teeceee eee ee nee 221 VAN GIASSEEESEIEBEINS—Sce BISINERESIEOMAS Eessennenee cee eeeecee a eteetes tee Eee eeeee ee 437 VASOUBZ; THERES A——Sec BROWINSTOEINGW, «=o. -cenceeteee teen tee cece eee eee 318 VASU-V.—SeerS ATNITAMIATE RIAT™ S seer ar eter te cicisi isnt sscrtnaas samen eietee anes eee ees eee ta eaeee eee 884 VASU, V., MALKIAT S. SAINI, and DAVID R. SMITH—Revision of the genus /ndostegia Malaise (ymenopteraslenthnredinidac) aingindiav. cee eceeeeeetee eee nee econ eee 997 VAZ_DE-MBEEO FERNANDO Zz —Sce OCAMPO: FEDERICOlIG@) a s--- ee te eee ee 912 WATETON WE — Sco NiBIR EB Rese Be ernserceete no acl area geome scan ac ein ee eee an en eee 458 WATSON TGIEETANDW2—See MIEE BREED OW GIEAS SYRa ereeees-saeeee nee eee eerie 825 WEBB DONA DAW SceDeWAI kee DNWARRID oe. eeeeeeeeeeee eee nee eee eee eee ee eeeeeee 126 WHARTON ROBERT Ac—See KIMANIENTOGUS SUSAN W 22tecee-soesce cee sees seen eenee 79 WHEELER, A. G., JR.—Chilacis typhae (Perrin) and Holcocranum saturejae (Kolenati) (He- miptera: Lygaeoidea: Artheneidae): updated North American distributions of two Palearctic Cattaile bus Stents weetaskh eee es eso sos Lone ae eee once e a «ore cine eae aa en eee See ee meee eee 24 WHEELER, A.G. JR.—New distributional and host-plant records for the Heteropterans Aufeius impressicollis Stal (Rhopalidae) and Prionosoma podopioides Uhler (Pentatomidae) in the southeastern United States, with notes on their western United States hosts ................... 138 WHEELER, A. G., JR.—Leptoypha elliptica McAtee and L. ilicis Drake (Hemiptera: Tingidae): New distribution records of seldom-collected lace bugs, with clarification of host-plant re- LatlOMShips eb Se dios tater eens etter oie oie s 5 a lates ee Sie ares o BE win ocale oes RIS Meare ee ene 687 WHEBLER; TERRY A>—See BEAULIEU; FREDERI©@ (2... ..cc0-clesnese ees eee: 300 WEIERINGS MICHAEL F—See HASTRIMERSMIGEHABIGIW.) 22tencccesnesecte seeeeetesaeeeneceee 663 WELT PSB —— Secu HIARRIS ONRBAGAR Teh cne se cene ose eae erate aie le oie srcieta cee eles aetare eae eee 655 WHITWORTH, TERRY—Two new species of North American Protocalliphora Hough (Dip- teras Calliphondae) stron bindimMestsaeeeeeeee- Meee eee reece eter reece eect eee ence eee seater eere 801 WILLIAMS, DOUGLAS J. and DOUGLASS R. MILLER—Systematic studies on the Antonina crawi Cockerell (Hemiptera: Coccoidea: Pseudococcidae) complex of pest mealybugs ...... 896 WILTERDING, JOHN H. Il and GEORGE J. BALOGH—Review of the North Amereican gray Pyla Grote (Lepidoptera: Pyralidae: Phycitinae) with description of a new species from western Umiteds S tatess i sree cis arerarava re hlssrs Osp pr. OU Dyers eaten eeraes: oA mee Dee ote ed ee Pe Ee MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 No.5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 _. No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi Ber Keer ete Tees Opa ant) mmm eee ana Ae SMR SI TE rg I DE Se No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- pogonidas): by Wall. Grogan. Ir and Wi\W. Wirth, 125) pp: 1979) No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. COT GED] MA ere er ae nee ee Pa arse RI eT ed 2 ol oc So Oy en Gl No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 jeyeys, NAS Ss sees Lee NP eh iin NS Se lek a eee ee ei es CEPR shai” Ae eed es oe Om Oe No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent BAR CUNWE Ie, (Giroyererny Ur IGFs) Foy on MEAS i) ea ree SERN Se et See eh ew ne ee eee eee ue ee No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 __. No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited Dba NordenandiAcs Menke: 2G ppilOOG) suet se eee ee 25.00 No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and Willian Grogan rs? Simp en USS iin sete Bae dA A de te No. 19. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by SHMOMAS ELEN Ty AA OMT) eS /u na wale y eet ina LL ica ea ae Wa ED Ye ae en lee A LIN ws No. 20. