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Mi Se Ma it nt ne nf bi : per, : i ; hy malt a : cal ea RR AL sl Dy tet Pd. ae Ri Rr ite, i bd Oa): Ay 4% Ty aicd | a a i ) , ' ae ae a A j | Mi ¥ ea. 4 "y ; F NTR VOL. 93 JANUARY 1991 NO. 1 (ISSN 0013-8797) ee PROCEEDINGS of the ENTOMOLOGICAL SOCIETY PUBLISHED QUARTERLY CONTENTS BOTOSANEANU, L. and O. S. FLINT, JR.—Some Helicopsyche Von Siebold species from Cuba and Hispaniola with conspicuous androconial systems (Insecta: Trichoptera: Heli- CODSYVCHIGAE) 2 ey Re ale erg ee ee a ade oie 32 Re AE eae 7 URE ewe eT Ws COKENDOLPHER, JAMES C. and STANLEY R. JONES—Karyotype and notes on the male reproductive system and natural history of the harvestman Vonones sayi (Simon) (Opiliones, GOSMETIAAS) ne ae es ees ee ee ree So. 11 5] ARE OME: SOM! Coen RNR NS CARB Se Ret eee DAVIS, DONALD R., RICHARD C. KASSULKE, KEN L. S. HARLEY, and JOHN D. GILLETT —Systematics, morphology, biology, and host specificity of Neurostrota gunniella (Busck) (Lepidoptera: Gracillariidae), an agent for the biological control of Mimosa pigra L. GRISSELL, E. E.—A revision of Nearctic Chalcedectini (Chalcidoidea: Pteromalidae) with a INE WeNVOLld) CHECKIIST# Seize hs Ap lac ae ee Meee kod 25) IRS at sted eae De OR ON ai oe HENDRICKSON, R. M. JR., J. J. DREA, and MIKE ROSE—A distribution and establish- ment program for Chilocorus kuwanae (Silvestri) (Coleoptera: Coccinellidae) in the United STALESTHs Serr en ee 7 Rian, Cals? SM aNeee me Tica tn Le ME Ie RMR oy Te Ey ae te HENDRICKSON, R. M. JR., F. GRUBER, G. MAILLOUX, and J. J. DREA—Parasite colo- nizations against Crioceris asparagi (L.) and C. duodecimpunctata (L.) (Coleoptera: Chrys- ‘omelidae)in INorthyAmenicatromel983: to 1988.0 eee ee ee os a HOEBEKE, E. RICHARD and A. G. WHEELER, JR.—Anthribus nebulosus, a Eurasian scale predator in the Eastern United States (Coleoptera: Anthribidae): Notes on biology, recog- MitionyanGvestablishmenitee mere: Gils tetany nk). aay Soa Me Na ent, a a ee Re MARSH, PAUL M.—Description of phytophagous Doryctine Braconid from Brasil (Hyme- MO Ptera-STACONIGAG) meg) eecg oe EN ee ta tee. RR MONNMIR Slee Nes eae Bec ahem SA ro eles MATHIS, WAYNE N. and AMNON FREIDBERG— Review of Afrotropical beach flies of the tribe Canacini and subfamily Nocticanacinae (Diptera: Canacidae) .................... McDANIEL, B. and A. BOE—Life history studies, host records, and morphological description of genitalia of Eurytoma tylodermatis Ashm. (Hymenoptera: Eurytomidae) from South DAKO LANES Sekt SOR ee ee eee Che Ce Plo ae eye emt pee hegre hs ede ot Re a ea LO Te (Continued on back cover) of WASHINGTON 176 86 197 67 THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1991 DAvip R. SmiTH, President NORMAN E. WOODLEY, Treasurer Wayne N. MaArtuis, President-Elect Gary STECK, Program Chairman M. ALMA SOoLIs, Recording Secretary RUSSELL B. STEWART, Membership Chairman Ho.uis B. WILLIAMS, Corresponding Secretary JEFFREY R. ALDRICH, Past President JAMES B. STRIBLING, Custodian RosBerRT D. Gorpbon, Editor THOMAS J. HENRY, Associate Editor Publications Committee DONALD R. DAVIS TERRY L. ERWIN GEORGE C. STEYSKAL F. CHRISTIAN THOMPSON Honorary President CurRTIS W. SABROSKY Honorary Members LouIs—E M. RUSSELL ALAN STONE THEODORE L. BISSELL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, NHB 168, Smithsonian Insti- tution, Washington, D.C. 20560. MEETINGS. — Regular meetings of the Society are held in the Natural History Building, Smithsonian Institution, on the first Thursday of each month from October to June, inclusive, at 8 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.— Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $20.00 (U.S. currency) of which $18.00 is for a subscription to the Proceedings of the Entomological Society of Washington for one year. PROCEEDINGS.-— The Proceedings are published quarterly beginning in January by The Entomological Society of Washington, % Department of Entomology, NHB-168, Smithsonian Institution, Washington, D.C. Members in good standing receive the Proceedings of the Entomological Society of Washington. Nonmember subscriptions are $50.00 per year, domestic, and $60.00 per year, foreign (U.S. currency), payable in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE P. 218 OF THE JANUARY, 1991 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560. Editor: Robert D. Gordon, Systematic Entomology Laboratory, ARS, % Department of Entomology, Smith- sonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Books for Review: T. Henry, Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 13 February 1991 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. PROC. ENTOMOL. Sox 93(1), 1991, py A REVISION OF NEARCTIC CHALCEDECTINI (CHALCIDOIDEA: PTEROMALIDAE) WITH A NEW WORLD CHECKLIST E. E. GRISSELL Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, % U.S. National Museum NHB 168, Washington, D.C. 20560. Abstract. — Four Nearctic species of Chalcedectini are recognized in two genera: Dry- adochalcis texana (Brues), Amotura hyalinipennis (Ashmead), n. comb., Amotura ma- culipennis (Ashmead), n. comb., and Amotura caelata Grissell, new species. Euchrysia (= Amotura) similis Girault is synonymized under A. hyalinipennis. Each previously de- scribed species is redescribed, illustrated for the first time, and the previously undescribed sexes for all species are recognized for the first time. A key to Nearctic taxa is presented, and known geographic and host data are summarized. The Neotropical species Dryado- chalcis superba DeSantis is discussed in relationship to its early confusion with D. texana. A list of 15 New World species (with 8 new combinations) in 3 genera is given. Key Words: The group now referred to as Chalcedec- tini (Hymenoptera: Pteromalidae: Cleo- nyminae) (Boucek 1988) has had a long and varied nomenclatural history. Although rel- atively distinct in habitus, placement of Chalcedectini in a hierarchical classification has been problematic. At various times they have been recognized as a subfamily of the Cleonymidae (Ashmead 1904), as a sub- family of Podagrionidae (Peck 1951), as a family (Burks 1958, 1967), and as a sub- family of Pteromalidae (Graham 1969). The current placement as a tribe of Cleonyminae (Boucek 1988) is accepted as the basis for this paper. Chalcedectini are an uncommonly col- lected group of wasps, presumed to be par- asitic on xylophagous beetles. Though rare, the few nearctic species are each widely dis- tributed throughout the Nearctic and northern Neotropical Regions. According to Boucek (1988) the Chalcedectini are cir- cumtropical in distribution with the major- Amotura, Dryadochalcis, wood-boring beetles, parasitic wasp ity of species found in the Neotropical Re- gion. In the recent Catalog of Hymenoptera in America North of Mexico, Burks (1979) placed 2 genera and 4 species in the tribe. More recently Boucek (1988) revised the generic limits in his treatment of the Aus- tralasian genera and species. As a result of his revision, none of the 4 Nearctic species are currently placed in the correct genus. The intent of this paper is to clarify the generic status of the four described Nearctic species (one of which is synonymized), de- scribe the previously unknown sexes of the three remaining species, describe one new species, and present a checklist of New World taxa with eight new combinations. Dryadochalcis superba, a Neotropical spe- cies, is discussed in relation to its early con- fusion with the Nearctic species D. texana. Although I have collected, reared, and ac- cumulated specimens for over 10 years, fewer than 300 nearctic specimens have been seen (from 15 museums). Material was bor- Z PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rowed from the following sources (abbre- viations given are used in the text): CAS, California Academy of Sciences, San Fran- cisco, California; CDFA, California De- partment of Food and Agriculture, Sacra- mento, California; CNC, Canadian National Collection, Ottawa, Canada; BMNH, The Natural History Museum, London, En- gland; DCD, D. C. Darling Collection, To- ronto, Canada; FSCA, Florida State Collec- tion of Arthropods, Gainesville, Florida; MBR, Mueso Argentina de Ciencias Na- turales Bernardino Rivadavia, Buenos Aires, Argentina; Halstead Collection, J. A. Hal- stead, Fresno, California; Hespenheide Col- lection, H. A. Hespenheide, Los Angeles, California; HUM, Hope University Muse- um, Oxford, England; MLP, Museo La Pla- ta, La Plata, Argentina; ROM, Royal On- tario Museum, Toronto, Canada; TAMU, Texas A & M University; UCD, University of California, Davis, California; UCR, Uni- versity of California, Riverside, California; UI, University of Idaho, Moscow, Idaho; USNM, National Museum of Natural His- tory, Washington, D.C. Abbreviations for morphological terms used in the text are: F = flagellomere, MOL = midocellar length (between mid- and hin- docelli), OOL = ocellocular length (between hindocellus and eye), POL = postocellar length (between hindocelli), Mt = metaso- mal tergum. In numbering the metasomal (= abdominal, gastral) terga, I follow the system used by Gibson (1989) wherein Mtl is the petiole (virtually invisible), Mt2 is the first visible tergum, and Mt7 bears the spira- cles. Terga may be difficult to count because in some species (e.g. Amotura caelata, Fig. 10) larger terga cover smaller ones, or terga are either fused (Mt8+9) or so closely ap- pressed as to appear fused. These conditions are described for each species. CHALCEDECTINI | have seen the types of 11 of the 15 known New World Chalcedectini as well as at least that many undescribed species from the Neotropical Region. Although Boucek (1988) revised the higher classification of the Australasian Chalcedectini, there is still much to learn about the New World forms. In this paper I recognize 3 New World gen- era: Dryadochalcis DeSantis, Amotura Westwood, and Chalcedectus Walker. Bou- cek (1988) pointed out that the genus Chal- cedectus was based solely upon the type- species Chalcedectus maculicornis Walker. This Brazilian species has the temple ex- panded into vertical crests with transverse ridges and differs from Amotura in no other discernable way (type in BMNH, exam- ined). I am not convinced that monotypic genera based upon such apparently autapo- morphic conditions serve a purpose in un- derstanding evolutionary pathways. Two of the species included in this study (i.e. A. caelata and A. maculipennis) might just as easily be placed in monotypic genera based upon autapomorphies. Given our limited knowledge of the Chalcedectini, their rare- ness, and the profusion of diverse morpho- logical types apparent in just the 4 Nearctic species it seems best, at present, not to frag- ment the group into what would be mono- typic genera. Until the New World species are revised, I follow the generic concepts of Boucek (1988) in order to maintain some semblance of nomenclatural stability. Chalcedectini are recognized by a com- bination of the following characters: hind- femur enlarged and with teeth along the ventral edge, hindtibia arched, eyes diverg- ing ventrally, antennae inserted nearer to the clypeus than the midpoint of the head, and in females the antenna has an apical spicula. The antennae of both sexes are of diagnostic value, but they vary in appear- ance depending upon how the specimens are prepared. In critical point dried material (i.e. life-like) the apical clavomere of the female is domed with a membranous sensory re- gion and the spicula is readily apparent. In males, the apical two clavomeres are diag- onally truncate and the truncation is de- veloped as a membranous sensory region. VOLUME 93, NUMBER 1 In air-dried specimens the sensory region of both sexes usually collapses. This results in females having a truncate or sunken clava and the spicula may fold over and not be immediately evident. In males, the sensory region invaginates and causes the apex to become triangularly asymmetrical. KEY TO NEARCTIC CHALCEDECTINI 1. Apex of hindtibia without spur, curved out- ward as pointed projection, inner margin not apparent and tibial comb absent (Fig. 26); in female, Mt8+9 laterally compressed and many times longer than wide (Fig. 11) Oe yen Dryadochalcis texana (Brues) — Apex of hindtibia with 2 spurs (1 of which may be difficult to see), essentially concave, inner and outer margins of tibial apex equally ap- parent and tibial comb present (Fig. 25); in female, Mt8+9 not laterally compressed and as wide or wider than long (Figs. 9, 10) ..... EGAN EN eo. dca Amotura Westwood ... 2 2. Hindfemoral teeth large, separated by gaps (Fig. 6); female: Mt2 deeply split medially, longi- tudinally strigate (Fig. 10); male: metasoma ap- pears composed of 2 equally long terga, Mt2 not medially split, with strong to faint longi- tudinal sculpturing basally, other tergum strongly transversely strigate Beals dence Amotura caelata Grissell, new species — Hindfemoral teeth minute, scarcely separated (Figs. 5, 7); female: Mt2 entire medially (Fig. 9), polished; male: with 4 or 5 apparent meta- somal terga which are weakly sclerotized and difficult to differentiate 3. Axillae and axillulae highly polished, posterior region of scutum with well developed patch of silver setae; forewing with postmarginal vein much shorter than marginal vein (Fig. 20) .. Re eke Oe Amotura maculipennis (Ashmead) — Axillae and axillulae sculptured, posterior re- gion of scutum without patch of silver setae; forewing with postmarginal vein much longer than marginal vein (Fig. 18) ............... Ree cea ee Amotura hyalinipennis (Ashmead) Dryadochalcis DeSantis Dryadochalcis DeSantis, 1970: 25. Type species. — Chalcedectes [sic] texanus Brues. Orig. desig. Although DeSantis (1970) described the genus Dryadochalcis for texanus this change was not noted in the most recent catalog (Burks 1979) where texanus was placed in Chalcedectus. The essential characters sep- arating Dryadochalcis and Amotura are stressed in the key, and I can add one ad- ditional differentiating character to separate them. In Dryadochalcis the distal segment of the maxillary palpus is right-angled in relation to its point of attachment (Fig. 24), but in Amotura the segment is only slightly curved (Figs. 21-23). Dryadochalcis superba De Santis (Fig. 12) Dryadochalcis texanus of DeSantis, 1970: 26 (Figs. 5, 6), 1 2 (description of misiden- tified female specimen as fexanus). Dryadochalcis superba DeSantis, 1977: 26. New name for misidentified female tex- anus of DeSantis, 1970 (nec male of Brues 1907). Holotype °, Santa Trinidad, Par- aguay, MBR. [Examined]. Discussion.—This Neotropical species is included here because of the confusion be- tween it and the Nearctic species texana. Brues (1907) described Chalcedectus tex- anus based upon a male collected in Texas. In 1970, DeSantis described what he thought was the female of texanus based upon a sin- gle female from Paraguay. He also trans- ferred texanus to the new genus Dryado- chalcis. In 1977, DeSantis stated that his 1970 description of the female of “‘texanus” was a misidentification and that this de- scription actually referred to a new species which he named superba. I have examined the type of superba, as well as female spec- imens from Argentina and Brazil, and I agree with DeSantis that these specimens repre- sent a species distinct from texana. The dif- ferences between the 2 are not great and are discussed under texana. Material examined.—In addition to the type specimen I have seen 6 @ as follows: BRAZIL: 2 2, Amazonas, vic. Manaus, Re- serva Ducke, 24—25 July 1981 and | August 1981, G. B. Fairchild, flight trap near pond (FSCA); 1 °, Tonatins (BMNH); | 2, Ron- 4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-8. Chalcedectini. 1-4, Heads, frontal and dorsal views. 1. Amotura busckii. 2. A. hyalinipennis. 3. A, maculipennis. 4. Dryadochalcis texana. 5-8, Hind femora, outer view. 5. 4. maculipennis. 6. A. caelata. 7. A. hyalinipennis. 8. D. texana. don, 5-IV-1962, F. Plaumann (USNM). ARGENTINA: | 2, Tucuman, Trancas-Ta- canas, 1-30-XI-1968, L. Stange (MLP), and 1 2, 11 km W. Las Cejas (El Solidad), L. Stange, Malaise trap (FSCA). Dryadochalcis texana (Brues) (Figs. 4, 8, 11, 1327, 24) Chalcedectes texanus Brues, 1907: 106-107, 1 6. Holotype 4, U.S.A., Brownsville, Texas, USNM #42714. [Examined]. Dryadochalcis texanus (Brues), n. comb.: DeSantis, 1970: 26. Female (first and only specimen for spe- cies).— Length 13.7 mm. COLOR: Metallic violaceous, with blue and green reflections, except as follows: antenna black; wing veins and tarsomeres 2—5 brown; tarsomere | white; basal '4 of hindtibia faintly mahog- any. Wings infuscate brown, most distinct beneath parastigma and marginal vein. SCULPTURE: Head and mesosoma most- ly umbilicately punctured, reticulate (retic- ulate-rugose) on head from dorsum of scrobes to venter of head, mesopleuron smooth in upper half, lower half transverse- VOLUME 93, NUMBER 1 Figs. 9-16. Chalcedectini. 9-11, Metasomas, dorsal view (Mt = metasomal tergum). 9. Amotura hyalini- pennis. 10. A. caelata. 11. Dryadochalcis texana. 12-13, Plical carina of propodeum. 12. D. superba, detail of carina only. 13. D. texana, left half of propodeum with expanded detail of carina. 14-16, Scutellum and metanotum, side view (Ax = axillula). 14. 4. hyalinipennis. 15. A. caelata. 16. A. maculipennis. ly strigate, median area of propodeum heavily rugose; submedian panels of pro- podeum shiny with a few transverse wrin- kles; Mt2 essentially polished, remaining terga with faint transverse striations; hind- femur shagreened with minute, widely spaced punctures. SETATION: Head and mesosoma generally covered with silvery pubescence, dense patch of recumbent setae present behind propodeal spiracle; follow- ing areas asetose: scrobes, mesopleuron, median and submedian areas of propo- deum; metasomal terga essentially asetose dorsally, laterally with sparse, short setae. STRUCTURE: Head in facial view (Fig. 4) with polished area of scrobes extended to less than 2 diameter of midocellus, area above this with few weak punctures; scrobes laterally sharp nearly entire distance; inter- antennal area laterally sharp; malar distance 0.7 intermalar distance (19:26) and 0.7 x eye height (19:25). Head, in dorsal view (Fig. 4), with OOL:POL:MOL ratio as 5:6:4, ocellocular area with anteriorly curving 6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ee LO FEZ Pe | ("7 ) ee GaP me S Pea aN: / WV \ | y) 4 x ) oS ee \ | 2 j 6 Figs. 17-26. Chalcedectini. 17-20, Forewing venation, dorsal view. 17. Dryadochalcis texana. 18. Amotura hyalinipennis. 19. A. caelata. 20. A. maculipennis. 21-24, Maxillary palpus, outer view. 21. A. hyalinipennis. 22. A. maculipennis. 23. A. caelata. 24. D. texana. 25-26, Hindtibial apex, left = inner view, right = outer view. 25. Amotura spp. 26. Dryadochalcis spp. carinae, vertex 0.3 head width (20:60). Eyes bare. Scape essentially cylindrical, 5 x as long as wide (55:11), ratio scape : pedicel : F1-9 : apical spicula as 55:18:8:20:12:14:12: 11:9:5:4:3, pedicel 3 x as long as wide (18: 6). Propodeal plica obscured by setae that continue from lateral panel upward over edge of submedian panel (Fig. 13); hindfe- mur (Fig. 8) 2x as long as wide (60:30, ex- cluding teeth), with 5 large teeth. Forewing almost evenly setose except oval bare area distal to basal cell, lower half of costal cell (on dorsal surface), and area behind cubital vein (basally), ratio SM:M:PM5:S as 70:31: 40:10 (Fig. 17). Metasoma (Fig. 11) with Mt2-8+9 in ratio of 19:5:6:9:15:30:39 in dorsal view (Mt7 and 8 appear fused dor- sally, lateral margins difficult to see), Mt2 slightly emarginate medially, otherwise ter- ga essentially straight on hind margin. Holotype male.—Body length 6.6 mm. Coloration as for female, except metasoma tinged orange in basal half. Otherwise as for female except as follows: antenna with ratio of scape : pedicel : FI-9 as 44:12:5:14:12:10: 8:7:5:4:4 (mght antenna missing beyond F1), pedicel 2 x as long as wide (12:6), metasoma weakly sclerotized, basal half smooth, distal half faintly alutaceous. Material examined.—In addition to the type male, I have seen one female from U.S.A., Texas, Cameron County, 2-3 mi. south of Southmost, 7-XII-1978, E. E. Gris- sell, A. S. Menke, malaise trap, xeric ham- mock (USNM). Discussion.—Since its original descrip- tion in 1907, Brues’ species texana has been known only from the male. In 1978, Arnold Menke and I trapped a female specimen within a 10 mile radius of the type locality and this is certainly the female of the spe- cies. It differs only in expected secondary VOLUME 93, NUMBER 1 sexual characters. Females of Dryadochalcis texana differ from those of superba in the following characters: in texana the propo- deal plica (Fig. 13) is obscured by setae that continue from the lateral panel upwards, over the panel, and onto the submedian panel, whereas in superba (Fig. 12) the plica is a well-defined, single ridge and the setae which cover the lateral panel do not cross over onto the submedian panel; in fexana the midocellus is separated from the dorsal apex of the scrobal basin by a nearly smooth area with a few ill-defined punctures 1m- mediately below the midocellus, whereas in superba there is a sculptured area subequal in width to the midocellus. These characters should work for the male of both species as well, but that of superba is still unknown. DeSantis (1977) cited several differences to separate the species involving antennae and hindfemora, but these have not proved ad- equate. Amotura Westwood Polychroma Westwood, 1874: 140. Type species.—Polychroma histrionica Westwood. Des. by Gahan & Fagan, 1923: 117. Preoc. by Polychroma Dejean, 1833 (Coleoptera). Amotura Cameron, 1884: 130-131. Type species.—Amotura annulicornis Cameron. Monotypic. Polychromatium Dalla Torre, 1897: 88. Re- placement for Polychroma Westwood. Episystole Girault, 1927: 317. Type species.—Episystole poeta Girault. Orig. Desig. Neochalcodectus Masi, 1936: 68. Type species.—Neochalcodectus sinaiti- cus Masi. Monotypic. The above synonymy is taken from Bou- cek (1988) who explained the nomenclatur- al history of the genus. Amotura is now the correct name for all nearctic species previ- ously placed in Euchrysia Westwood (Burks 1979). Euchrysia (sensu strictu) was syn- onymized with Agamerion by Boucek (1988). Girault (1917) published a k ‘y to the 3 North American species of Evchrysia, but this is no longer useful. Amotura hyalinipennis (Ashmead), NEw ComBINATION (Figs. 2397,.9, 14.°18> 21) Euchrysia hyalinipennis Ashmead, 1896: 231, ?6 2. Lectotype ¢ (here designated), Argus Mts., California, USNM No. 3334. [Examined]. Euchrysia similis Girault, 1917: 15, 1°. Ho- lotype 2, Lawrence, Kansas, USNM No. 20464. [Examined]. NEw Synonymy. As explained in the variation and discus- sion sections below, this species appears to be geographically variable. Therefore I have redescribed the species based upon the lec- totype of hyalinipennis as the standard form against which variation is measured. Lectotype female.— Body length 4.0 mm. COLOR: General body color black with metallic green reflections especially on face, pronotum, thoracic venter, basal '2 of Mt6; hindfemur tinged purple; yellowish brown are: F1-5, palps, wing veins, tarsomeres 4— 5; tarsomeres 1-3 whitish. SCULPTURE: Coarsely reticulate (reticulate-rugose) on head from dorsum of scrobes to venter of head, largest pits ca 0.75 x diameter of mi- docellus (on frons reticulation much finer and granular in nature, scrobes transversely reticulate); mesosoma almost evenly retic- ulate dorsally; transversely strigate are pos- terior angle of sidelobe (scutum) between notaulus and axilla, and Mt5-6 basally; me- dian area of propodeum heavily rugose; submedian panels of propodeum with a few transversely rugose wrinkles; alutaceous to weakly reticulate are: lower '2 of pronotum laterally, tegula (weakly), venter of meso- soma, lower 73 of mesopleuron, axillula (weakly), legs (especially hindfemur), Mt2- 4, Mt8—9 (extremely weak, terga appear pol- ished in most views), posterior /2 Mt5-6, and entire Mt7; smooth are subalar area, 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON upper '3 of mesopleuron, dorsellum, nucha. SETATION: Head and mesosoma generally covered with silvery pubescence except fol- lowing areas bare: scrobes, mesopleural de- pression, mesepimeron, median and sub- median areas of propodeum (laterally with a patch of decumbent setae below spiracle); at least some of the pubescence near hind- femoral teeth noticeably longer than else- where on femur; tergal setation as follows: Mt2-6 without dorsal setae, Mt7-9 with sparse dorsal setae, laterally Mt2-5 with a few setae along dorsal edge, Mt6-9 evenly setose. STRUCTURE: Head in facial view with scrobes extended to within 1.5x di- ameter of midocellus, scrobes well defined laterally, fading dorsally into frons, inter- antennal area without median carina, lat- erally well defined, distance between outer margins of hindocelli greater than distance between inner margins of toruli (1.7 x, Fig. 2), malar distance 0.9 x intermalar distance and 0.6 x eye height (20:32), clypeus straight, in dorsal view ratio of OOL:POL:MOL as 1:4:3, frontovertex 0.3 head width (36: 120), scape cylindrical in basal '4, gradually flattening ventrally in upper 1, lateral ca- rina present only on mesal margin in upper ’3, scape 5.3 x as long as wide (53:10), ratio scape : pedicel : Fl1—-9 : apical spicula as 53: 20:8211211:1'0:9°8:7:6:6:6; pedicel 2.9.x as long as wide (20:7). Pronotum nearly ver- tical in lateral view, scutellum dorsally flat- tened, laterally with angled edge which sets off poorly defined, inward sloping face which is obscure where met by dorsellum (Fig. 14), hindfemur (Fig. 7) with numerous, saw-like teeth (each tooth tiny and not free from the next); forewing evenly setose except oval bare area distal to basal cell and area behind cubital vein at base of wing, ratio SM:M: PM:S as 50:18:40:6 (Fig. 18). Metasoma (Fig. 9) in dorsal view widest in basal '4, Mt2-9 with ratio of 23:4:5:9:20:12:7:5 (Mt8 and 9 may appear fused), terga essentially straight on hind margins. Male.—Body length 1.9-3.0 mm. Body color predominantly black with metallic blue or green reflections (or brassy red in south Florida specimens). Otherwise as for female except as follows: antenna with ratio of scape : pedicel : Fl—9 as 37:14:4:9:6:6:5:5:5: 4:7, pedicel 2 x as long as wide (14:7), F2-9 covered with recurved, evenly spaced, silver setae which give the flagellum a shiny ap- pearance in some lights; metasoma weakly sclerotized, Mt2 smooth, Mt3-8 alutaceous. Type material.—Euchrysia hyalinipennis was described from an unspecified number of specimens. Six localities were listed by Ashmead for his new species, and 6 females have type labels in the USNM collection. I consider these the only syntypical material, and designate the specimen from Argus Mts., California as lectotype. This specimen is the only complete one remaining but has its right antenna, left foreleg (beyond coxa), and right hindleg on a slide along with parts of the holotype of E. similis. Euchrysia similis was described from | female which is missing the left wings and right midleg. The left fore- and hindlegs (beyond coxae) and left anten- na are on a slide. Other material examined.—I have ex- amined 70 females and 138 males as follows (specimens in CAS, CDFA, CNC, DCD, FSCA, ROM, TAMU, UCD, UI, USNM): CANADA: Ontario, Quebec. UNITED STATES: California (2 paralectotypes), Ar- izona, New Mexico, Texas (paralectotype), Kansas (holotype—similis), Missouri (par- alectotype), Florida, Georgia, North Caro- lina, South Carolina, Virginia, District of Columbia, New Jersey, and Massachusetts. MEXICO: Baja California Sur, San Luis Po- tosi, Chiapas, and Yucatan. Distribution.—This is a widespread and relatively common species known from southern Canada (Ontario) to southern Mexico (Chiapas). Variation. — Females vary in length from 3.0 to 5.0 mm. Color has been used to dis- tinguish species in Amotura, but it is not always reliable. In females, mesosomal col- or varies from mostly blackish with metallic green or blue-green to brassy-violet (in some VOLUME 93, NUMBER 1 Florida specimens). Girault (1917) separat- ed his species similis from hyalinipennis pri- marily by the infuscated forewing of the for- mer and the hyaline forewing of the latter. Additionally, similis had flagellomeres all black while hyalinipennis had F2-5 reddish brown. In examining almost 200 specimens from the United States and Mexico it is not possible to segregate these two forms by these criteria. Basically, specimens from areas bordering the 40th parallel and from higher elevations tend to have dark antennae and infuscated wings (similis) while specimens from lower latitudes and elevations tend to have reddish brown or yellowish flagello- meres and no wing spot (hyalinipennis). Specimens from southern California (type locality) have F1-5 yellowish, or occasion- ally F2-4 are yellowish and F5 is darker yellow basally fading to black apically. Specimens from Texas, Florida, and lower elevations of Mexico have only F2-4 yel- lowish with F5 always black. Interpretation of the brownish (or yellowish) coloration, however, is fairly subjective. In some in- stances the color is outstanding in contrast to the remainder of the flagellum. In others, flagellomeres 2—4 are merely slightly lighter in color relative to the remainder. In all males the antenna is completely black (but with conspicuous silver setae). I see little practical application in using antennal col- oration for delimiting species. Interpreta- tion of the forewing infuscation is equally subjective in my opinion. It is not a matter of a distinct spot or its position, but rather the degree of intensity. Even the darkest in- fuscation is pale at best, and only visible under certain lighting (e.g. transmitted backlighting). In females the distance between the outer margins of the hindocelli averages 1.4 x longer than the distance between inner mar- gins of the toruli (1.3-1.7x, n = 10), the width of the frontovertex compared to the breadth of the head averages 0.30 x (0.29- 0.31, n = 10) and is independent of body size, and the malar to intermalar ¢ averages 1.0 x (1.0-1.1, n = 10). I have seen exceptionally flattened indi- viduals of this species in series of specimens from the same locality. In some cases the propodeum is nearly in the same plane as the scutellum. This arises, I believe, due to the host tunnel (in stems or twigs) being too shallow for normal pupation of the wasp. If pupation occurs, the resultant adult would be deformed. Flattened specimens have been collected as free-living individuals. Based upon available specimens, the fol- lowing descriptive material may be added to the description for the lectotype female: pronotum in dorsal view without apical ca- rina, with median polished longitudinal line (almost a carina), nearly diagonal strigose sculpture radiates from this line; scutellum laterally with distinct carina which fades to- wards posterior apex; hindcoxa without dorsal carina, outer face alutaceous and ase- tose. Hosts.— There are few specific rearing re- cords for this species. The published records (Burks 1979) include: Buprestidae: Chrys- obothris deleta Leconte, C. mali Horn; Cer- ambycidae: Oncideres rhodosticta Bates. Records taken from examined specimens are: Polycaon confertus LeConte (Bostrich- idae) in Manzanita (California), and Dicer- ca and Chrysobothris (Buprestidae) in Rhamunus californica Eschscholtz, ex stems of fire-damaged Desmanthus illinoensis (Michaux) MacMillan (Texas), ex pecan twig (Texas), ex citrus limbs (Florida), ex Gle- ditsia stems (New Jersey), ex Quercus du- mosa Nuttall stems (California), ex dead stems of Pinus ponderosa Douglas and Pseudotsuga menziesii (Mirbel) Franco (California), and ex mesquite stems (Cali- fornia). Discussion. — Yoshimoto (1984) reported a “possible new species of Chalcedectus” as the first Canadian record for the Chalce- dectinae (= Chalcedectini). I have examined those specimens and they are Amotura hy- alinipennis. Darling (1988: 2814, 2819) dis- 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cussed the phylogenetic implications of the labrum of this species (as Euchrysia). I have seen a voucher specimen (ROM collection) from this study and it is correctly identified. Amotura hyalinipennis can easily be dis- tinguished from other nearctic species by characters given in the key. It is most similar to the Puerto Rican species A. busckii which is known only from 5 specimens. A simple color character separates both sexes of both species: in A. hyalinipennis the scutellum is concolorous with the scutum, usually black to blackish green, whereas in A. busckii the scutellum contrasts markedly in color with the scutum (the former brilliant metallic brassy red to green, the later brilliant blue to purple-blue). It is not known if these color extremes represent geographic variation or not. The only morphological differences that I can find are on the head of females (cf. Figs. 1, 2). In 4. hyalinipennis the distance between the outer margins of the hindocelli is about 1.4 times greater than the distance between the inner margins of the toruli (range 1.3-1.7, n = 10, Fig. 2), whereas in A. busckii this distance is about 0.9 times (range 0.8-0.9, n = 4, Fig. 1). Also in fe- males, the width of the frontovertex com- pared to the face width is greater in A. hy- alinipennis (0.29-0.31, n = 10) than in A. busckii (0.20—0.25, n = 5). These differences are not striking and may not be valid. I have seen only | male of 4. busckii and they do not hold up for males. Amotura maculipennis (Ashmead), NEw CoMBINATION (Figs23754 116; 20-22) Euchrysia maculipennis Ashmead, 1896: 231, 6. Lectotype 4 (here designated), San- ta Cruz Mountains, California, USNM No. 3335. [Examined]. Female.—Body length 2.1-3.0 mm. COLOR: Body metallic green to copper, some specimens with purple reflections (es- pecially from Florida); brown are: scape and pedicel (both may have metallic reflections), apex of club, legs (except femora may be brownish yellow), wing veins, and well-de- fined infuscation beneath marginal vein extending to hind margin of wing; flagello- meres and tegula brownish yellow. SCULP- TURE: Densely reticulate (fine to granular) are: head from dorsum of scrobes to venter of head, scutum, dorsum of scutellum; high- ly polished are: axillae, axillulae, and apical vertical face of scutellum; polished with wi- despaced pits are: sides of head and frons; median propodeum vertically carinate, sub- median panels polished and with irregular carinae; Mt2-3 essentially polished, re- mainder weakly alutaceous as are legs. SE- TATION: Body covered with short, silvery pubescence except bare are: scrobes, me- sopleuron, median and submedian areas of propodeum, dorsum of Mt2-8 + 9; posterior half of scutum covered with dense, silvery pubescence, much more noticeable than elsewhere on body. STRUCTURE: Head in facial view (Fig. 3) with frons flat, meeting lower face at angle, scrobes ending at least 4 or 5 diameters before midocellus, scrobes well defined to top, interantennal area well defined laterally, malar distance subequal to intermalar distance and ca. 0.6 x eye height, clypeus straight, in dorsal view (Fig. 3) ratio of OOL:POL:MOL ca. 1:4:3, frontovertex ca. 0.33 x head width, scape almost 6 x as long as widest point in lateral view, ratio scape : pedicel : F1-9: apical spicula as 40: 13:3:6:6:6:6:5:4:4:3:3, pedicel ca. 2 x as long as wide. Pronotum vertical in lateral view, with prominent, sharp carina across ante- rior margin, scutellum bulging higher than scutum, apically with vertical face almost as wide as dorsellum (Fig. 16), hindcoxa with delicate dorsal carina (more prominent basally), hindfemur ventrally on outer edge with numerous, minute teeth each distinct from the next (Fig. 5); forewing essentially bare from parastigma to base of wing, basal vein with setal line, ventral surface of costal cell with several complete setal rows, ratio SM:M:PM5&S as 45:20:11:5 (Fig. 20). Meta- VOLUME 93, NUMBER 1 soma about as for hyalinipennis (as in Fig. 9), all terga straight on hind margins. Male. — Body length 1.9-2.3 mm. Differ- ing from female only in that the forewing is essentially hyaline. Type material.—The number of speci- mens in the type series was not stated, but the word “‘types’”’ was used. In the USNM there are 3 male specimens all from Santa Cruz Mountains, California and with USNM “type” labels. I consider this to be the extant type series. As no holotype was designated, I have designated one specimen as lectotype with a handwritten lectotype label. Other material examined.—I have seen 23 2 and 7 4 of this species from the follow- ing localities: MEXICO: 1 @. Coahuila, 33 mi. SE Saltillo, nr. Jame, 25 July 1963, H. & A. Howden (CNC). U.S.A.: ARIZONA: 1 ¢. Cochise Co., Huachuca Mts., Sunny- side, 31 August 1981, 6234 ft., D. C. Darling (DCD). CALIFORNIA: 1 ¢, Los Angeles Co., Claremont, Baker (USNM); Tulare Co., Ash Mountain Power Station #3, 3 June and 11 August 1984, R. D. Haines (2 9, 1 38), same but 1 and 8 October 1982, R. D. Haines, J. A. Halstead (1 2, 1 4) (all Halstead Collection). FLORIDA: Alachua Co., Gainesville, 1 6, 28 December 1972-4 Jan- uary 1973, H. V. Weems, Jr., insect flight trap, 1 9, 24 March, W. H. Pierce, Malaise trap (FSCA), 2 9, 2 6, 17-30 April 1987, D. Wahl, Malaise trap (CNC), 2 2, 4-18 June 1987, D. Wahl (CNC); Dade Co., 1 2, Fuch’s Hammock, near Homestead, T. S. Dickel and H. V. Weems, Jr., flight trap (FSCA), 1 9, S. Miami, 21 February—1 June 1986, S. & J. Peck, flight intercept trap (CNC). MARYLAND: 1 2, Montgomery Co., Be- thesda, ex Calllirhytis] crypta (Ashmead) (Cynipidae) (USNM). NEW MEXICO: 1 8, Lincoln Co., 4 km W Alto, 25 July 1982, G. Gibson (CNC). NORTH CAROLINA: 1 2, Northampton Co., 7 km S. Jackson, 10 July/23 September 1987, BRC Hym. Team, flight intercept trap, bald cypress swamp (CNC). TEXAS: 1 2, Brazos Co., College Station, 25 March 1967, J. C. Schaffner (TAMU); 4 9, 2 6, Cameron Co., | mi. | Southpoint Ranch, 5-6 July 1982, G. Gib son (CNC); 1 2, Montgomery Co., 8 mi. S. Conroe, 6-12 April, 1987, Wharton, Wang, Praetorius (TAMU); 1 2, Somervell Co., 10 mi. W. Glen Rose, 2 May 1975, J. C. Schaff- ner (TAMU). Distribution.— This species, although un- commonly collected, appears to be wide- spread across the entire southern half of the United States and extends into northern Mexico (Saltillo). Host.—The record cited above for Cy- nipidae is possible, though suspect. Several other species of Amotura have been asso- ciated with oak, but without specific host association. Amotura hyalinipennis was reared from stems of oak and A. caelata was taken feeding at honeydew of oak galls. The true host is most likely a beetle of some sort, but this remains to be demonstrated. Discussion.