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- PATS LECTED ION SHEN) et) to eaten oe aed NS 1s et OA AN 8s Oe eae EN Ted oe ae ie ate No. 21. New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. Leura al CO oj opel MOXS Rosh 5 ib Nhat apie oe SEGAL ND i he) Sine AE TN en eerie Re BES SED Ia No. 22. Systematics of the North American Species of 7richogramma Westwood (Hymenoptera: linichorrammatidac) spy Onn MeimtO: 28 / ppl O99) ss eee ee ee eee No. 23. Revision of the Net-Winged Midges of the Genus Blepharicera Macquart (Diptera: Blepha- riceridae) of Eastern North America, by Gregory W. Courtney. 99 pp. 2000 ____-----------------------__- $10.00 15.00 40.00 30.00 $15.00 15.00 15.00 15.00 12.00 5.00 25.00 25.00 25.00 25.00 18.00 12.00 12.00 28.00 14.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume to non-members and $25.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10 percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Treasurer, Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. CONTENTS (Continued from front cover) NIETO NAFRIA, J. M., M. P MIER DURANTE, and J. ORTEGO—Pentamyzus Hille Ris Lambers, a Neotropical genus of the tribe Macrosiphini (Hemiptera: Aphididae: Aphidinae), with the description of a new species 918 OCAMPO, FEDERICO C. and FERNANDO Z. VAZ-DE-MELLO—A new species of Daimothoracodes Petrovitz (Coleoptera: Scarabaeoidea: Hybosoridae) and a key to species of the genus POLHEMUS, DAN A.—Two new species of orsilline Lygaeidae (Heteroptera) from the Hawaiian Islands PRADO, PAULO INACIO, THOMAS M. LEWINSOHN, ADRIANA M. ALMEIDA, ALLEN L. NORRBOM, BRUNO D. BUYS, ANTONIO CARLOS MACEDO, and MARCELO B. LOPES—tThe fauna of Tephritidae (Diptera) from capitula of Asteraceae in Brazil PRICE, ROGER D. and ROBERT C. DALGLEISH—The chewing louse genus Kaysius Price and Clayton (Phthiraptera: Amblycera: Menoponidae) from the Passeriformes (Aves) 993 PRICE, ROGER D. and ROBERT M. TIMM—Two new species of the chewing louse genus Gliricola Mjoberg (Phthiraptera: Gyropidae) from Peruvian rodents 863 SAINI, MALKIAT S., DAVID R. SMITH, and V. VASU—The sawfly genus Kambaitina Malaise (Hymenoptera: Tenthredinidae) in India 884 VASU, V., MALKIAT S. SAINI, and DAVID R. SMITH—Revision of the genus /ndostegia Malaise (Hymenoptera: Tenthredinidae) in India 997 WILLIAMS, DOUGLAS J. and DOUGLASS R. MILLER—Systematic studies on the Antonina crawi Cockerell (Hemiptera: Coccoidea: Pseudococcidae) complex of pest mealybugs NOTES LATTIN, JOHN D. and JAMES R. LABONTE—Cardiastethus luridellus Fieber (Hemiptera: Heteroptera: Anthocoridae), a non-indigenous anthocorid discovered in Oregon OVRUSKI, SERGIO M.—New records of fruit fly parasitoids (Hymenoptera: Braconidae, Figitidae, Pteromalidae) for La Rioja Province, northwestern Argentina RABAGLIA, ROBERT J. and GAYE L. WILLIAMS—Two species of western North American Hylesinus Fabricius (Coleoptera: Scolytidae) new to the eastern United States ROBBINS, RICHARD G., STEVEN G. PLATT, and JAMES E. KEIRANS—First report of Hyalomma marginatum isaaci Sharif (Acari: [xodida: Ixodidae) from the Union of Myanmar, with a concurrent collection of H. hussaini Sharif SCOTT, DEBRA A. and ROBERT A. WHARTON—First report of Hesperoctenes fumarius Westwood (Heteroptera: Polyctenidae) from the Island of Dominica BOOK REVIEWS ACORN, JOHN—Tiger Beetles: The Evolution, Ecology, and Diversity of the Cicindelids, by David L. Pearson and Alfried Vogler BALL, GEORGE E.—Carabidae (Insecta: Coleoptera): Catalogue, by A. Larochelle and M.-C. Lariviere, /n Crosby, T. K., Series Editor, Fauna of New Zealand OBITUARY SLATER, JAMES A., THOMAS J. HENRY, and A. G. WHEELER, JR.—Halbert Marion Harris (1900-2000): Biographical sketch, described taxa, and list of publications on Heteroptera .. 1074 SOCIETY MEETINGS HECKMAN Bole Ne DEMS Da Cc. Bound-To-Please* 2003 N. MANCHESTER, INDIANA 46962 Nae ce a, veagbatente year t vt Pa sot NPT ran unten bactteae ete at $ hae are" ereneegep ww iseg Pej reeana SAP UANE