—Amotura maculipennis is distinctive among New World Chalcedec- tini based upon the highly polished axillae and axillulae, the relatively short postmar- ginal vein (compared to marginal), the flat- tened and angled frons, the bulging scutel- lum, the patch of silver setae on the posterior of the scutum, and the basally bare fore- wing. No other known species has any of these characters, and the species is so dis- tinctive that it does not appear to be related to any other New World taxon. Amotura caelata Grissell, NEw SPECIES (rigss.6,. 1021571925) In the following description, numbering of metasomal terga is based upon the true tergal position, not the apparent number. Thus, visible tergum 3 is actual metasomal tergum 7 (Fig. 10). Mt3—5 are weakly scler- otized and are covered by Mt2. Holotype female.— Body length 6.6 mm. COLOR: Entirely black with faint purplish blue reflections on face and mesosomal dor- sum; brown are: flagellomeres 1-4, wing 12 veins, tarsomeres 3-5: whitish are: tarso- meres 1-2, basodorsal spot on hindfemur (Fig. 6). SCULPTURE: Coarsely reticulate (reticulate-rugose) on head from dorsum of scrobes to venter of head, largest pits ca. 0.5 x diameter of midocellus, on frons re- ticulation much finer and granular in na- ture, scrobes transversely reticulate; meso- soma evenly reticulate dorsally; Mt2 (Fig. 10) longitudinally strigate, transversely stri- gate are anterior 74 of Mt6 and Mt7; median propodeum heavily rugose vertically, sub- median panels of propodeum heavily ru- gose transversely; alutaceous to weakly re- ticulate are legs (especially hindfemur); smooth are subalar area, upper *4 of me- sopleuron, nucha. SETATION: Entire body covered with short silvery pubescence ex- cept asetose are: scrobes, mesopleuron, me- dian and submedian areas of propodeum (laterally densely covered with semierect se- tae below spiracle), dorsum of metasoma; pubescence on hindfemoral teeth (both in- ner and outer row) noticeably longer than elsewhere on femur (Fig. 6). STRUCTURE: Head in facial view with scrobes extending to within 1.5 diameters of miudocellus, scrobes well defined laterally fading dorsally into frons, interantennal area without me- dian carina, laterally well defined, malar distance 1.0 x intermalar distance and 0.6 x eye height, clypeus straight, in dorsal view ratio of OOL:POL:MOL as 4:15:13, front- overtex 0.36 head breadth, scape cylin- drical in basal ', gradually flattening ven- trally in apical 4 and with lateral carina on mesal margins, scape 4.5 x as long as widest point in lateral view, ratio scape : pedicel: Fl-9 : apical spicula as 46:17:7:10:9:8:6:5: 4:3:3:3, pedicel 4.3 x as long as wide. Pro- notum nearly vertical in lateral view, with- out anterior carina, scutellum dorsally flat, apically with angled edge that sets offinward sloping face as wide as dorsellum (Fig. 15), hindcoxa without dorsal carina, hindfemur ventrally on outer edge with 9 or 10 prom- inent teeth and several minute sawlike teeth (Fig. 6); forewing evenly setose except oval 7 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bare area distal to basal cell and area behind cubital vein bare for 3 length of hindmargin of wing from base to apex, ratio SM:M:PM:S as 100:23:61:10 (Fig. 19). Metasoma (Fig. 10), in dorsal view, widest in basal 3, only 4 terga visible (Mt2, 6, 7, 8+9) with ratio of 8:12:7:5, Mt2 deeply emarginate, re- mainder straight on hind margin. Male.— Body length 2.9-4.3 mm. As for female except metasoma which differs as follows: Mt2 and Mt7 occupy almost entire length of metasoma (Mt3-6 hidden under Mt2, rarely one may be visible as narrow band), each comprising about half meta- somal length, Mt2 without median emar- gination, at least basal half lightly sculp- tured but sculpture may reach apex, Mt7 strongly transversely strigate (as in female). Type material.—Holotype 2, USNM; U.S.A., California, “Pinon Flat, Santa Rosa M,” 21-V-41, D. J. and K. N. Knull Collrs. Paratypes, 16 °, 33 6 as follows: U.S.A.: ARIZONA: Graham Co., | 2, 0.9 mi. along rd. to Mariyilda Canyon from hwy. 666, H. B. Leach (CAS); Pima Co., 1 4, Florida Wash, | August 1979, C. L. Bellamy (Hes- penheide Collection); Santa Cruz Co., 1 9, 2 mi. N. Nogales, 2 September 1981, 3865 ft., on Helianthus, D. C. Darling (Cornell University Collection). CALIFORNIA: Tu- lare Co., 3 2, 2 6, Ash Mountain, Kaweah Power Station (No. 3), May, Aug., Sept., and Oct. 1982-83, J. A. Halstead, R. D. Haines, D. J. Burdick (Halstead Collection); Monterey Co., | 2, Carmel Valley road (1 mi. NW Arroyo Seco road), 11 May 1967, H. J. Leach, ex dead Quercus containing Psoa quadrisignatus (Horn) (Bostrichidae); Riverside Co., 1 2, 4 6, San Timoteo Can- yon, 13-14 Sept. 1972, M. Wasbauer, A. Hardy, at honeydew of Disholcaspis eldor- adensis (Beutenmeuller) (Cynipidae) galls on Quercus lobata Nee (CDFA); Imperial Co., 1 2, Westmoreland, 31 May 1930, P. H. Timberlake, on Phacelia (UCR); 1 9, 1.5 mi. W. Winterhaven, 18 June 1978, C. Bel- lamy (Hespenheide Collection). FLORIDA: Alachua Co., 2 , 2 mi. N. Gainesville, 27 VOLUME 93, NUMBER 1 August and 25 September 1974, H. Davis and W. Jetter, ramp trap; Highlands Co., 1 2, 4 6, Archbold Biological Station, 19-22 March 1987, D. Wahl (American Ento- mological Institute, CNC); Marion Co., 1 9, 4 6,9 mi. SSW. Ocala, Kingsland Country Estates, 19 September to 19 October 1975, J. Wiley, Malaise trap in turkey oak (FSCA); Monroe Co., 1 6, Big Pine Key, 10 April 1970, W. W. Wirth, Malaise trap (USNM), also 1 2, 1-31 October 1985 and 2 4, 1-31 February 1986, S. & J. Peck, mangrove hardwood transition (CNC); Osceola Co., | 2, Ocala, 24 Oct. 1914 (USNM). NEW MEXICO: Dona Ana Co., | 2, 5 mi E. Las Cruces, 12 April 1965, R. M. Bohart (UCD); Hidalgo Co., 1 6, 4 mi. S. Rodeo, 28 June 1969, V. D. Roth (Hespenheide Collection) and 1 2, Rodeo, 28 August 1959, 4000 ft., H. E. Evans (DCD); 4 4, 9.3 mi. W. Animas, 26-30 July 1982, G. A. P. Gibson, sweeping Chilopsis linearis (Cavanilles) Sweet (CNC). TEXAS: Cameron Co., 2 4, 2.3 mi. S. South- most, 8 Dec. 1978, E. E. Grissell, A. S. Menke, ex stem Desmanthus illinoensis (USNM); Uvalde Co., 1 2, 3 mi. NW Uval- de, 4 May 1977, T. Eichlin, W. Wasbauer, Malaise trap (CDFA). MEXICO: Baja: | 4, 4.6 mi. E. Valle de Trinidad, 26 May 1979, C. L. Bellamy (Hespenheide Collection); 1 6, 25 mi. W. La Paz, 30 August 1959, K. W. Radford, F. G. Werner (CAS). OAXA- CA: | 46, Puerto Escondido, 15 July 1985, J. Woolley, G. Zolnerowich (TAMU); SAN LUIS POTOSI: 2 4, 7 mi. S. Ciudad Valles, 20 December 1970, P. H. and M. Arnaud (CAS); SONORA: 1 ¢, 1 4, 10 mi. SE. Al- amos, 29 June 1963, F. D. Parker, L. A. Stange (UCD). Other material examined (too poor to designate as type material). ARIZONA: 1 2, Portal, 18 June 1956, R. and K. Dreisbach (USNM). CALIFORNIA: 1 ¢, Los Gatos, 28 October 1918, H. E. Burke, ex pupal cell Chrysobothrus mali Horn (Coleoptera: Bu- prestidae) (USNM). Etymology.—From the Latin caelo, meaning engraved, in reference to the turing of the metasoma. Hosts. — Amotura caelata has been reared from a pupal cell of the buprestid Chryso- bothrus mali, from dead Quercus containing Psoa quadrisignatus, and from dead stems of Desmanthus illinoensis. It would appear to be a parasite of wood boring beetles. Variation.—Females of caelata vary in length from 4.5 to 6.6 mm. The coloration appears stable throughout the entire geo- graphic range, except that Florida speci- mens are weakly purplish overall rather than black. In females the yellow spot on the apex of the hindtibia varies from '/, to ¥, the tibial length. In males the color is less obvious (% to % tibial length) and, in a few cases nearly disappears. Rarely, in some females, one of the hidden metasomal terga (Mt3-S) is vis- ible in the emargination of Mt2. Discussion. —It is of interest that both 4. caelata and A. hyalinipennis have been reared from the same plant stems and lo- cality (Desmanthus illinoensis in South- most, Texas) at the same time. These two species and A. maculipennis have also been collected at the same locality in California (Tulare County, Ash Mountain Power Sta- tion), but not at the same time: A. caelata was collected in May, September, and Oc- tober; A. hyalinipennis in August; and 4A. maculipennis in October. Darling (1988: 2814, 2819-2820, 2832) discussed the phylogenetic implications of the labrum of this species which he referred to as “Chalcedectes [sic] sp.” He illustrated the free labrum (his Fig. 20) which he con- sidered the plesiomorphic state among Chalcidoidea. I have seen the specimen (in DCD collection) upon which his discussions were based and can confirm that it is Amo- tura caelata. Among Nearctic species, A. caelata su- perficially resembles 4. hyalinipennis. It 1s, however, distinct in the structure of the me- tasoma (Fig. 10, especially the emargination of tergum | and the strigate sculpture), the apex of the scutellum (Fig. 15, brim-like, 14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON angled apex), and the hindfemora (Fig. °6, enlarged, separated teeth). These characters are unique among Nearctic species, but the character states of the tergal emargination, scutellum, and hindfemur can be found in Neotropical species in different combina- tions. Because the tropics seem to be the center of diversity for Chalcedectini, a thor- ough analysis of characters and states must await a study of Neotropical species. Checklist of New World Chalcedectini I have seen the types of all specimens except guaraniticus Strand (discussed by Boucek 1959), septemdentatus pallidipes Roman, and annulicornis Cameron. The type locality follows depository. Amotura annulicornis Cameron.—Amotura annuli- cornis Cameron, 1884: 131. BMNH: Nic- aragua. annulipes (Ashmead) n. comb.—Chalce- dectes annulipes Ashmead 1904: 483. USNM: Brazil, Matto Grosso. busckii (Ashmead) n. comb.—Euchrysia busckii Ashmead, 1900: 256. USNM: Puerto Rico. caelata Grissell.—See text. guaraniticus (Strand) n. comb.—Polychro- matium 16-dentatum var. guaraniticum Strand, 1911: 95. Berlin Zoological Mu- seum: Paraguay. histrionicus (Westwood) n. comb.—Poly- chroma histrionica Westwood, 1874: 141. HUM: Brazil, Para. hyalinipennis (Ashmead).—See text. lanei (DeSantis) n. comb.—Chalcedectus lanei DeSantis, 1970: 22-23. MLP: Bra- zil, Serra do Navio. maculipennis (Ashmead).—See text. regalis (Westwood) n. comb.—Polychroma regalis Westwood, 1874: 141. HUM: Bra- zil, Amazonas. sedecimdentatus (Westwood) n. comb.— Polychroma 16-dentata Westwood, 1874: 141. HUM: Brazil, Para. septemdentatus (Westwood) n. comb. — Polychroma 7-dentata Westwood, 1874: 142. HUM: Brazil, Amazonas. 97-dentatus pallidipes Roman. Chal- cedectus 7-dentatus var. pallidipes Ro- man, 1920: 12. Type depository ?: Bra- zil, Amazonas. (Syn. questionable, see Boucek 1959.) Chalcedectus Walker maculicornis Walker.—Chalcedectus mac- ulicornis Walker, 1852: 47. BMNH: Bra- zil, Para. Dryadochalcis DeSantis texana (Brues).—Chalcedectes texanus Brues, 1907: 106-107. USNM: U.S.A., Texas. superba DeSantis.—Dryadochalcis superba DeSantis, 1977: 26. MBR: Paraguay, Santa Trinidad. ACKNOWLEDGMENTS During the overly long course of this study I have had many people to thank (abbre- viations refer to museums listed in the in- troduction). Of special note are L. DeSantis (MLP), Z. Boucek (BMNH), and M. Gra- ham (HUM) who loaned me valuable type material; M. Fritz (MBR) who located spec- imens that were not easily found; and the following curators who represent the spirit behind their museum’s acronym: Paul Ar- naud (CAS), D. Chris Darling (ROM), Gary Gibson and Carl Yoshimoto (CNC), Saul Frommer (UCR), Robert Schuster (UCD), Lionel Stange, J. Wiley, and Harold Green- baum (FSCA), Marius Wasbauer (CDFA), and Jim Woolley (TAMU). Additionally I thank Chris Darling, Jeff Halstead, and Henry Hespenheide for loaning material from their private collections. I thank Bob Gordon and Dick White for helping with names of Coleoptera. I am also grateful for the reviews of this manuscript provided by R. V. Peterson, P. M. Marsh, D. C. Darling, and G. A. P. Gibson. VOLUME 93, NUMBER 1 LITERATURE CITED Ashmead, W.H. 1896. Descriptions of new parasitic Hymenoptera. Transactions of the American En- tomological Society 23: 179-234. 1900. VI. Report upon the Aculeate Hy- menoptera of the islands of St. Vincent and Gre- nada, with additions to the parasitic Hymenoptera and a list of the described Hymenoptera of the West Indies. Transactions of the Entomological Society of London 1900 (2): 207-367. . 1904. Classification of the chalcid flies. Mem- oirs of the Carnegie Museum 1(4): v-ix, 225-551. Boucek, Z. 1959. On Chalcedectus sinaiticus (Masi) from the near east, and Ch. guraniticus (Strand), from Paraguay, and new synonymy. Acta Ento- mologica Musei Nationalis Pragae 33:483-486. 1988. Australasian Chalcidoidea. C.A.B. In- ternational, Wallingford, U.K. 832 pp. Brues, C. T. 1907. Notes and descriptions of North American parasitic Hymenoptera IV. Bulletin of the Wisconsin Natural History Society 5: 96-111. Burks, B. D. 1958. Chalcidoidea, pp. 62-84. Jn Krombein et al. Hymenoptera of America north of Mexico, synoptic catalog. United States De- partment of Agriculture, Monograph No. 2, First Supplement. . 1967. Chalcidoidea, pp. 213-282. Jn Krom- bein et al. Hymenoptera of America north of Mex- ico, synoptic catalog. United States Department of Agriculture, Monograph No. 2, Second Supple- ment. . 1979. Pteromalidae, pp. 768-835. Jn Krom- bein et al. Catalog of Hymenoptera in America north of Mexico. Smithsonian Institution Press, Washington, D.C. Cameron, P. 1884. Biologica Centrali-Americana. Hymenoptera 1: 81-144. Darling, D.C. 1988. Comparative morphology of the labrum in Hymenoptera: The digitate labrum of Perilampidae and Eucharitidae. Canadian Journal of Zoology 66: 2811-2835. Dalla Torre, C.G. 1897. Zur Nomenclature der Chal- cididen-Genera. Wiener Entomologische Zeitung 16: 83-88. DeSantis, L. 1970. Nota sobre dos Calcedectinos Neotropicales. Revista del Museo de La Plata (n.s.) 11: 21-26. . 1977. Nota sobre el genero ““Dryadochalcis.” Neotropica 23: 10, 26. Gahan, A. B. and M. M. Fagan. 1923. The type spe- cies of the genera of Chalcidoidea or chalcid-flies. Bulletin of the United States National Museum 124: 1-173. Gibson, G. A. P. 1989. Phylogeny and classification of Eupelmidae, with a revision of the world genera of Calosotinae and Metapelmatinae. Memoirs of the Entomological Society of Canada 149: 1-121. Girault, A. A. 1917. The North American species of Euchrysia. Bulletin of the Brooklyn Entomological Society 12: 15. 1927. Notes on and descriptions of chalcid wasps in the South Australian Museum. Records of the South Australian Museum 3: 309-338. Graham, M. W.R.de V. 1969. The Pteromalidae of northwestern Europe. Bulletin of the British Mu- seum (Natural History), Entomology, Supplement 16: 1-908. Masi, L. 1936. Nuovo genere di Chalcodectini rac- colto nei Sinai. Bollettino della Societa Entomo- logica Italiana 68: 67-71. Peck, O. 1951. Chalcidoidea, pp. 410-594. Jn Muese- beck et al. Hymenoptera of America north of Mex- ico, synoptic catalog. United States Department of Agriculture, Monograph No. 2. Roman, A. 1920. Wissenschaftliche ergebnisse der schwedischen entomologischen Reise des Herrn Dr. A. Roman in Amazonas 1914-1915. 3. Hy- menoptera. 2. Chrys:didae and Chalcididiae. Ar- kiv for Zoologi 12(19): 1-30. Strand, E. 1911. Neue und wenig bekannte exotische Arten der Chalcididengattungen Megastigmus Dalm., Mesodiormorus Strand (n.g.), Polychro- matium D.T. und Leucospis F. Wiener Entomo- logische Zeitung 30: 93-99. Walker, F. 1852. VII. Notes on Chalcidites, and de- scriptions of various new species. Annals and Magazine of Natural History 10(2nd ser.): 45-48. Westwood, J. O. 1874. Thesaurus Entomologicus Oxoniensis. Oxford. xxiv, 205 pp, + 40 pls. Yoshimoto, C. M. 1984. The families and subfam- ilies of Canadian chalcidoid wasps. The insects and arachnids of Canada, Part 12, Agriculture Canada Publication 1760. 149 pp. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 16-44 SYSTEMATICS, MORPHOLOGY, BIOLOGY, AND HOST SPECIFICITY OF NEUROSTROTA GUNNIELLA (BUSCK) (LEPIDOPTERA: GRACILLARIIDAE), AN AGENT FOR THE BIOLOGICAL CONTROL OF MIMOSA PIGRA L. DONALD R. DAvis, RICHARD C. KASSULKE, KEN L. S. HARLEY, AND JOHN D. GILLETT (DRD) Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (RCK & KLSH) CSIRO Division of Entomology, Long Pocket Laboratories, Private Bag No. 3, Indooroopilly, Australia 4068; (JDG) De- partment of Primary Industries and Energy, PO Box 858, Canberra, Australia 2601. Abstract. —The systematics and morphology of the gracillariid moth, Neurostrota gun- niella (Busck, 1906), from Mexico are reviewed. The biology and host specificity of this moth were studied in quarantine facilities in Brisbane, Australia. Eggs were usually laid singly on the ventral surface of mature leaves of Mimosa pigra L. The first two larval instars were sap-feeding miners in the leaf pinnules. Instars three to eight were tissue- feeding stem borers. Pupation occurred outside the stem within a silken cocoon festooned with frothy globules excreted by the larva. One hundred species of plants, either closely related to M. pigra or of economic or environmental importance, were tested in host specificity studies as part of an assessment of the suitability of N. gunniella as a biological control agent for M. pigra. The only plants which supported development of N. gunniella were the two introduced weeds, M. pigra and M. pudica, and four Australian Neptunia species. Larval mortality was high on the Neptunia species. These studies showed that N. gunniella would not cause damage to any plant other than the target weed, M. pigra in Australia. N. gunniella was first released near Darwin, Australia in February, 1989. Key Words: Lepidoptera biology, Gracillariidae, Neurostrota, Mimosa pigra, biological control, host specificity Mimosa, Mimosa pigra L. (Legumino- laide River, with major infestations on the sae), iS native to tropical America (Burkart 1948) where it usually occurs as small clumps of multi-stemmed plants growing in seasonally flooded habitats (Fig. 1). It prob- ably entered the Northern Territory of Australia during the 20 years prior to 1891 (Miller & Lonsdale 1987), underwent a pop- ulation explosion in the late 1970’s and now infests 45,000 ha of wetlands (Lonsdale and Segura 1987). Dense to scattered thickets of M. pigra cover the floodplains of the Ade- Finniss, Mary, and East Alligator Rivers (Pitt and Miller 1989, Miller 1982). M. pi- gra is a designated Class ““A” noxious weed in the Northern Territory (Noxious Weeds Ordinance 1962), is a serious weed in Thai- land (Napompeth 1982) and is spreading in Burma, Laos, Kampuchea, Vietnam, In- donesia and Malaysia (Napompeth, B., pers. comm.). In the Northern Territory it pre- vents access for irrigation and stock water- ing, makes livestock mustering difficult and VOLUME 93, NUMBER 1 interferes with the use of watercourses for recreation (Miller et al. 1981). On flood plains it out-competes other plants and se- riously reduces dry season grazing for cattle and buffaloes. There and in Southeast Asia it occurs as huge, impenetrable, near mono- specific thickets. Two other Mimosa species, M. invisa Mart. and M. pudica L. have been intro- duced into and become weeds in Australia (Kleinschmidt and Johnson 1977). M. in- visa 1s confined to Queensland and M. pu- dica is widespread throughout Queensland, northern New South Wales and the North- ern Territory. A program for biological control of MM. pigra in Australia was initiated by CSIRO and the Northern Territory Department of Primary Industry and Fisheries in 1979 and extended to Thailand through a program of the Australian Centre for International Ag- ricultural Research (ACIAR) in 1983. In in- vestigations in Mexico, adults of a gracil- lariid moth Neurostrota gunniella (Busck) frequently emerged from tips of M. pigra and occasionally from tips of another le- gume native to Mexico, Neptunia plena (L.) Benth. Following preliminary tests in Mex- ico for specificity to M. pigra, N. gunniella was imported into Australia for detailed evaluation in quarantine. This paper gives details of studies on the systematics and morphology of N. gunniella conducted at the Smithsonian Institution as well as studies of its biology and host spec- ificity carried out by the Division of Ento- mology, CSIRO, Australia. SYSTEMATICS AND MORPHOLOGY Neurostrota gunniella (Busck) Gracilaria [sic] (Dialectica) gunniella Busck, 1906: 731. Acrocercops gunniella (Busck) Meyrick, 1912a: 16, 1912b:. 44. Barnes and McDunnough, 1917: 188. Neurostrota gunniella (Busck) Ely, 1918: 38, 41, 68. McDunnough, 1939: 97. Vari 1961: 41. Davis, 1983: 9; 1984: 26 Neurostrata [sic] gunniella (Busck) Ely, 1918: 41. Busck, 1934: 179. Adult (Fig. 2).— Forewing length: 4, 3.3- 4.4 mm; 2, 3.2-4.6 mm. Small moth with dark coppery brown forewing possessing a single, oblique, cream white stria at distal third of costa and hind margin bordered with cream. Male valva rhombiform, sim- ple; seventh abdominal segment with a sin- gle pair of elongate hair pencils and a more basal tuft of short, spatulate sex scales. Fe- male bursa with a pair of diamond shaped signa. Head (Fig. 3): Vestiture smooth, scales curving down over frons; vertex cream to pale buff at middle, becoming darker brown laterally; a patch of brownish fuscous scales at anterior rim of eye below antenna; lower frons mostly silvery white. Antenna ca. |.2 x length of forewing, covered with dark brown scales dorsally and dull white ventrally. Ocellus absent. Interocular index ca. 1.1. Mandible absent. Maxillary palpus mod- erately long, four segmented, silvery white dorsally, grayish brown ventrally. Haustel- lum elongate, over 2 x length of labial pal- pus, coiled in repose. Labial palpus slender, elongate, over 2 vertical diameter of eye, curved dorsally; vestiture silvery white dor- sally, grayish brown ventrally and laterally. Thorax: Pronotum dark coppery brown with a broad cream to pale buff median stripe; venter dark brown laterally, silvery white over coxa. Wing venation as shown in Fig. 4. Female frenulum consisting of two bristles (Figs. 9-11), male of one bristle, with basal fusion of ca. 5—6 setae (Figs. 12-14). Both sexes with a secondary pseudofrenu- lum arising near termination of Sc and con- sisting of usually 3—4 piliform scales tightly overlapping (Figs. 9, 15-18). Pseudofrenu- lum coupling with a row of stiff, specialized scales located along the ventral hind margin in the anal area of the forewing (Figs. 19, 20). Forewing mostly dark coppery brown 18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1,2. 1, A dense concentration of Mimosa pigra near Acapulco, Mexico. 2, Neurostrota gunniella, adult 2. forewing length 4.2 mm. VOLUME 93, NUMBER 1 Figs. 3-8. Neurostrota gunniella. 3, Head, frontal view (0.25 mm). 4, Wing venation, pf = pseudofrenulum. 5, Male genitalia, ventral view (0.25 mm). 6, Aedoeagus. 7, Female genitalia, lateral view (0.5 mm). 8, Ventral view. (Scale lengths in parentheses.) 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with a single, cream-white stria extending obliquely from distal third of costa about half the distance to tornus; a single, narrow, silver fascia transversing subapex; fringe with a pair of dark brown striae curving around apex; hind margin bordered with cream to tornus; fringe along hind margin uniformly dark gray. Hindwing uniformly dark gray. Legs dark brownish fuscous with apex of tibia and most of tarsal segments banded with cream; tarsi of hindleg mostly dull white to pale buff. Abdomen: Dark brownish fuscous dor- sally, mostly cream to white ventrally, with 5 slender, dark fuscous, oblique bands lat- erally across segments A2-6. Male with a single pair of elongate hair pencils and a more basal tuft of short, spatulate sex scales lateral on A7. A7 greatly reduced. Tergite 8 enlarged, well defined, hoodlike, extending caudad of lateral sex scaling on A7. Male genitalia (Figs. 5, 6): Uncus absent. Vinculum, triangular with broad, lateral projections dorsad to base of valva. Valva rhombiform, with outer margin relatively straight and oblique. Aedoeagus relatively simple, lightly sclerotized, equalling length of valva; cornuti absent; phallobase 1.5 x length of aedoeagus. Female genitalia (Figs. 7, 8): Anterior and posterior apophyses equally developed, short. Ductus bursae shorter than length of corpus bursae; seminal duct arising midway along ductus. Corpus bursae well devel- oped, ovoid, with a pair of roughly dia- mond-shaped signa. Egg (Figs. 27-31).—Elongate-ovoid, with a minutely dimpled chorion; average length 0.357 mm, width 0.164 mm (n = 20). Mi- cropyle with 3-6 oval rings clustered around a minute central depression. Larva (Figs. 32-60, 63-70).— Hyper- metamorphic; first two instars with highly modified, depressed body for sap-feeding in leaf pinnules; maximum length, 1.2 mm. Remaining six instars of typical caterpillar form with cylindrical body, boring in stems of host. Length of largest larva 8.0 mm; maximum body width | mm; maximum head width 0.54 mm; body color instars 1- 7 cream, instar 8 dark blue-green, later re- verting to cream. SAP-FEEDING INSTARS Head: Greatly depressed, triangular (Figs. 32, 39). Most setae lost or reduced; dorsal cranium with only two small pairs of setae preserved, P2 near basal lateral margin and L1 above median (third) stemma. Labrum (Fig. 33) reduced, 0.64 the width of labial lobe, deeply clefted about 0.25 its length; only one pair of setae preserved; venter of labrum with dense concentration of short epipharyngeal spines (Fig. 37). Mandibles large, greatly flattened, with three primary cusps dorsally; innermost cusp broad with serrated margin; outermost cusp with a dor- sal groove and a fourth, shorter cusp arising ventrad. Labial lobe (Figs. 34, 35) with an- terior margin slightly depressed at middle; anterior margin and dorsal surface densely covered with short hypophyrangeal spines (Fig. 38) similar to epipharynx; spinneret vestigial, reduced to a minute, flushed open- ing ca. 0.86 wm in diameter. Ventral cra- nium with only a single pair of substem- matal setae (SS3?). Maxillary and labial palpi absent. Antenna reduced, with three rela- tively stout sensilla basiconica and three shorter, more slender sensilla (Fig. 36). Five pairs of stemmata present in an irregular lateral row; an anterior row of three contig- uous stemmata followed by two more wide- ly spaced stemmata posteriorly. Body: Setae extremely reduced. Legs, pro- legs, and crochets absent. TISSUE-FEEDING INSTARS Head: Approximately round (Figs. 40, 51) with full complement of mouthparts, dark reddish brown. Frons (Fig. 64) moderately long, about 0.66 the distance to epicranial notch. Ecdysial line terminating at epicra- nial notch. Chaetotaxy relatively complete; all three MD setae present; AF2 absent; P2 reduced; L1 arising immediately caudad and VOLUME 93, NUMBER 1 21 Figs. 9-14. Neurostrota gunniella, wing structure. 9, Hindwing, female, showing basal frenulum and more distal pseudofrenulum (270 um). 10, Base of female frenulum (23.1 wm). 11, Surface detail of female frenulum (1.5 wm). 12, Base of male frenulum (10 wm). 13, Surface of male frenulum (3 um). 14, Detail of Fig. 13 (1.5 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 9.) tO NO PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Piit-J\ aS Figs. 15-20. Neurostrota gunniella, wing structure. 15, Male pseudofrenulum (120 um). 16, Base of pseu- dofrenulum (17.6 um). 17, Overlapping scales of pseudofrenulum (6 um). 18, Surface detail of pseudofrenulum showing overlapping scutes (1.5 um). 19, Subanal retinaculum (SR) of ventral forewing (100 um). 20, Detail of scale (a) of Fig. 19 (2 um). (Scale lengths in parentheses: bar scale for all photographs = Fig. 15.) VOLUME 93, NUMBER 1 23 Figs. 21-26. Neurostrota gunniella, life history. 21, Leaf mines of sap-feeding larva on Mimosa pigra. 22, Frass ejected from stem burrow. 23, Larva inside tunnel within stem of M. pigra. 24, Moth larva killed by parasitic tachinid larva, E/fia n. sp. 25, Pupa inside cocoon spun on leaflets of M. pigra. 26, Cocoon spun on stem of M. pigra, Note frothy bubbles excreted by larva. 24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . reg le Orr hud i Sar Fama? i dls SSS “> ‘pie. > Figs. 27-32. Neurostrota gunniella. 27, Egg (120 wm). 28, Surface of egg chorion (23.1 um). 29, Micropyle end (67 um). 30, Detail of micropyle (15 um). 31, Detail of micropyle (17.6 um). 32, Sap-feeding larva, dorsal view of head (60 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 27.) VOLUME 93, NUMBER 1 25 Figs. 33-38. Neurostrota gunniella, sap-feeding larva. 33, Labrum and mandibles, dorsal view (21.4 um). 34, Ventral view of head (50 wm). 35, Labium and antennae, ventral view (30 um). 36, Antenna, ventral view (7.5 um). 37, Anterior view of mouthparts (27 wm). 38, Labium hypopharynx, ventral view; note aperture of vestigial spinneret (4.3 wm). (Scale lengths in parentheses; bar scale for all photographs = Fig. 33.) 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 39-44. Neurostrota gunniella, larva. 39, Sap-feeding larva, lateral view (60 um). 40, Tissue-feeding (boring) larva, anterior view of head (176 wm). 41, Detail of Fig. 40 (100 um). 42, Detail of mouthparts (60 um). 43, Labial palpi and spinneret (38 wm). 44, Maxilla (25 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 39.) VOLUME 93, NUMBER 1 27 Figs. 45-50. Neurostrota gunniella, tissue-feeding larva. 45, Labrum, ventral view (43 um). 46, Detail of Fig. 45; EP = epipharyngeal sclerite (17.6 um). 47, Lateral view of ventral side of labrum (8.6 wm). 48, Head, ventral view (176 wm). 49, Detail of mouthparts, ventral view (60 um). 50, Head, dorsal view (100 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 45.) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 51-56. Neurostrota gunniella, tissue-feeding larva. 51, Lateral view of head (176 um). 52, Detail of stemmatal region (75 um). 53, Antenna (25 um). 54, Distal view of antennal apex (13.6 wm). 55, Thorax, ventral view of T1 and 2 (231 wm). 56, Pretarsus, lateral view of T2 (15 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 51.) VOLUME 93, NUMBER 1 Figs. 57-62. Neurostrota gunniella, sap-feeding larva and cocoon. 57, Abdominal proleg, A4 (60 um). 58, Lateral view of A9-10 (200 um). 59, Ventral view of A9-10 (200 um). 60, Anal proleg showing vestigial crochet (38 wm). 61, Surface of cocoon ornamented with frothy globules secreted by larva (38 um). 62, Detail of globule (120 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 57.) 30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON = < 3 J) NY Buses I< Figs. 63-70. Neurostrota gunniella, morphology of tissue-feeding larva. 63, Chaetotaxy of pro- and meso- thorax, abdominal segments 1, 5, 7-9. 64, Head dorsal view (0.2 mm). 65, Head, ventral view. 66, Head, lateral view. 67, Abdominal segments 8-10, dorsal (0.2 mm). 68, Mandible (0.1 mm). 69, Labrum, dorsal (0.1 mm). 70, Labrum ventral; ES = epipharyngeal sclerite. (Scale lengths in parentheses.) VOLUME 93, NUMBER 1 between first and second stemmata. Six stemmata present, arranged in an oval (Figs. 52, 66). Antenna relatively short; sensilla as in Figs. 53-54. Labrum (Figs. 69, 70) with M1 absent; three pairs of epipharyngeal spines present, the lateral spine the most reduced; epipharyngeal sclerite (Figs. 45- 47) thin, ovoid with entire margin, and raised slightly above epipharyngeal mem- brane. Mandible (Fig. 68) with three large medial cusps and three smaller, marginal cusps. Maxilla as in Figs. 42, 44. Spinneret (Figs. 43, 49) tubular, moderately elongate. Labial palpus with a relatively long, basal segment bearing one short sensillum and a minute apical segment bearing a larger sen- sillum. Thorax: Pronotal plate variable, light to dark brown. L group bisetose on T1, trise- tose on T2-3 with L2 and 3 minute. SV bisetose on T1, unisetose on T2-3. Legs (Fig. 55) well developed, with coxal plates well separated; pretarsal claw (Fig. 56) with large, triangular, axillary spine. Abdomen: Dorsal and subdorsal plates of A8-10 dark brown. Al-2, 8-9 with eight pairs of primary setae; L2 absent. A3-5 with 10 pairs of primary setae; SV series trise- tose. Crochets (Fig. 57) arranged in two scat- tered groups on A3-S, an anterior row of 2-3 spines and a posterior concentration of ca. 20 spines (Fig. 57); prolegs and crochets absent on A6. SV series bisetose on A7; L1 together with SD1-2 on spiracular plate; L series trisetose on A9; SD1 with D1-2 on dorsal plate. Anal plate (A10) with four pairs of setae (Figs. 58, 67); anal prolegs (Figs. 59, 60) nearly devoid of crochets, each proleg partially subdivided into two rounded tu- bercules with 1—2 small spines on each tu- bercule (Fig. 60). Pupa (Figs. 71-82).—Length of largest pupa 5.3 mm; maximum width 1.0 mm. Vertex with a small, conical, multipointed apical process (cocoon cutter). Two pairs of small, nearly contiguous setae from lower frons near dorsal margin of labrum. Anten- na long and straight, extending almost a 31 fourth its length beyond abdomen. Wing sheath extending to caudal margin of AS. Hindleg slightly surpassing caudal apex of abdomen. Chaetotaxy as in Fig. 72; all setae extremely short. Dorsum of A2—6 with dense concentration of small, stout spines. Venter of A9 with a transverse row of four small stout spines; A10 with a pair of similar spines dorsally and two spines laterally (Figs. 80- 82). Cocoon (Figs. 25, 26, 61, 62).—Consist- ing of a firm white sheet of silk over some crevice in the stem or among the leaf pin- nules, ornamented externally by 30-40 minute frothy balls approximately 0.3 mm in diameter. Type.— Lectotype, ° (present designation, Davis) USNM. Type locality.— Brownsville, Texas. Material examined.—COSTA RICA: Guanacaste: Tempisque: | 4, 18 Jan, slides USNM 29938, 29939. CUBA: Santiago de Cuba: Santiago de Vegas [Cuba]: 2 4, 1 8, 22-23 Oct, ex Mimosa asperata, slide USNM 29936. MEXICO: Guerrero: Aca- pulco: 4 4, 3 2?, em. 10-25 Nov, DRD 554.1, ex Mimosa pigra; 2 8, 5 2, em. 12 May-23 Jun, ex Mimosa pigra; | 6, em. 6 Jun, ex Mimosa sp.; ca. 15 larvae, 8 Feb. ex Mimosa pigra; ca. 40 larvae, 28 Oct-1 Nov, ex MI- mosa pigra. Barra Vieja, Acapulco: | 2, em. 19 Jul, ex Mimosa pigra. Puerto Marquez, Acapulco: 6 4, 7 2, em. 21 Apr—30 Jun, ex Mimosa pigra, slides USMN 28687, 28804; 2 6,em. 25 Jun, ex Neptunia plena. Tabasco: Villahermosa: 1 2, 18 Mar, ex Mimosa pi- gra. Veracruz: Catemaco: | 2, 16 Mar., ex Mimosa pigra, slide USNM 29933. Vera- cruz: 1 9, 14 Mar, ex Mimosa pigra. UNIT- ED STATES: Texas: Cameron Co:: Brownsville: 1 @ (lectotype), Jun, slide USNM 29931; 11 2 (paralectotypes), Jun. San Benito: 6 2, 24-31 Jul; 2 2, Aug; 2 2, 8- 15 Sep. Southmost: 1 6, 27 Oct. Fort Bend Co: Brazos Bend State Park: 1 9, 3 Aug. Harris Co: Houston: 1 4, 13 Jul, slide USNM 28440. All specimens in USNM. Distribution.— Widespread in subtropi- 32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 71,72. Neurostrota gunniella, pupa. 71, Ven- tral view (0.5 mm). 72, Lateral view. (Scale length in parenthesis). cal or tropical, moderately wet to semi-arid habitats wherever the primary host, Mi- mosa pigra occurs; from southern Texas to Costa Rica and Cuba. Hosts.— Leguminosae: Mimosa pigra L. and Neptunia plena Benth.; for secondary or potential hosts see Table 3. BIOLOGY Rearing methods.—Stems of M. pigra in- fested with larvae of N. gunniella were col- lected at Barra Vieja and Pie de la Cuesta in the state of Guerrero, Mexico and shipped by air freight to Australia on 19th Septem- ber 1986 and 5th December 1986, respec- tively. On arrival larvae were removed from these stems, transferred to potted plants of M. pigra and held in the quarantine facility at CSIRO, Long Pocket Laboratories, Bris- bane. From this stock, a colony of N. gun- niella was established on potted plants of M. pigra in a quarantine insectary air con- ditioned to 26 + 1°C for 14 hours/day and 21 + 1°C for the remainder of the day. Rel- ative humidity was maintained above 55%. Overhead fluorescent lighting supplement- ed oblique natural lighting. Newly emerged adults were fed a solution of ascorbic acid, (2 gms), honey (15 mls) and water (180 ml). Five pairs of fed adults were placed in a cage 45 x 45 x 90 cm high, with 4-8 potted plants of M. pigra. After the eggs laid on the plants had hatched, the larvae were provided with fresh plants of M. pigra as required, until development was completed. Biological studies.—Studies were carried out in “Conviron” constant temperature cabinets at 25 + 0.5°C with a 14:10 pho- toperiod. The number of eggs/female/day (fecundity) was measured and preferred ovi- position sites determined by counting the number of eggs laid on different parts of the plant. Egg incubation time and the size of eggs were measured. The number and du- ration of each larval instar was recorded by examining larvae every day and measuring the width of their head capsules. Duration of pupal stage and adult longevity were also measured. Eggs were usually deposited singly on the ventral side of mature leaves growing at ei- ther the first or second node from the tip. The most favoured site being on the sec- ondary rachis adjacent to a leaf hair. Eggs hatched after 4-5 days. First and sec- ond instar larvae mined pinnules (Fig. 21), with as many as 5 being mined by each larva. Third instar larvae usually entered the primary rachis where a pinna was at- tached and tunnelled towards the stem where they fed either in the petiole bulb or in the VOLUME 93, NUMBER 1 Figs. 73-78. Neurostrota gunniella, pupa. 73, Head, dorsal view (231 um). 74, Head, ventral view (231 um). 75, Detail of frons (86 um). 76, Detail of frontal process (cocoon cutter) (30 um). 77, Head, lateral view (176 um). 78, Dorsal view of A2 (176 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 73.) 34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SA ee ’ WAP Figs. 79-82. Neurostrota gunniella, pupa. 79, Dorsal spines of A3 (27 wm). 80, Caudal view of A10 (86 um). 81, Lateral view of A9-10 (100 um). 82, Ventral view of A9-10 (86 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 79.) stem (Fig. 23). Sometimes they exited and re-entered the stem at another node or near a prickle. Frass was often visible where the larvae entered the stem (Fig. 22). There were eight larval instars. The final instar con- sisted of three distinct phases based on be- havioral and/or color changes. The first phase was the typical, tissue-feeding form; in the second phase the larva usually exited from the stem, spun a cocoon, and changed from a dark, blue-green color back to the cream color of the earliest instars. In the final phase, herein referred to as the prepupa (Fig. 83, Table 1), the larva was inactive for ca. 24 hours and then pupated. If disturbed, the prepupa moved in a spiralling motion similar to that of the pupa. Most mature larvae exited from the stem and pupated in a slender cocoon usually spun between pinnules (Fig. 25), or in crevices on or within the stem (Fig. 26). The outside of the cocoon was ornamented with small, pearly-white, frothy balls (Figs. 26, 61, 62) discharged from the anus of mature larvae and attached by projecting their posterior through slits in the cocoon. These slits were then covered with additional silk and the larvae entered the prepupal stage. Adults emerged 7 days after pupation. The dura- tion of life-cycle stages of N. gunniella and VOLUME 93, NUMBER 1 eee ee 35 Adult emergence aE) eee Prepupa So fe. CViIII' Insta eee CVI Insta = VI Instar a i. V Instar pk. IV Instar | a Ill Instar | Il Instar Ls | Instar O 5 10 15 20 25 30 35 40 STAGE DURATION ( DAYS ) Fig. 83. Duration of life-cycle stages of Neurostrota gunniella and the proportional distribution of the stages for each day. The histograms show the proportion of each stage and adult emergence present on each day. the proportional distribution of stages for each day of development is shown in Fig. 83. Adults when resting had their antennae lying back along the body and extending beyond the wings; the body was usually held so that the head was slightly depressed; the fore- and mid-legs were held together to the end of the tibia but then separated at an angle of ca. 45°, and the hind legs were held back along the body before being slightly splayed. In addition to the typical frenular differences, males could be distinguished from females by differences in the appear- ance of scales arising from the terminal seg- ments of the abdomen. In females the scales formed a cylinder that enveloped the ovi- positor whereas in males the scales overlaid the valvae. The ratio of reared males to fe- males was 1:1. Adults fed on liquid nutrient solution. The age-specific fecundity and sur- vival of adult females is shown in Fig. 84. The duration, number and development time of the stages of N. gunniella at 25 + 0.5°C is summarized in Table | and Fig. 83. Host specificity tests. —In Mexico, stems of leguminous plants growing adjacent to M. pigra were dissected and examined for N. gunniella. These plants included seven Mimosa spp., two Acacia spp., Schrankia distachya DC., Prosopis juliflora (Swartz) DC., Parkinsonia aculeata L., Leucaena leucocephala (Lam.) de Wit, and Neptunia plena. N. gunniella was not encountered in any of these plants except N. plena. One potted plant of Mimosa sp. (not M. pigra), growing in the laboratory grounds was found infested with larvae of N. gunniella. Larvae in N. plena and Mimosa sp. completed de- velopment. 36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 20 MEAN EGGS / FEMALE / DAY 1 | . 1 ete 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 10 on NO. FEMALES SURVIVING 17 18 19 AGE OF FEMALES ( DAYS ) Fig. 84. Age-specific fecundity and survival of Neurostrota gunniella females (n = 20). In Australia, host specificity was deter- mined by observing the oviposition of adult females and feeding by first instar larvae. These behavioral characteristics of N. gun- niella were studied using 100 species of plants representing 18 plant families and included 69 species of Leguminosae (Table 2). These are the same species used in stud- ies with other insects already released for the biological control of M. pigra in Aus- Table 1. Duration of development (days) of Neu- rostrota gunniella on Mimosa pigra at 25 + 0.5°C (n = 20). Stage XY Range Egg (incubation) 4.0 4-5 Larvae Ist instar 2.0 a 2nd instar 230) = 3rd instar DES 2-3 4th instar 2:3 2-3 5th instar 2.8 2-4 6th instar Sal 2-4 7th instar 3.4 2-4 8th instar 33 2-7 Prepupa 1.0 _ Pupa 7.0 = Adult longevity 15.6 11-19 Total life cycle 8322 28-40 Number of eggs/female 85.6 49-128 tralia (Kassulke et al. 1990) with the addi- tion of four Australian species of Neptunia and one of Adenanthera, Archidendron and Pararchidendron. Species of Neptunia were added because N. gunniella had been reared occasionally from N. plena in Mexico (rec- ords of CSIRO exploratory unit, Mexico). The other three genera were added to in- crease the range of woody Australian native legumes tested. Larval feeding. — Excised pinnae of M. pi- gra each with at least ten eggs due to hatch the next day, were placed on the first mature leaf of each test plant and on a plant of M. pigra. Those species on which larvae fed were held until larvae either died or com- pleted development. This procedure was re- peated until all plant species had been tested at least 3 times. In these tests, neonate lar- vae fed and completed development on 6 species: M. pigra, M. pudica, Neptunia di- morphantha Domin, N. gracilis Benth., N. major (Benth.) Windler and N. monosper- ma F. Muell. In one trial with Acacia po- dalyriifolia A. Cunn. ex. G. Don and an- other with Glycine max (L.) Merr., one larva survived and developed to the 3rd instar before dying. On another 37 plant species, larvae fed briefly in some trials before dying while still in the first instar (Table 3). VOLUME 93, NUMBER 1 Table 2. Plants tested to determine host specificity of Neurostrota gunniella. Family Genus/Species Common Name 1. Taxonomically unrelated plants Anacardiaceae Annonaceae Bignoniaceae Caricaceae Compositae Cruciferae Lauraceae Malvaceae Myrtaceae Poaceae Proteaceae Rosaceae Rubiaceae Rutaceae Simmondsiaceae Solanaceae Theaceae i) Leguminosae Mangifera indica Annona reticulata Jacaranda mimosifolia Carica papaya Helianthus annuus Lactuca sativa Brassica oleracea var. botrytis Persea americana Gossypium hirsutum Eucalyptus maculata E. miniata E. tereticornis E. tetrodonta Leptospermum longifolium Psidium guajava Syzygium luehmannii Saccharum officinarum Sorghum vulgare Zea mays Macadamia ternifolia Fragaria x ananassa Malus sylvestris Prunus persica Rosa sp. Coffea arabica Citrus limon C. reticulata C. sinensis Simmondsia chinensis Lycopersicon esculentum Camellia sinensis . Taxonomically Related Plants Acacia aulacocarpa . auriculiformis . cincinnata . crassicarpa deanii x decurrens . difficilis . dimidiata . holosericea . latescens . leptocarpa . Mangium . oncinocarpa . podalyriifolia . spectabilis Adenanthera pavonina Albizia lebbeck Arachis hypogaea Archidendron hendersonii BRS JSS ISPS ISSR SRR fe SUI mango custard apple jacaranda pawpaw sunflower lettuce cauliflower avocado cotton spotted gum forest red gum Darwin stringy bark tea tree guava small leaf water-gum sugar cane sorghum maize Queensland nut strawberry apple peach rose coffee lemon mandarin orange jojoba tomato tea hickory wattle ferny wattle war-roon silver wattle Pilliga wattle red sandalwood woman’s tongue peanut Table 2. Continued. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Genus/Species Bauhinia galpini B. variegata Caesalpinia ferrea Cajanus cajan Calopogonium mucunoides Cassia alata C. artemisioides C. fistula C. mimosoides Cathormion umbellatum Centrosema pubescens Delonix regia Desmanthus virgatus Desmodium tortuosum Dichrostachys cinerea D. spicata Enterolobium contortisiliquum Erythrophleum chlorostachys Glycine max Lablab purpureus Leucaena diversifolia L. lanceolata L. leucocephala L. macrophylla L. pallida L. shannonii Lysiphyllum hookeri Macroptilium atropurpureum Medicago sativa Mimosa invisa M. pigra M. pudica Neptunia dimorphantha N. gracilis N. major N. monosperma Pararchidendron pruinosum Peltophorum pterocarpum Piliostigma malabaricum Pisum sativum Pongamia pinnata Pueraria phaseoloides Samanea saman Sesbania formosa Stylosanthes hamata cv. Verano Tamarindus indica Vicia faba Vigna mungo V. radiata V. unguiculata Common Name orchid tree leopard tree pigeon pea calopo ringworm cassia cassia golden shower centro poinciana desmodium Cooktown ironwood soybean lablab bean leucaena white bauhinia siratro lucerne giant sensitive plant mimosa, giant sensitive plant common sensitive plant native sensitive plant pea pongamia puero raintree stylo tamarind broadbean mung bean mung bean cowpea VOLUME 93, NUMBER 1 Table 3. Larval feeding, oviposition, and development of Neurostrota gunniella. Larval Trials Adult Trials No. with No. with No. Trials Feeding Stage Reached No. Trials Eggs Stage Reached Acacia crassicarpa 3 3 lst Instar 3 0) — A. leptocarpa 4 3 Ist Instar 4 0 _ A. mangium 4 4 Ist Instar 5 ] Ist Instar A. oncinocarpa 3 1 Ist Instar 3 0 a A. podalyriifolia 3 2 Ist Instar* 10 1 Ist Instar Adenanthera pavonina 3 2 Ist Instar 3 (0) — Albizia lebbeck 3 l Ist Instar 4 0 ~ Arachis hypogaea 3 3 Ist Instar 4 0 _ Brassica oleracea 3 2 Ist Instar 3 0 — Cajanus cajan 3 Ist Instar 3 0 — Cassia alata 3 2 Ist Instar 4 0 _ C. artemisioides 3 1 Ist Instar 4 0 — C. fistula 3 1 Ist Instar 3 0 _ C. mimosoides 3 1 Ist Instar 4 0 — Delonix regia 3 2 Ist Instar 3 0) a Desmanthus virgatus 3 i lst Instar 4 1 Ist Instar Desmodium tortuosum 3 2 Ist Instar 3 0 — Glycine max 3 3 Ist Instar* 4 0 ~ Gossypium hirsutum 3 2 Ist Instar 3 0 _ Helianthus annuus 3 | Ist Instar 3 0 = Lablab purpureus 4 1 Ist Instar 4 0 _ Lactuca sativa 3 l Ist Instar 3 0 _ Leucaena diversifolia 3 1 Ist Instar 3) 0 — L. lanceolata 3 l Ist Instar 3 0 _ L. leucocephala 3 l Ist Instar 4 0 - L. macrophylla 3 1 Ist Instar 3 0 _ L. shannonii 4 4 Ist Instar 6 1 Ist Instar Lycopersicon esculentum 3 1 Ist Instar 4 0 = Macroptilium atropurpureum 3 2 lst Instar 4 0 _— Malus sylvestris 4 3 Ist Instar 3 0 _ Medicago sativa 3 1 Ist Instar 4 0 — Mimosa pigra 5) 5 Adult 5 5 Adult M. pudica 3 1 Adult 4 4 Adult Neptunia dimorphantha 3 2 Adult 3 3 Adult N. gracilis 3 l Adult 4 3 Adult N. major 3 3 Adult 3 3 Adult N. monosperma 3 ” Adult 5) 3 Adult Piliostigma malabaricum 3 2 Ist Instar 3 0 — Pongamia pinnata 3 l Ist Instar 3 0) _ Prunus persica 3 1 lst Instar 3 0 = Samanea saman 3 1 Ist Instar 4 0 Sorghum vulgare 3 1 Ist Instar 3 0 ~ Tamarindus indica 3 1 Ist Instar 4 0 Vigna mungo 3 2 Ist Instar = 0 — V. unguiculata 3 3 Ist Instar 4 0 — * | larva reached 3rd Instar. 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 4. Development of Neurostrota gunniella on five test plants and number of adults emerged in F1 generation from five gravid females in no choice tests. Development (Days) Egg to Adult Plant Species 55 Range Mimosa pigra 36 31-48 M. pudica 42 37-46 Neptunia dimorphantha 35 31-48 N. gracilis 40 33-37 N. major 48 34-63 N. monosperma 35 31-47 % Larval Adults Emerged F, Generation Mortality 36 Range <25 95 52-132 >70 18 1-51 >85 22 5-38 >90 3 0-6 >96 5 2-7 >70 10 0-39 Oviposition.— Plant species which were fed on by larvae in feeding tests were further tested for selection by females for oviposi- tion in the absence of M. pigra. These no choice tests were repeated at least three times. Five pairs of newly emerged adults were fed nutrient solution and placed in a cage with a potted test plant for five days. A fine mist of water was sprayed into the cage each morning and afternoon for adults to drink. Plants were examined for the pres- ence of eggs and those with eggs were ex- amined daily for larval feeding. Observa- tions continued until larvae either died or completed development. Species of Acacia which have bipinnate leaves in the juvenile stage were tested with and without pinnules. To avoid any possible disruption of behav- ior no M. pigra was present in the laboratory when these tests were carried out. Eggs were laid and larvae completed de- velopment on the same 6 plant species as in the larval feeding tests, i.e. M. pigra, M. pudica, N. dimorphantha, N. gracilis, N. major and N. monosperma. Eggs were also laid on Acacia mangium Willd. in one of five trials, on 4. podalyriifolia in one of ten trials, on Desmanthus virgatus (L.) Willd. in one of four trials and on Leucaena shan- nonii Donn. Smith in one of six trials, but in all cases larvae died in the first instar after mining in a few pinnules (Table 3). Fecundity and development.—Oviposi- tion averaged 86 eggs per female on M. pigra (n = 20). Oviposition could commence the first night after emergence but most eggs were laid during the second night. Eggs were difficult to see and accurate egg counts were not made on ™. pudica, N. dimorphantha, N. gracilis, N. major and N. monosperma. The number of larval mines that developed following egg-laying by five gravid females on each of these species varied from an av- erage of 128 on M. pigra, to 26 on M. pu- dica, 37 on Neptunia dimorphantha, 25 on N. gracilis, 60 on N. major and 7 on N. monosperma (n = 3). On these species, there was little difference in the duration of larval development (Table 4). Adults obtained from larvae that developed on these plants were returned to plants of the same species to determine if successive generations could develop. Successive generations were able to develop on these six species but larval mortality was greater than 96% on N. major, 90% on N. gracilis, 85% on N. dimorphan- tha, and 70% on M. pudica and N. mono- sperma. Mortality on M. pigra was less than 25% (Table 4). Larvae caused no significant damage to the Neptunia spp. whereas they always caused heavy damage to M. pigra and occasionally moderate damage to M. pudica. OBSERVATIONS IN MEXICO AND CENTRAL AMERICA In Mexico and Central America, M. pigra has two varieties, M. pigra var. pigra L. and M. pigra var. berlandieri (Gray. ex Torr.) B. L. Turner (Turner, 1959). Both varieties are attacked by N. gunniella, and in Mexico and Costa Rica all stands of M. pigra examined VOLUME 93, NUMBER 1 Table 5. Parasitoids associated with Neurostrota gunniella in Mexico. Diptera Chloropidae Fiebrigella sp. Fiebrigella new sp. nr. catalpae (Malloch) Tachinidae Elfia new sp. undescribed Hymenoptera Perilampidae Perilampus sp. Braconidae Apanteles sp. Bracon sp. Hypomicrogastar sp. Orgilus sp., probably new sp. Phanomeris sp. Chalcididae Brachymeria sp. Eulophidae Elasmus sp. Horismenus sp. were infested with the moth. Typically a 20 cm piece of stem would be infested by 5 larvae. PARASITIZATION OF N. GUNNIELLA In Mexico, field collected stems of M. pi- gra infested with larvae of N. gunniella were caged in the laboratory for emergence of parasitoids (Table 5). Many more para- sitoids than adult moths emerged, with E/fia n. sp. (Tachinidae) being by far the most numerous. Figure 24 shows N. gunniella be- ing attacked by a larva of E/fia n. sp. The chloropid, Fiebrigella sp., emerged only in- frequently and may not represent a parasitic relationship. Sabrosky (1950) suggested that the rather random appearance of this spe- cies in such rearings may indicate they are scavengers. DISCUSSION Morphology and systematics. — The stem- boring habit of the larva was first reported by Busck (1934), based upon rearings in Cuba by H. R. Otero. Busck also described a second species of Neurostrota (pithecolo- 41 biella) from Cuba, a stem borer in the le- guminous tree Pithecellobium saman (Jacq.) Benth. However, on the basis of the male genitalia, pithecolobiella does not appear congeneric with the type of the genus, N. gunniella. A second, undescribed species of Neurostrota has been collected at light near Tamazunchale, Mexico by the senior author (Davis). Another species, similar to the lat- ter, is also known to bore in stems of Inga vera Willd., (Leguminosae) in Puerto Rico, thus bringing the number of known species of Neurostrota to three. Although no further published reports have been found on stem- boring Gracillariidae, other than the above references by Busck and the gall-making or shoot-mining habits of a few other Gracil- lariidae, it appears from collecting records that a true stem-boring habit has arisen per- haps several times within tropical/subtrop- ical Gracillariidae. One unusual morphological feature pres- ent in the adults of N. gunniella, is the de- velopment of a supplementary wing cou- pling mechanism in both sexes. In addition to the frenular bristles (single in the male and double in the female), these moths pos- sess a prominent pseudofrenulum arising from the costal margin of the hindwing near the termination of Sc. The pseudofrenulum couples with a row of stiff, specialized scales located along the ventral hind margin in the anal area of the forewing. Because of its lo- cation, it is proposed that this accessory ret- inaculum be termed the subanal retinacu- lum, in comparison to the subcostal and subdorsal retinaculum described by Till- yard (1918) and Braun (1924). The longi- tudinal scale ridges of the subanal scales in Neurostrota are strengthened by the fusion of all scutes (Fig. 20), similar to the sub- dorsal retinacular scales described for Opo- stegidae (Davis, 1989). In N. gunniella, the pseudofrenulum is composed of three to four, tightly ap- pressed, piliform scales (Figs. 15, 16), usu- ally accompanied by four or five much shorter, broader costal scales which overlap 42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON more basally. Although often appearing to be fused (as in the case of the male frenu- lum), the piliform scales are separated throughout their length and closely overlap each other (Fig. 17). In contrast to the more specialized, cylindrical bristles of the male and female frenulum, the longitudinal ridg- es of the pseudofrenulum are not entire but instead are subdivided into overlapping scutes (Fig. 18). Busck (1934) and others have noted the presence of similar pseudofrenula in a few other gracillariid genera (Micrurapteryx, Neurobathra, and Parectopa), and Kumata (in litt.) has observed their presence in Can- opomorpha, Cuphodes, Epicephala, Liocro- byla, Stomphastis, and a few Acrocercops. A similar pseudofrenulum has been illus- trated in Heliozelidae (Common 1970, Fig. 36.15B). Less specialized, pseudofrenular scales have also been noted in several genera of Cosmopterigidae (Hodges 1978, Figs. 1- 3). Within this family and according to ge- nus, a graduated series varying from a rel- atively long row of costal scales to a more concentrated group of a few piliform scales can be present. The ventral surface of the labrum in the tissue-feeding instars of N. gunniella bears the typical lepidopterous complement of three pairs of spinose, epipharyngeal setae. Also present is a pair of flat, oval, platelike structures (Figs. 45-46) which may be ho- mologus to the epipharyngeal sclerites de- scribed recently by Leidy and Neunzig (1989). These sclerites are more distinct and laterad in N. gunniella than in the species of Dioryctria figured by Leidy and Neunzig. Like the associated setae, the epipharyngeal sclerites may occur throughout most fami- lies of Lepidoptera and have been observed by the senior author in several genera of Psychidae and Tineidae. In all species of Tineinae studied thus far, the outer margins of the epipharyngeal plates are serrated, lat- eral, and typically project beyond the edge of the labrum. Host specificity.—A particularly large number of plants were studied including representatives of all taxa closely related to M. pigra. Although newly-emerged larvae fed slightly on a number of plants, the larvae died before completing the first instar. The only established Australian plants which supported development of N. gunniella were M. pigra, M. pudicaand four species of Nep- tunia, N. dimorphantha, N. gracilis, N. ma- jor and N. monosperma. Larval mortality was lower and plant damage was much more severe on M. pigra than on any of these plants. Any damage to the weed M. pudica would be regarded as beneficial in Australia. In Mexico, N. gunniella was observed to attack N. plena occasionally, but only when this plant was growing in a thicket of M. pigra (records of CSIRO Exploratory Unit). Ex- tensive testing of the four Neptunia species endemic to Australia, N. dimorphantha, N. gracilis, N. major and N. monosperma, showed that they may occasionally act as hosts for N. gunniella if it was to become established in Australia for biological con- trol of M. pigra. However as larval mortal- ity was very high, N. gunniella is unlikely to have any significant affect on these Nep- tunia species or to persist on them. Fur- thermore, the range of M. pigra in Australia does not overlap to any significant extent, that of these plants. N. dimorphantha and N. monosperma are reported from open grasslands in northern and eastern Australia where M. pigra does not occur. N. major occurs along rivers in northern Australia (Windler 1966) but is uncommon in areas infested with M. pigra. N. gracilis is a rare plant on floodplains in the Northern Ter- ritory (C. G. Wilson, pers. comm.) and is more common in open grasslands of Queensland and northern New South Wales. In Mexico the full impact of N. gunniella on the growth of M. pigra plants was limited by a suite of larval parasitoids. Away from these natural enemies the potential of N. gunniella as a biological control agent is es- VOLUME 93, NUMBER 1 timated to be high. The first release of this insect (100 adults and 1200 eggs) was made at the lower Adelaide River flood plain, 69 km east southeast of Darwin, Northern Ter- ritory, Australia on February 2, 1989. Sub- sequently Napompeth (pers. comm.) re- ported that in quarantine studies in Thailand, N. gunniella fed on Neptunia oleraceae Lour., an important aquatic veg- etable, and was not approved for release against M. pigra. ACKNOWLEDGMENTS We thank Vichai Malikul and Young Sohn, Department of Entomology, Smith- sonian Institution for the line drawings and Susann Braden and Brian Kahn, Smithson- ian SEM Lab, and Victor Krantz Smithson- ian Photographic Laboratory for photo- graphic assistance. The final draft of the manuscript was prepared by Silver West. Michael Day, Glynn Maynard and Hugo Miranda provided valuable technical assis- tance. Parasitoids were kindly identified by E. E. Grissell, P. M. Marsh, C. W. Sabrosky, M. E. Schauff and N. E. Woodley, research entomologists of the Systematic Entomol- ogy Laboratory, BBII, USDA. Gail Rob- ertson and Lyall Skilling assisted with il- lustrations. This study was supported financially by the Australian Centre for In- ternational Agricultural Research and the Northern Territory Department of Primary Industry and Fisheries. LITERATURE CITED Barnes, W. and J. McDunnough. 1917. Check List of the Lepidoptera of Boreal America. Herald Press, Decatur, Ill. viii + 392 pp. Braun, A. F. 1924. The frenulum and its retinaculum in the Lepidoptera. Annals of the Entomological Society of America 18(3):L 234-256, pl. 23. Burkart, A. 1948. Las especies de Mimosa de la flora Argentina. Darwiniana 8(1): 9-231. Busck, A. 1906. Tineid moths from southern Texas, with descriptions of new species. Proceedings U.S. National Museum 30: 721-736. 1934. Microlepidoptera of Cuba. Entomo- logica Americana 13(4): 151-217. Common, I. F. B. 1970. Lepidoptera, pp. 765-866. 43 In CSIRO, The Insects of Australia. Melbourne University Press, Carlton, Victoria. Davis, D. R. 1983. Gracillariidae, p. 9. In R. W. Hodges et al., eds. Check List of the Lepidoptera of America North of Mexico. E. W. Classey Ltd. and the Wedge Entomological Research Founda- tion, London. 1984. Gracillariidae, p. 26. Jn J. Heppner, ed. Atlas of Neotropical Lepidoptera. Checklist: Part | (Micropterigoidea-Immoidea). Dr. W. Junk Publishers, The Hague, Boston, Lancaster. 1989. Generic revision of the Opostegidae, with a synoptic catalog of the world’s species (Lep- idoptera: Nepticuloidea). Smithsonian Contribu- tions to Zoology, No. 487. 97 pp., 320 figs., 1 map. Ely, C. R. 1918. A revision of the North American Gracilariidae [sic] from the standpoint of vena- tion. Proceedings of the Entomological Society of Washington 19(1-4): 29-77. Hodges, R. W. 1978. Gelechioidea, Cosmopterigi- dae. Fasc. 6.1, 166 + x pp., pls. 1-6. In R. B. Dominick et al. The Moths of North America. E. W. Classey Ltd. & The Wedge Entomological Re- search Foundation, London. Kassulke, R. C., K. L. S. Harley, and G. V. Maynard. 1990. Host specificity of Acanthoscelides quad- ridentatus and A. puniceus (Col.: Bruchidae) for biological control of Mimosa pigra (with prelim- inary data on their biology). Entomophaga 35: 85- 96. Kleinschmidt, H. E. and R. W. Johnson. 1977. Weeds of Queensland. Queensland Department of Pri- mary Industries. 188 p. Leidy, N. A. and H. H. Neunzig. 1989. Taxonomic study of the larvae of six eastern North American Dioryctria (Lepidoptera: Pyralidae: Phycitinae). Proceedings of the Entomological Society of Washington 91(3): 325-341. Lonsdale, W. M. and R. Segura. 1987. A demograph- ic study of native and introduced populations of Mimosa pigra. Proceedings of the Eighth Austra- lian Weeds Conference 1987, pp. 163-166. In Lemerle, D. and A. R. Leys, eds. Weed Society of New South Wales, Haymarket. McDunnough, J. 1939. Checklist of the Lepidoptera of Canada and the United States of America. Part 2, Microlepidoptera. Memoirs of the Southern California Academy of Sciences 2(1): 1-171. Meyrick, E. 1912a. Genera Insectorum, Lepidoptera Heterocera (Tineae) Fam. Gracilariadae [sic], Fasc. 128, p. 1-36, 1 col. pl. M. Wytsman, Tervuren, Belgium. 1912b. Lepidopterorum Catalogus. Pars 6, Adelidae, Micropterigidae, Gracilariadae [sic], p. 25-68. W. Junk, Berlin. Miller, I. L. 1982. The distribution and threat of Mimosa pigra in Australia, pp. 38-50. In G. La- 44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mar Robert and D. H. Habeck, eds. Proceedings of the International Symposium on Mimosa pigra Management, Chiang Mai. Miller, I. L.and W. M. Lonsdale. 1987. Early records of Mimosa pigra in the Northern Territory. Plant Protection Quarterly 2: 140-142. Miller, I. L., L. Nemestothy, and S. E. Pickering. 1981. Mimosa pigra in the Northern Territory. Tech- nical Bulletin No. 51, Department of Primary Pro- duction, Northern Territory. Napompeth, B. 1982. Background, threat, and dis- tribution of Mimosa pigra L. in Thailand, pp. 15—- 26. In G. Lamar Robert, D. H. Habeck, eds. Pro- ceedings of the International Symposium on Mi- mosa pigra Management, Chiang Mai. Pitt, J. L. and I. L. Miller. 1989. A review of survey techniques for the detection of weeds with partic- ular reference to Mimosa pigra L. in Australia and Thailand. Plant Protection Quarterly 3: 149-155. Sabrosky, C. W. 1950. A synopsis of the chloropid genera Chaetochlorops and Eugaurax (Diptera). Journal of the Washington Academy of Sciences 40(6): 183-188. Tillyard, R. J. 1918. The Panorpoid complex. Part 1. The wing-coupling apparatus, with special ref- erence to the Lepidoptera. Proceedings of the Lin- naean Society of New South Wales 53:286-319. Turner, B. L. 1959. The Legumes of Texas. Univer- sity of Texas Press, Austin and London. 284 pp. Vari, L. 1961. South African Lepidoptera, Vol. 1, Lithocolletidae. Transvaal Museum, Memoir No. 125 x1x 238) pp! 2 apls: Windier, D. R. 1966. A revision of the genus Nep- tunia (Leguminosae). Australian Journal of Bot- any 14: 379-420. Now Available A Handbook of Families of Nearctic Chalcidoidea by E. Eric Grissell and Michael E. Schuaff Morphology Biology Distribution Collecting * & & & Family Systematics 20 families and their major subfamilies Distinguishing Characters Pictorial key with over 120 illustrations Systematic Overview Literature Resources Biological Overview Collecting and Mounting e Spiral Bound for easy use; 86 pp. Price — $10.00 includes postage, prepaid orders only. Published by and for sale from: Entomological Society of Washington c/o Department of Entomology NHB-168, Smithsonian Institution Washington, D.C. 20560 PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 45-50 ANTHRIBUS NEBULOSUS, A EURASIAN SCALE PREDATOR IN THE EASTERN UNITED STATES (COLEOPTERA: ANTHRIBIDAE): NOTES ON BIOLOGY, RECOGNITION, AND ESTABLISHMENT E. RICHARD HOEBEKE AND A. G. WHEELER, JR. (ERH) Department of Entomology, Cornell University, Ithaca, New York 14853; (AGW) Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Penn- sylvania 17110. Abstract. —The Eurasian Anthribus nebulosus Forster is a scale predator that was intro- duced from Europe and released in Virginia in the late 1970s for potential control of pest species. New records of this natural enemy of soft scales are given for Connecticut, Massachusetts, and New York, where it was collected on spruce (Picea spp.) infested with the bud scale Physokermes hemicryphus (Dalman). It is suggested that this anthribid is adventive in the northeastern states and that populations were established before its intentional introduction into Virginia. The Old World distribution and habits are sum- marized, and characters facilitating its recognition in the Nearctic fauna are provided. Key Words: Insecta, biological control, Coccoidea, Physokermes, immigrant insects During detection surveys for immigrant arthropods conducted in the northeastern United States, we discovered Anthribus nebulosus Forster, a Eurasian scale preda- tor. On 29 May 1989, numerous adults were beaten from the lower branches of a Colo- rado blue spruce (Picea pungens Engelm.) growing adjacent to the Wesleyan Univer- sity campus, Middletown, Connecticut. The tree was infested with the introduced bud scale Physokermes hemicryphus (Dalman). Although long identified as the European P. piceae Schrank, the Old World bud scale introduced into North America has been shown to be P. hemicryphus (Williams and Kosztarab 1972, Gill 1988). Additional an- thribids were found in mid-July from west- ern Connecticut and Massachusetts to east- ern New York. We consider 4. nebulosus to be adventive in the northeastern United States, even though this predator had been intentionally introduced from Europe into Virginia (Kosztarab and Kozar 1983). Adults were collected in Hungary in 1975 and 1977, reared and evaluated for their biocontrol potential against various scale insects, and released on the Virginia Tech campus at Blacksburg in fall 1978 and 1979. These beetles were placed on Norway spruce (P. abies (L.) Karst.) infested with P. hemicry- phus. Monitoring of these trees indicated establishment of A. nebulosus by 1979, and a scale predation rate of about 30% by Au- gust 1981. Surveys in the surrounding area revealed a small anthribid population on Norway spruce about | km east of the re- lease site during fall 1981 and spring 1982; in fall 1982, a few adults were collected on Norway spruce at Radford, about 13 km southwest of Blacksburg. Field releases of the predator (300 adults) at Virginia Beach in fall 1981 apparently did not result in es- tablishment (Kosztarab and Kozar 1983). Here, we review the Old World distri- bution of A. nebulosus, and provide new 46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON igen: in 1989. Ml, denotes known release sites in Virginia. North American records. Scanning electron photomicrographs are included for salient morphological characters of 4A. nebulosus and Trigonorhinus spp., indigenous an- thribids that could be mistaken for A. ne- bulosus. We also speculate on the origin of northeastern populations of this Old World beetle. Distribution.—Anthribus nebulosus has Known distribution of Anthribus nebulosus in eastern North America. @, denotes collection localities been reported from the following Palearctic countries: Bulgaria, Finland, France, Ger- many, Hungary, Poland, and the USSR (Ukraine to eastern Kazakhstan). It occurs commonly in areas of deciduous forests, spruce forests, orchards, and on ornamental trees infested with prey scale insects. In the United States it has been known only from Virginia (Fig. 1), where adults originating VOLUME 93, NUMBER 1 47 Figs. 2-6. Rostra and tarsal segments of Anthribidae. 2, Dorsal aspect of rostrum of Ormiscus walshi (LeConte). 3, Rostral apex of Trigonorhinus sticticus (Boheman); arrow denotes central prolongation. 4, Rostral apex of Anthribus nebulosus Forster. 5, Metatarsus of T. sticticus; arrow denotes separate lobes of third tarsomere. 6, Metatarsus of A. nebulosus; arrow denotes connate or fused lobes of third tarsomere. from material collected in Europe were re- leased to help control various species of soft scales (Coccidae). As a result of our recent collecting, the following locality records are new for A. nebulosus (Fig. 1): UNITED STATES: CONNECTICUT: Middlesex Co., Middle- town, Wesleyan Univ., 28-V-1989, 6-VI- 1989, 15-VII-1989, and 1-X-89, ex Picea pungens, Middletown, Indian Head Cem- etery, 15-VII-1989, ex P. abies; East Hamp- ton, 15-VII-1989, ex P. abies; Tolland Co., Hebron, 15-VII-1989, ex P. abies; Somers, 15-VII-1989, ex P. abies; Storrs, Univ. Connecticut, 15-VII-1989, ex P. abies; Windham Co., Willimantic, 15-VII-1989, ex P. abies. MASSACHUSETTS: Berkshire Co., nr. Stockbridge, 16-VII-1989, ex P. abies; Hampden Co., East Longmeadow, 15-VII- 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1989, ex P. pungens; Hampshire Co., Am- herst, Amherst College, 16-VII-1989, ex P. abies, Amherst, Univ. Massachusetts, 16- VII-1989, ex P. glauca; Northampton, Smith College, 16-VII-1989, ex P. abies. NEW YORK: Dutchess Co., Annandale- on-Hudson, Bard College, 16-VII-1989, ex P. abies; Fishkill, Fishkill Rural Cemetery, 16-VII-1989, ex P. abies; Red Hook, 16- VII-1989, ex P. abies; Orange Co., War- wick, 22-VII-1989, ex P. abies; Ulster Co.., Kingston, St. Mary’s Cemetery, 22-VII- 1989, ex P. abies. Voucher specimens (C.U. Lot #1193) are deposited in the collections of Cornell Uni- versity (Ithaca, NY) and Pennsylvania De- partment of Agriculture (Harrisburg, PA). Recognition features.—Among the an- thribid taxa that occur in the eastern United States, the immigrant Anthribus nebulosus would most likely be confused with species of the genus Trigonorhinus. Several mor- phological and biological features charac- teristic for each genus were described by Valentine (1960). Species of Trigonorhinus can be readily recognized by the dorsal ros- tral surface being progressively narrowed from base to apex (Fig. 3), with the central apical portion distinctly produced beyond the corners and slightly emarginate (Fig. 3). The lobes of the third tarsal segment are separate, giving the appearance of a deeply lobed third segment (Fig. 5). In contrast, species of Anthribus lack the apical prolon- gation of the rostrum (Fig. 4), and the lobes of the third tarsal segment are connate or fused (Fig. 6). Trigonorhinus species are strictly plant feeders, as the majority of the Anthribidae, whereas species of Anthribus (formerly Brachytarsus Schoenherr) have larvae that are predacious on eggs of certain lecaniine scale insects (Valentine 1960). Anthribus nebulosus keys nearest to Tri- gonorhinus in Valentine (1960) because of the narrowed rostrum. Couplet 19 of that work is modified as follows to accommo- date this established immigrant. 19. Rostrum, excluding mandibles, quadrate, or slightly widened from base to apex (Fig. 2) Rostrum, excluding mandibles, narrowed from base to apex (Figs. 3-4) Rostral apex distinctly produced at middle, with central prolongation slightly emargin- ate (Fig. 3); lobes of third tarsomere separate (Rigs, 5) eee Trigonorhinus Wollaston (21 North American species, sensu Valentine 1960) Rostral apex truncate, lacking central pro- longation (Fig. 4); lobes of third tarsomere connate or fused (Fig. 6) ..... Anthribus Forster (1 North American species, A. nebulosus Forster) 19a. Biology and seasonal history.—Koszta- rab and Kozar (1983) thoroughly summa- rized the literature that documents Anthri- bus nebulosus as an effective predator of at least 15 species of scale insects in Europe and Central Asia. Three of these scales are also considered pest species in the eastern United States, i.e. the European fruit leca- nium, Parthenolecanium corni (Bouche); a nut scale, Eulecanium tiliae (L.); and a spruce bud scale, Physokermes hemicry- phus. They noted the potential for A. ne- bulosus to suppress populations of other pest scale insects in North America because it feeds on closely related scales in Europe. For additional references to aspects of bi- ology and seasonal history of A. nebulosus in Europe and in Virginia, the reader is re- ferred to Kosztarab and Kozar (1983) and Kosztarab and Rhoades (1983). Discussion.—In New England, the Eur- asian predator A. nebulosus is common on Norway spruce; we also collected it on Col- orado spruce and on white spruce (Picea glauca (Moench.) Voss). Available evidence suggests that populations of 4A. nebulosus in the northeastern states are adventive rather than associated with releases made in Vir- ginia. In many of the areas surveyed in southern New England, 20 or more adults could be quickly collected from a scale-in- fested spruce. Beetles were more difficult to obtain in eastern New York. The presence of large numbers in parts of Connecticut and VOLUME 93, NUMBER 1 Massachusetts seem to point to long-stand- ing populations rather than recent popula- tion outbreaks. The principal prey of this adventive pred- ator is the introduced Physokermes hemi- cryphus, a soft scale first collected in North America near Hartford, Connecticut, in 1906. Other early records of this scale are from Massachusetts, New Hampshire, New York, Pennsylvania, Ontario, and Wiscon- sin (Fenton 1917). Thus a suitable food source has existed in New England through- out the twentieth century. Although clearly speculative, we suggest the anthribid could have been introduced into the eastern states as early as the late nineteenth or early twentieth century. Be- fore passage of the Plant Quarantine Act of 1912, millions of seedlings and larger plants entered the United States each year (e.g. Sy- mons 1911), including insect-infested spruce trees (Marlatt 1912). The low dispersion rate reported for A. nebulosus in Virginia (Kosztarab and Kozar 1983) is evidence that the widespread New England populations are of long-standing origin. Even if some of the adults released in fall 1978 and 1979 had dispersed widely, it seems unlikely that the beetle could have become so abundant and widespread in New England within ten years. A mass appear- ance of A. nebulosus in New England due to passing weather conditions also cannot be dismissed, but any origin of northeastern populations based on long-range movement from Virginia seems implausible. Our surveys for the anthribid in New York just west of the Hudson River, and in parts of Maryland, New Jersey, Pennsylvania, Virginia, and West Virginia, were negative. These surveys were made on scale-infested spruces during the period of known adult activity, as reported by Kosztarab and Ko- zar (1983). This suggests that 4. nebulosus may not be established in these areas, or at least is not as widespread and abundant as in southern New England. Anthribus nebulosus apparently repre- 49 sents another example of a natural enemy that was accidentally introduced and that became established before it was considered for intentional introduction and release by biological control workers. Mason (1978) provided examples of Old World ichneu- monid parasitoids already established in North America before their intentional in- troduction for biocontrol of insect pests. Some similar cases are found in Clausen’s (1978) world review of introduced natural enemies. Goeden (1983) gave several ex- amples of the fortuitous introductions of in- sects later released for biocontrol of weeds. Goeden (1983) and Schroeder and Goeden (1986) advocated prerelease inventories of weeds targeted for biocontrol to minimize such “‘surprise”’ detections of adventive spe- cies. But A. nebulosus was not among the known enemies of Virginia coccids (Wil- liams and Kosztarab 1972:209-210), and Kosztarab and Kozar (1983) thus had no reason to suspect that the anthribid oc- curred anywhere in North America before it was collected in Hungary and released in Virginia. ACKNOWLEDGMENTS We thank Michael Kosztarab (VPI&SU, Blacksburg, VA) and Richard E. White (USDA, PSI, Systematic Entomology Lab., Washington, DC) for providing us with background information on the release of Anthribus nebulosus in Virginia, and James K. Liebherr (Cornell University) for criti- cally reading a draft of this paper. LITERATURE CITED Clausen, C. P. (ed.). 1978. Introduced parasites and predators of arthropod pests and weeds: A world review. United States Department of Agriculture, Agricultural Research Service, Agriculture Hand- book no. 480. 545 pp. Fenton, F. A. 1917. Observations on Lecanium corni Bouche, and Physokermes piceae Schr. Canadian Entomologist 49: 309-320. Gill, R. J. 1988. The scale insects of California. Part 1: The soft scales (Homoptera: Coccoidea: Coc- cidae). California Department of Food and Agri- 50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON culture, Technical Series in Agricultural Biosys- tematics and Plant Pathology 1. 132 pp. Goeden, R. D. 1983. Critique and revision of Harris’ scoring system for selection of insect agents in bi- ological control of weeds. Protection Ecology 5: 287-301. Kosztarab, M. and F. Kozar. 1983. Introduction of Anthribus nebulosus (Coleoptera: Anthribidae) in Virginia for control of scale insects: A review. Vir- ginia Journal of Science 34: 223-236. Kosztarab, M. and M. Rhoades. 1983. Food con- sumption, mating behavior, and shelter selection of Anthribus nebulosus Forster (Coleoptera: Anthribidae), an introduced predator of scale in- sects in Virginia. Virginia Journal of Science 34: 237-250. Marlatt, C. L. 1912. Some recent new importations. Journal of Economic Entomology 5: 73-77. Mason, W.R.M. 1978. Ichneumonoid parasites (Hy- menoptera) accidentally introduced into Canada. Canadian Entomologist 110: 603-608. Schroeder, D. and R. D. Goeden. 1986. The search for arthropod natural enemies of introduced weeds for biological control—In theory and practice. Bio- control News and Information 7: 147-155. Symons, T. B. 1911. A practical method of inspecting imported seedlings. Journal of Economic Ento- mology 4: 263-265. Valentine, B. D. 1960. The genera of the weevil fam- ily Anthribidae north of Mexico (Coleoptera). Transactions of the American Entomological So- ciety 86: 41-85. Williams, M. L. and M. Kosztarab. 1972. The insects of Virginia no. 5. Morphology and systematics of the Coccidae of Virginia with notes on their bi- ology (Homoptera: Coccoidea). Virginia Polytech- nic Institute and State University, Research Di- vision Bulletin 74. 215 pp. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 51-61 A NEW SPECIES OF STENELMIS (COLEOPTERA: ELMIDAE) FOUND WEST OF THE MISSISSIPPI RIVER Kurt L. SCHMUDE AND HARLEY P. BROWN (KLS) Department of Entomology, University of Wisconsin, Madison, Wisconsin 53706; (HPB) Department of Zoology, The University of Oklahoma, Norman, Oklahoma 73019. Abstract. — Adults of a new Nearctic riffle beetle, Stenelmis occidentalis, are described. Its range is restricted to areas west of the Mississippi River where it is one of only a few species in this genus to occur in Mexico, and the only Stene/mis known to occur in the northwestern U.S., Montana, Oregon, and Utah. Diagnostically useful characters include size, pronotal sculpturing, elytral color pattern, comparative lengths of the tarsomeres, metatibial spinous ridge on males, and the distinctive male genitalia. Adults are most similar to adults of Stenelmis decorata, and differences between the two, and adults of other species, are discussed. Adults occur in several types of lotic situations and they are found on a variety of substrates; they are also readily attracted to lights. Key Words: States, Mexico Stenelmis Dufour is the most diverse and widespread Nearctic genus of Elmidae (riffle beetles), with 30 described species occurring throughout most of North America north of Mexico (Brown 1983, 1987, White 1982). The senior author is revising the genus in North America, and this paper is part of that study. Adults of the following species are being described at this time to make the name available for a concurrent revisionary study of the riffle beetles of mid-America and the West Indies (Spangler and Santiago, in prep). This “new” species of Stene/mis actually has been known for some time. Texas spec- imens have been mentioned in the literature as Stenelmis sp. (near decorata) as early as 1963 by Burke, and by Stewart et al. (1973) and Poole and Stewart (1976) as Stenelmis mexicana(us), one of Brown’s manuscript names. In studies on a Montana stream, Gore (1975, 1977, 1978) referred to this species as Stenelmis sp. a, and specimens Elmidae, Stene/mis occidentalis, new species, riffle beetle, western United from Oregon have also been mentioned (Brown 1972). But structural differences be- tween northern and southern individuals, and their similarity to adults of other spe- cies, some of which are yet undescribed, prevented the junior author from describing the species separate from a revisionary study. MATERIALS AND METHODS Extensive collecting was accomplished in the southern range of the species by the ju- nior author, and additional material was ex- amined from the following collections: CASC—California Academy of Sciences, San Francisco, D. H. Kavanaugh & R. Brett CNCI— Canadian National Collections, Ot- tawa, Ontario, L. LeSage INHS—Illinois Natural History Survey, Urbana, K. C. McGiffen MCZC— Museum of Comparative Zoology, Harvard University, Cambridge, Mas- sachusetts, S. R. Shaw, D. Furth, S. Pratt 52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LSUC—Louisiana State University, Baton Rouge, C. B. Barr, J. B. Chapin NMNH—National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C., P. J. Spangler ODAC—Oregon Department of Agricul- ture, Salem, R. L. Westcott PERC—Purdue University, West Lafay- ette, Indiana, A. Provonsha TAMU-—Texas A&M University, College Station, H. R. Burke, E. G. Riley UGAM-— University of Georgia Museum of Natural History, Athens, C. L. Smith UMRM-— University of Missouri, Colum- bia, R. L. Blinn, K. B. Simpson USUC-— Utah State University, Logan, W. J. Hanson WSUC— Washington State University, Pullman, R. S. Zack CBB—Chery] B. Barr, Sacramento, Califor- nia HPB—Harley P. Brown, Oklahoma Muse- um of Natural History, Norman JRD—Jack R. Davis, Texas Water Com- mission, Austin DLG— Daniel L. Gustafson, Montana State University, Bozeman EGR—Edward G. Riley, Texas A&M Uni- versity, College Station KLS— Kurt L. Schmude, University of Wis- consin, Madison WDS-— William D. Shepard, California State University, Sacramento A Hitachi S-570 scanning electron mi- croscope was used at 5 kV or 10 kV to obtain the SEM photographs. Drawings of the gen- italia were made using a Prado Universal (Leitz Wetzlar) microprojector. All mea- surements were made at 72 x magnification with an ocular micrometer on a Leitz Wetz- lar dissecting microscope. Abbreviations and explanations of measurements used in the text and Table | are listed below: IOW—minimum interocular width; PL—maximum length of pronotum mea- sured along mesal plane; PW —maximum width of pronotum; PW/PL-—ratio of pronotal width vs length; EL—maximum length of elytra measured from anterior margin to apex along elytral suture with beetle’s venter situated in horizontal plane; EW—maximum width of elytra; EL/EW —ratio of elytral length vs width; PE length—length of pronotum and elytra, measured separately, and summed. Stenelmis occidentalis Schmude and Brown, NEw SPECIES Stenelmis sp. (near decorata) Burke, 1963: 3s Stenelmis sp. a Gore, 1975: 215, 1977: 115, 1978: 147. Stenelmis mexicana Stewart et al., 1973: 960 [nomen nudum]. Stenelmis mexicanus Poole and Stewart, 1976: 154 [nomen nudum]. Holotype male.— Head: IOW: 0.38 mm. Median dark band between eyes narrowed posteriorly; each light band equal in width to dark band at point between eyes. Anten- nae and palpi testaceous. Antennal and pronotal lengths equal. Pronotum (Figs. 1, 2): PL: 1.00 mm, PW: 0.91 mm. Widest just behind middle; sides narrowed toward base with a slight sinua- tion before base; abruptly narrowed before middle, then subparallel to apex. Median sulcus deep, widest and deepest anteriorly, gradually narrowed toward base. Median costae prominent, progressively narrowed and elevated toward base. Sulcus and costae obsolete in anterior 0.25 and before basal margin. Lateral tuberculi prominent, clearly separated by shallow, oblique lateral de- pression. Anterior tubercle round; posterior tubercle more elongate, weakly costate, and obsolete well before basal margin. Granules densely but uniformly distributed over dor- sal surface, except less dense in median sul- cus; granules of two or more sizes, smallest granules less than one-half the size of larg- est. 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Stenelmis occidentalis pronota; scale lines 0.5 mm. 1, alt—anterior lateral tubercle; plt— posterior lateral tubercle; mc—median costae; ms—median sulcus; old—oblique lateral depression; Texas specimen. 2, Montana specimen. Scutellum: Sixteen granules that are slightly larger than those on pronotum scat- tered over surface; smaller granules present also. Elytra (Fig. 3): EL: 2.30 mm, EW: 1.24 mm. Base of interval 3 moderately costate for 0.20 elytral length; more costate ante- riorly but obsolete before basal margin. Background color brown; each elytron faint- ly vittate. Medial portion of vitta diffuses with background color; anterior and pos- terior portions brighter. Vitta does not over- lap humeral angle; confined anteriorly be- tween striae 4-5; medially expanded to include portion of interval 4; includes stria 3 posteriorly; ends beyond lateral carina but before apical margin. Interval 1, and much of posterior portion on outside of lateral carina, also lighter than background color. Venter: Apical emargination of last ster- num wide and deep, but narrower than api- cal width of 5th metatarsomere. Legs (Figs. 4, 5): Each femur with nu- merous granules of different sizes scattered over surface. Femur and most of each tibia same color as pronotum; apex of tibia and each tarsus testaceous. In dorsal view, basal half of each Sth tarsomere narrow with sides subparallel, next quarter length dilated, api- cal quarter length subparallel. Fifth tarso- mere of pro- and mesotarsi slightly longer (0.03 mm) than their preceding four tarso- meres combined; 5th metatarsomere equal in length to its four preceding tarsomeres. Apicoventral margin of 5th tarsomere slightly convex. Conspicuous spinous ridge on distal half of inner margin on mesotibia. Distal half of metatibia with a weak spinous ridge on inner margin. Genitalia (Figs. 6, 7): Penis widest at base, constricted toward middle where it becomes swollen, then subparallel to its rounded apex. Each paramere with basodorsal portion wide, elongate, nearly truncate distally, and longer than apicodorsal portion; inner dor- sal margin slightly divergent distally, and VOLUME 93, NUMBER 1 2 oe & “7 & i. La. fl i" t* ie. ; i i ’ 4 Mi Fig. 3. Stenelmis occidentalis elytra; scale line 0.5 mm; c—costae on third intervals; Montana specimen. abruptly angled toward outer margin at apex; outer margin subparallel basally and con- vergent apically with a shallow sinuation before apex; inner ventral margin slightly sinuate apically, then broadly concave to base. Allotype.—IOW: 0.39 mm, PL: 1.05 mm, PW: 0.91 mm, EL: 2.40 mm, EW: 1.25 mm. Essentially similar, but larger. Sides of pro- notum more sinuate at base, more conver- gent at apex. Color of interval | on elytra not noticeably lighter than background col- or. Fifth tarsomere of pro- and mesotarsi slightly longer (0.04 mm) than preceding four tarsomeres combined. Without sexual char- acters (spinous ridge) on meso- and meta- tibiae. Variations.— Variation within the type series is minimal. The antennae can be slightly shorter than the pronotum. Some specimens are essentially and faintly bi- Stenelmis occidentalis meso- and me- Figs. 4, 5. tatibiae of male; scale lines 0.1 mm, insert scale lines 0.5 mm; Montana specimen. 4, mesotibia with con- spicuous spinous ridge on inner margin. 5, metatibia with weak spinous ridge on inner margin. 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON aE 5 Te dorsal; parameres spread wider than natural; scale line 0.25 mm. 6, Mexico specimen. 7, Montana specimen. Stenelmis occidentalis male genitalia; maculate, with narrow anterior and poste- rior maculae confined to interval 5, and interval 3 to stria 5, respectively. The meso- and metatibial spinous ridges on males can be slightly less or more conspicuous. Variation of specimens from the south- eastern portion of the range is similar to that found within the type series. Pronotal and elytral sculpturing can be less pronounced. As few as nine large granules occur on the scutellar surface. Elytral color pattern varies from nearly immaculate, narrowly (interval 5) to broadly (interval 4 to stria 5) maculate or vittate. Background color of elytra is light to dark brown to black; diffusion of the tes- taceous markings with the light brown back- ground color is common. In males, the Sth tarsomere of the pro- and mesotarsi can be even longer (0.04-0.06 mm) than the pre- ceding four tarsomeres combined. In both sexes, the 5th metatarsomere can vary from being 0.03 mm shorter to 0.03 mm longer than the preceding four tarsomeres com- bined. The weak metatibial spinous ridge on males is inconspicuous to distinct. Male genitalia vary slightly in size and the medial swelling of the penis is reduced and not dis- tinctly evident in some. In general, northern specimens are larger. We found a male and female from South Dakota and a female from Utah with PE lengths more than four millimeters; these are the largest Stenelmis recorded from North America. In northern specimens the antennae are distinctly shorter than the comparatively longer pronotum. The an- terolateral pronotal margin can be sinuate. The pronotal and elytral sculpturing are more pronounced. Up to 27 large granules occur on the scutellar surface. The elytra are maculate to vittate and the testaceous mark- ings are usually wider and cover as much as a part of interval 3 to stria 5 anteriorly, and striae 2-5 posteriorly. When vittate, the medial portion varies considerably in width, from narrow (striae 4—5) to wide (interval 3 to stria 5), and it is usually fainter than other portions, while it nearly diffuses with the background color in some beetles. Pos- terior area on the outside of the lateral ca- rina can also be lighter than the background color. Lengths of the Sth tarsomere on the pro- and mesotarsi of females are 0.03—-0.06 mm longer than the preceding four tarso- meres combined, but are equal to 0.05 mm longer in males. The Sth metatarsomere is 0.01 mm shorter to 0.01 mm longer in fe- males, and 0.04 mm shorter to equal in males. The metatibial spinous ridge on males, although low, is distinct on most in- dividuals. Besides being larger, the penis of most individuals is even more swollen me- dially, and the outer dorsal margin of the parameres can be more deeply sinuate be- fore its apex. Diagnosis.—This species cannot be in- serted into the adult keys of Sanderson (1938) or Brown (1972) because the distin- guishing characters are not in accordance with the key’s structure. Rearranging the keys to accommodate adults of S. occiden- talis would not be prudent at this time be- cause several new, undescribed species must also be included, and a reorganization of the VOLUME 93, NUMBER 1 key based, in part, on different characters is required. Instead of providing a key, we dis- tinguish adults of S. occidentalis from adults of similar species in the discussion below. Although individuals in its northern range make this species the largest Stene/mis in North America, some specimens from southern Texas are relatively small and could easily be confused with specimens of other species. Adults of S. occidentalis are most similar to those of S. decorata San- derson and S. n. sp. (identified as S. bicari- nata in Barr and Chapin 1988), but can be separated by comparing measurements and ratios of the head, pronotum, and elytra as listed in Table 1. In addition, the more pro- nounced pronotal sculpturing, the elongate but not distinctly costate basolateral tuber- cle, and the dense but uniform pronotal granulation, with different-sized granules, are distinctive characters. The elytral mark- ings of most specimens do not contrast as sharply with the background color as they do in individuals of most species. The com- bination of the 5th tarsomere on the pro- and mesotarsi most commonly being longer than their preceding four tarsomeres, while it is shorter or subequal on the metatarsi, along with the large mesotibial spinous ridge and smaller metatibial ridge in males, is di- agnostic. The male genitalia, without a lat- eral flange on the penis, is very similar to the genitalia of most species in Sanderson’s (1938) crenata group, but especially to S. decorata in his sinuata/humerosa group. The constricted and then swollen medial area on the penis is unique, and when combined with its rounded apex and the characteris- tics of the parameres, males are easily iden- tified. Type data.— Holotype, allotype, 33 para- types: TEXAS: Bastrop Co., Bastrop State Park, 24 June 1963, at light, S. G. Wellso. Holotype and allotype are on permanent loan to the NMNH from TAMU. Paratypes will be deposited in the following collec- tions: 15 TAMU, 4 NMNH, 4 CASC, 2 CNCTI, 2 INHS, 2 Snow Entomological Mu- Sif seum, University of Kansas, 2 Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico, D.F., 2 HPB. Additional specimens examined. — An ad- ditional 2626 specimens were examined. Following each collection record, the num- bers of individuals are in parentheses and immediately precede depository institu- tions or private collections, which are ab- breviated as in Methods section. Arkansas. Scott Co. Mill Cr., 5 mi E Y City, 9 Jul 1965, J. L. Donahue (1 KLS). Washington Co. 24 Jul 1940 (1 INHS). Arizona. Yuma, HornColl H3951 () “—MCZC); Yuma; HornColl H (1 MCZC). Colorado. Mesa Co. Cameo Power Plant, 8 Jul 1977, A. A. Bork- si(2 UMRM; | KLS). Louisiana. Grant Par. Stuart Lk. Rec. Area, 22-29 Jul 1982, light trap, EGR (2 LSUC). Montana. Tongue R., 21 Sep 1978, R. Oswald (26 HPB). Bighorn Co. Busby, Rosebud Cr., 29 Aug 1989, DLG (34 DLG; 16 KLS). Carter Co. Alzada, Lit- tle Missouri R., 30 Aug 1989, DLG(1 KLS). Chouteau Co. Ft. Benton, Missouri R., R. Oswald, summer 1978 (2 HPB). Golden Valley Co. Ryegate, Musselshell R., 1 Sep 1989, DLG (18 DLG; 15 KLS). Musselshell Co. Melstone, Musselshell R., 1 Sep 1989, DLG (2 KLS). Rosebud Co. Ashland, Tongue R., 30 Aug 1989, DLG (19 DLG; 13 KLS). Birney, Tongue R., 1975, J. A. Gore (7 KLS). Mouth of Rosebud Cr., 31 Aug 1989, DLG (43 DLG; 28 KLS). Rose- bud Cr., 10 Jun 1976, S. Baril (51 HPB). Nebraska. Dawes Co. White R., 8 mi NE Chadron, 17 Jun 1984, W. P. McCafferty, A. V. Provonsha (6 PERC; 6 KLS). Nevada. Clark Co. Moapa, Muddy R., 12 Dec 1986, WDS (2 KLS; 4 WDS), Muddy R. & Warm Springs outflow stream, 6.5 mi NW Moapa, 12 Dec? 1986, CBBC (CBB). Lincoln: Go: Ash Springs outflow streams, U.S. Hwy 93, 125 Dee 1986..CBB (7-ESUCG: 16;:CBB: 9 KLS; 10 WDS). Oklahoma. Caddo Co. Ana- darko, Washita R., 20 Jul 1963, HPB (7 HPB). Verden, Washita R., 31 May 1971, HPB (45 HPB). Cleveland Co. Norman, 25 Aug 1975, 3 Jun—13 Aug 1976, light trap, 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON WDS (3 WDS). Comanche Co. East Cache Cr., 1 Jul 1963, HPB (36 HPB). Custer Co. Washita R., 2 mi W Clinton, 5 Nov 1977, WDS (5 WDS), 8 mi E Cordell, 5 Nov 1977, WDS (6 WDS). Garvin Co. Washita R. N Wynnewood, | Sep 1967, HPB (21 HPB). Grady Co. Washita R., 6 Oct 1963, HPB (1 HPB). Johnston Co. Tishomingo, Penning- ton Cr., 25 Jul 1972, HPB(1 HPB). Latimer Co. 5 mi W Red Oak, 2 Jul 1977, K. Stephan (1 HPB)! Eove Co. Mud’ Cr, 5 Jul 1963; HPB (1 HPB). Major Co. creek W Cleo Springs, 1800 ft elev., 5 Sep 1971, HPB (10 HPB). McClain Co. S. Canadian R., New- castle, 18 Aug 1976, WDS (2 WDS). McCurtain Co. Glover R., 25 Jul 1972, HPB (1 HPB). Stephens Co. Wild Horse Cr., 26 Jul 1962, HPB (34 HPB). Washita Co. Cor- dell, 4 Jul 1969, HPB (30 HPB). Oregon. Malheur Co. Ontario, Exp. Sta., 2 Aug 1960, light trap, F. P. Larson (2 ODAC; 2 HPB). South Dakota. Fal/ River Co. Cheyenne R., U.S. Hwy 38, 11 Jun 1975, W. P: Mc- Cafferty, A. V. Provonsha, B. L. Heath (4 PERC; 3 KLS). Texas. Anderson Co. 10 mi SW Elkhart, 23 Jul 1960, light trap, H. R. Burke (3 TAMU). Salmon, 27 Jun 1975, light trap, H. R. Burke (4 TAMU). Trinity R., U.S. Hwy 287 nr Cayuga, 6 Oct 1987, JRD (2 JRD). Bastrop Co. Bastrop St. Pk., 24 Apr 1962, light trap, S. G. Wellso (10 CASC; 10 TAMU; 6 INHS), 13 Jun 1969, S. Peck (5 INHS). Bee Co. Aransas R. E Papalote, 29 Apr 1982, JRD (3 JRD; 2 KLS). Bexar Co. San Antonio, 2 Jun 1958, J. F. Lawrence (1 HPB; 2 MCZC), 4 Jun 1975, M. Druckenbrod (3 NMNH). Blanco Co. John- son City, 5 Oct 1966, HPB (2 HPB). John- son City, Pedernales R., 27 Aug 1967, HPB (9 HPB). Bosque Co. 1 mi N Clifton, 6 Jul 1967, 19 Aug 1970 (13 TAMU). 2 mi W Iredell, 1 Jul 1966 (2 TAMU). Morgan, Steele Cr., 23 Dec 1974, 30 Dec 1976, WDS (4 WDS). Brazos Co. Bryan, 16-21 Jul 1965, 28 May-12 Jul 1966, light trap, J. C. Schaff- ner (17 TAMU), 25 Jun-8 Aug 1988, EGR (7 TAMU; 4 EGR). Cedar Cr., Oct 1971, R. G. Phelps (1 TAMU). College Station, 10 May 1964, H. R. Burke (1 TAMU), 5- 10 Aug 1968, 1 Jun 1970, V. V. Board (30 TAMU), 1966, M. H. Sweet (4 TAMU), 21 Aug 1972, light trap, W. E. Clark (1 TAMU), 17 Jun 1989, EGR (4 TAMU). Little Brazos R., Hwy 21; 3: Jul 1970, V. Board) é& R- Phelps (18 TAMU). Caldwell Co. Plum Cr., U.S. Hwy 183, 11 Sep 1987, CBB (3 LSUC; 4 CBB). Coke Co. NE Bronte, 29 May 1969, HPB (25 HPB). Colorado Co. Columbus (29 NMNH). Coleman Co. Coleman, 29 May 1969, HPB (1 HPB). Comal Co. Guadalupe R., 1.8 mi E Spring Branch, Rte 33, 16 Dec 1989, R. S. Zack (15 WSUC). Guadalupe R. St. Pk., 11 Sep 1987, light trap, CBB (3 LSUC; 13 CBB; 1 KLS). Wimberly, Blanco R. 25 May 1969, HPB (1 HPB). Comanche Co. De Leon, Sabanna R., 29 May 1969, HPB (8 HPB). Concho Co. Concho R. S Lowake, 23 Jul 1982, JRD (8 JRD; 4 KLS). Cooke Co. Gainesville, Elm Fk. Trinity R., 21 Jun 1978, WDS (1 WDS). Coryell Co. Gatesville, 20 Aug 1970, HPB (8 HPB). Crosby Co. Crosbyton, 22 Apr 1972, HPB (1 HPB). Denton Co. Argyle, Denton Cr., 28 Apr 1939, J. A. & H. H. Ross ( 8 INHS). Denton Cr., 12 mi S Denton, 21 Jun 1978 (3 WDS). 3 mi S Denton, U.S. Hwy 35, 22 Jul 1975 (12 WDS). Elm Fk., 15 Nov 1970, D. McNeely (7 WDS). DeWitt Co. Coleto Cr. nr Weser, 28 Mar 1974, HPB (29 HPB). NE Westhoff, 28 Mar 1974, HPB (77 HPB). Brook S Westhoff, 28 Mar 1974, HPB (9 HPB). Duval Co. Sepulveda Ranch, 4 Jul 1970, C. W. Griffen (1 NMNH). Ellis Co. Waxahachie, 31 Aug 1967, HPB (7 HPB). Fort Bend Co. Missouri City, 31 May 1977, light trap, EGR & LeDoux (8 EGR; 2 KLS). Frio Co. Pearsall, 26 Jun 1970, J. W. Smith (1 TAMU). Gillespie Co. Pedernales R., 4 mi S Fredericksburg, 23 Jun 1978, WDS (4 WDS). Goliad Co. Goliad, San Antonio R.., 23 Jul 1962, H. R. Burke (1 TAMU). Small creek E Goliad, 29 Mar 1974, HPB (2 HPB). Gonzales Co. Gonzales, Guadalupe R., 28 Mar 1974, HPB(57 HPB). Palmetto St. Pk., 7 Jun 1969, Board & Hafernik (61 TAMU), 14 Jun 1969, M. W. Sanderson (125 INHS), VOLUME 93, NUMBER 1 30 May 1977, light trap, EGR (6 LSUC; 20 NMNH,; 50 UMRM;; 39 HPB; 11 EGR; 25 KLS), 30 May 1983, 27 Apr 1988, CBB (3 LSUC; 5 CBB), San Marcos R., 31 May 1983, CBB (2 CBB). Grimes Co. Navasota, Navasota R., 31 Aug 1967, HPB (9 HPB). Navasota R., Hwy 158, 15 Aug 1962, Reeves & Burke (1 TAMU). Guadalupe Co. Shertz, 25 Jun 1985, light trap, D. W. Sundberg (2 LSUC; 12 CBB). Hamilton Co. Hico, N. Bosque R., 22 Jun 1978, WDS (45 WDS). Leon R., 6 mi N Hamilton, 22 Jun 1978, WDS (133 WDS). Harris Co. Houston, Buf- falo Bayou, 16 Jul 1985, JRD (6 JRD; 2 KLS). Seabrook, 19 Aug 1970, light trap, WDS (1 WDS). Hays Co. Buda, Onion Cr., 17 Sep 1960, H. Chambers (8 TAMU). San Marcos, San Marcos R., 30 Sep 1964, 9 Aug 1969, HPB (4 HPB). San Marcos R. 12 Jul 1967, light trap, G. S. Tulloch (1 TAMU), 27 Aug 1974, J. Kolb (2 NMNH). Hidalgo Co. Bentsen-Rio Grande St. Pk., 18 Jun 1969, Board & Hafernik (1 TAMU), 9-10 Jun 1975, light trap, R. Turnbow (6 HPB; 8 UGAM), 17-18 May 1979, 4 Oct 1986, 15-16 Oct 1988, EGR (2 NMNH; 3 TAMU; 2 EGR). Santa Ana Nat. Wldlf. Ref., 23 Oct 1987 (2 EGR). Hill Co. Blum, Nolan Cr., 23 Dec 1974, WDS (1 WDS). Kaufman Co. Trinity R., Hwy 34 nr Rosser, 26 Aug 1987, 12 Oct 1988, JRD (20 JRD; 7 KLS). Kendall Co. Boerne, Cascade Caverns, 20 Jul 1981, W.E. Steiner (2 NMNH). Mission, 17 Jul 1981, light trap, W. E. Steiner (5 NMNH). Lavaca Co. Navidad R., 10 & 12 mi N Hal- lettsville, 23 Apr 1961, 28 Jul 1961, H. R. Burke (2 TAMU). Limestone Co. Groes- beck, Navasota R., 18 Oct 1969, HPB (2 HPB). Live Oak Co. 12 mi N Mathis, 7 Aug 1971, light trap, W. H. Tyson (1 NMNH). Whitsett, Atascosa R., 25 Aug 1962, J. Haf- ernik (1 TAMU). Llano Co. Castell, Llano R., 1 Oct 1960, H. R. Burke (2 TAMU). McCulloch Co. San Saba R., 16 mi S Brady, 13 Jun 1963, D. C. & K. A. Rentz (363 CASC; 26 KLS). McLennan Co. Bosque R. nr Waco, 14 May 1961, B. McDaniel (2 TAMU). Medina Co. Hondo, Hondo Cr., 59 25 May 1969, HPB (4 HPB). Menard Co. Menard, May 1946, L. J. Bottimer (6 CNCI). Montgomery Co. Conroe, W. Fk. San Ja- cinto R., 25 Jun 1967, HPB (2 HPB). The Woodlands, 1-10 Jul 1978, R. Turnbow (2 UGAM), 24-30 Jun 1979, J. E. Wappes (1 TAMU). Nacogdoches Co. Douglas, Ange- lina R., 10 Aug 1969, HPB (1 HPB). Nolan Co. Sweetwater, 10 Jul 1937 (1 TAMU). Nueces Co. Corpus Christi, 26 Jun, 1 Jul 1970, light trap, C. W. Griffen (6 TAMU). Palo Pinto Co. Brazos R., Hwy 4, 30 May 1973, B. Stark (2 HPB), 29 Jul 1982, WDS (3 WDS), 4 mi W Mineral Wells, 27 Jun 1978, WDS (63 WDS). Dark Valley, 29 Jul 1970, D. McNeely (2 WDS). 4—5 mi SW Graford, 25 Aug 1981, WDS (53 WDS; 15 KLS). Parker Co. Dennis, Brazos R., 24—25 Jun 1989, EGR (1 TAMU). Robertson Co. Camp Cr., 10 Jun 1962, light trap, V. House (1 TAMU). Runnels Co. Ballinger, Colo- rado R. Rte 83, 19 Dec 1989, R. S. Zack (5 WSUC). San Patricio Co. 6 mi W Edroy, | Jul 1970, C. W. Griffen (4 TAMU). Lake Corpus Christi St. Pk., 9 Jun 1969, Board & Hafernik (5 TAMU). San Saba Co. San Saba R., 5 mi W San Saba, 22 Jun 1978, WDS (2 WDS). Somervell Co. Glen Rose, Paluxy R., 23 Dec 1974, WDS (8 WDS). Tarrent Co. Benbrook, Trinity R., 27 May 1971, 13 Jun 1974, WDS (50 WDS). 5 mi S Benbrook, 21 Jun 1978, WDS (7 WDS). Tom Green Co. Concho R., 14 mi E San Angelo, 12 Jun 1961, H. R. Burke (2 TAMU). Travis Co. Austin, Onion Cr., 19 Aug 1970, 4 Jul 1978, HPB (5 HPB). Vic- toria Co. Raisin, Coleto Cr., 29 Mar 1974, HPB (5 HPB). sandy creek W Inez, 29 Mar 1974, HPB (12 HPB). San Antonio R., 16 mi S Victoria, 20 Aug 1962, H. R. Burke (4 TAMU). Victoria, 26 Jun 1965, P. J. Spangler (3 NMNH). Washington Co. Washington-on-the-Brazos St. Pk., 12 Jun 1969, M. W. Sanderson (21 INHS). Whar- ton Co. Wharton, 19 Aug 1970, light trap, WDS (1 WDS). Williamson Co. San Gabriel R., 6 mi E Georgetown, | Oct 1960, H. R. Burke (5 TAMU), Taylor, 16 Jun 1967, J. 60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 8. see section on distribution. E. Hafernik (2 TAMU). Wilson Co. Cibolo Cr., 8 May 1980, JRD (34 HPB). Zapata Co. Falcon St. Pk., 11 Jun 1975, light trap, R. Turnbow (7 UGAM). Falcon St. Rec. Area, 5 Jun 1983, light trap, CBB & D. A. Rider (25 LSUC; 47 CBB; 16 KLS). Utah. Box Elder Co. 5 Oct 1971, D. W. Shaw (1 HPB), 5 Jul 1972, R. Newell (2 HPB). Deep Cr., 3 Jan 1971, D. W. Shaw (2 HPB). Field- ing, Malad R., 14 Aug 1954, A. Gaufin (4 HPB). Cache Co. Logan, 24 Jul 1974, G. F. Knowlton (1 USUC). Duchesne Co. Ft. Du- chesne, Uinta R., 31 Mar 1973, 5 Oct 1974, B. P. Stark (14 HPB). Grand Co. Arches Nat. Mon., Moab, 12 Jul 1964, J. T. Doyen (1 HPB). Uintah Co. Uinta R., Hwy 40, 23 May 1974, B. P. Stark (2 HPB). Ashley Cr. nr Vernal, 1 Apr 1973, 5350 ft, B. P. Stark (7 HPB). Wayne Co. 14 mi S Hanksville, 25 Jul 1968, at light, H. F. Howden (1 CNCI). Weber Co. Ogden, 10 Aug 1960, G. Distribution of Stenelmis occidentalis. Star represents type locality. Arizona record is questionable; F. Knowlton (1 HPB). Mexico. Coahuila Rio San Rodrigo, E El Remolino, 26-27 May 1969, HPB (3 HPB). Villa Fuente, 26 May 1969, 19 Aug 1970, HPB (27 HPB). Zaragoza, 26 May 1969, HPB (14 HPB). Distribution (Fig. 8).—Stenelmis occiden- talis is the most common and widespread species of Stene/mis restricted to areas west of the Mississippi River, and one of only a few found in Mexico. It is also the only species known to occur in the northwestern U.S. (MT, OR, UT), an area formerly be- lieved to be devoid of the genus. Although there are large areas in its range from which we have not seen specimens, we suspect a lack of comprehensive collecting is the cause of its apparent absence. Two specimens from Yuma, Arizona were found in the Horn- LeConte Collection, MCZC. Since these venerable beetles have never been men- tioned in the literature, and we are unaware VOLUME 93, NUMBER 1 of any other records from Arizona, their label data may be questionable, but not 1m- possible. Habitat.— Adults occur in several types of lotic situations and on a variety of sub- strates. In southeastern Montana, a popu- lation of adults was found on rocks in Rose- bud Cr., a shallow, very warm, five-foot wide ditched creek, with minimal flow; the creek supported a massive algal population and the beetles were covered with algae. Further downstream aggregations of 10-20 individ- uals were found in depressions on rocks, just above the anaerobic stain on the rocks. In contrast, adults were also found in the shal- low, turbid, 30-foot wide Musselshell R. where they occurred in aggregations on large rocks in fast current near the shore. Others were collected from woody debris and over- hanging shoreline vegetation in the Tongue R., a deep, turbid, 50-foot wide river. In southern Nevada, adults were found in a small stream and spring outfall. The water was clear and warm, and there was a mod- erate to fast current with a sand-gravel-rock substrate; filamentous algae, woody debris, and travertine were abundant. In Texas and Oklahoma, S. occidentalis was most com- monly collected from rocks and submerged wood in sandy streams. The type series was collected at a light trap. Newly emerged adults are readily attracted to lights. Etymology.—Named after its wide- spread, western distributional pattern. ACKNOWLEDGMENTS We extend our gratitude to William L. Hil- senhoff for his invaluable assistance and ed- itorship, and Paul J. Spangler for reviewing and improving the manuscript. For provid- ing beetles to study, we are grateful for the cooperation by the people and their insti- tutions mentioned in the methods section, including James A. Gore for sending voucher specimens collected during his studies. Ad- ditional habitat information was affably fur- nished by Cheryl B. Barr, Daniel L. Gustaf- 61 son, and William D. Shepard. Also, we thank Daniel K. Young for his support, and both he and Paul J. Johnson for their interest in this research. Melissa J. Curtis provided co- pious assistance with the SEM. Support was provided by the Graduate School and the College of Agriculture and Life Sciences, University of Wisconsin, and by the Theo- dore Roosevelt Memorial Fund, American Museum of Natural History, New York. LITERATURE CITED Barr, C. B. and J. B. Chapin. 1988. The aquatic Dry- opoidea of Louisiana (Coleoptera: Psephenidae, Dryopidae, Elmidae). Tulane Stud. Zool. Bot. 26: 89-164. Brown, H. P. 1972. Biota of freshwater ecosystems identification manual no. 6. Aquatic dryopoid beetles (Coleoptera) of the United States. Water Pollution Control Research Series, U.S. Environ. Prot. Agency, Washington, D.C. 92 pp. . 1983. A catalog of the Coleoptera of America north of Mexico, family: Elmidae. U.S. Dep. Agric., Agric. Handb. No. 529-50. 33 pp. . 1987. Stenelmis cheryl: new name for a well- known riffle beetle (Coleoptera: Elmidae). Ento- mol. News 98: 111-112. Burke, H. R. 1963. Notes on Texas riffle beetles (Co- leoptera, Elmidae). Southwest. Nat. 8: 111-114. Gore, J. A. 1975. Fall-winter composition of the ben- thic macroinvertebrates of the Tongue River, Montana. Proc. Fort. Union Coal Field Symp. 2: 212-225. 1977. Reservoir manipulations and benthic macroinvertebrates in a prairie river. Hydrobio- logia 55: 113-123. 1978. A technique for predicting in-stream flow requirements of benthic macroinvertebrates. Freshwater Biol. 8: 141-151. Poole, W. C. and K. W. Stewart. 1976. The vertical distribution of macrobenthos within the substra- tum of the Brazos River, Texas. Hydrobiologia 50: 151-160. Sanderson, M. W. 1938. A monographic revision of the North American species of Stenelmis (Dry- opidae: Coleoptera). Univ. Kans. Sci. Bull. 25: 635-717. Stewart, K. W., G. P. Friday, and R. E. Rhame. 1973. Food habits of hellgrammite larvae, Corydalus cor- nutus (Megaloptera: Corydalidae), in the Brazos River, Texas. Ann. Entomol. Soc. Am. 66: 959-963. White, D. S. 1982. Stenelmis morsei, a new species of riffle beetle (Coleoptera: Dryopoidea: Elmidae) from South Carolina. Coleopt. Bull. 36: 170-174. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 62-66 A NEW SPECIES OF JAPANAGROMYZA (DIPTERA: AGROMYZIDAE) FROM FLORIDA, WITH A KEY TO NORTH AMERICAN SPECIES BRIAN M. WIEGMANN Department of Entomology, University of Maryland, College Park, Maryland 20742. Abstract.—The new species Japanagromyza polygonivora is described from Florida, based on specimens reared from blotch mines in a smartweed, Polygonum sp. (section Persicaria). This 1s the first record of an agromyzid attacking this plant in North America. A revised key to the North American species is provided to help separate this species from other known members of the genus. Key Words: Japanagromyza, originally described for six species from Japan (Sasakawa 1958), is now known from nearly all zoogeographic regions. It is represented in the Indopacific by 16 species (Sasakawa 1963), 7 in Aus- tralia (Spencer 1977), and 20 in the Neo- tropical region (Spencer and Stegmaier 1973). In North America, only 5 species were recorded until Spencer and Steyskal (1986) added an additional one from the Western United States, which significantly broadened the generic concept. The latter authors also provided a revision of the North American species. Larvae of this genus attack a wide variety of hosts. Nearly all make blotch mines and pupate in the soil (Spencer and Steyskal 1986). In North America, species of Japan- agromyza have been recorded mining leaves of Fabaceae and Fagaceae. Although in Eu- rope, Agromyza pittodes Hendel is known to attack Polygonum viviparum and A. po- lygoni Hering is recorded from Polygonum bistorta, no North American species of agro- myzid has ever been associated with plants in the family Polygonaceae. The discovery of Japanagromyza poly- gonivora, new species, from Florida, mining Agromyzidae, Japanagromyza, Polygonum, smartweed, leaf-miner the leaves of a smartweed Polygonum sp. (sect. Persicaria), is of considerable interest taxonomically, as well as biologically. This fly lacks the prescutellar bristles and has 2 pairs of dorsocentrals showing its possible relationship with J. perpetua and J. des- modivora. Because species of the genus nor- mally have only 2 pairs of dorsocentrals, a character common in the genus Melanagro- myza, and it frequently possesses prescu- tellars, a character of the genus Agromyza, it is often considered intermediate between these two groups. These characters vary sig- nificantly throughout the group, however, and their phylogenetic utility must be left uncertain until a more detailed analysis is undertaken. The presence of two apical midtibial bristles indicates its placement in the Japanagromyza. The distinctive shapes of the epandrium and surstylus indicate that it is new. In this paper, I describe as new Japan- agromyza polygonivora, give a brief generic diagnosis, and provide a key to the North American species of the genus. METHODS In this paper, morphological terminology follows the convention used by Spencer and VOLUME 93, NUMBER 1 Steyskal (1986) to allow for effective com- munication and comparison of male geni- talic features. There has been considerable debate over the homologies of the structures of the male genitalia of higher Diptera (Griffiths 1972, McAlpine 1981, Wiegmann 1989), leading to alternative terminologies for these structures. My use of the terms epandrium and surstyli (McAlpine 1981, Spencer and Steyskal 1986) (= periandrium and gonostyli of Griffiths 1972) is not an endorsement of either genitalic homology theory, but reflects the usage of other work- ers in Agromyzidae (i.e. Spencer 1976). Descriptions of characters were made us- ing a Wild Optics dissecting scope at 25x. Measurements were made with an objective micrometer. Genitalia were dissected cleared in KOH, and preserved in glycerin for fur- ther study. Leaf mines were collected from Brevard Co., near Melbourne, Florida on 15-v-1986 by K. Hibbard and F. Smith, and adults were reared by H. Weems. Emer- gence was on or near 10-x1i-1986. For in- formation on rearing Agromyzidae from their mines see Spencer and Steyskal (1986). Japanagromyza Sasakawa, 1958: 138. Type species Agromyza duchesneae Sasakawa, 1954: 106, original designation. Diagnosis: Robust black flies, 1.5—3.0 mm long; frons normally black or brownish, ap- proximately equal to width of eye, as long as or narrower than wide; upper orbitals 2; lower orbitals 2; gena narrow, 0.05-0.12 of eye height; I st flagellomere rounded to oval; arista bare or with microscopic pubescence; mesonotum normally with 0+2 dorsocen- trals; acrostichals 8 to 10 rows; prescutellars 1 pair, sometimes lacking; halter white or yellow, often with black base; wing 1.5—2.5 mm long; costa reaching M1+2; foretibia with one lateral bristle, midtibia with two; male genitalia with epandrium ventrally ar- ticulated to hypandrium; surstylus directed inwards or downwards, often with several stout spines or terminal uncus; cercus vari- able often with stout internal spines, or long 63 Figs. 1-3. Japanagromyza polygonivora Wieg- mann n. sp. male. 1, Head, lateral view (bar = 0.07 mm); 2, Epandrium with (A) surstylus, (B) cercus, lat- eral view (bar = 0.03 mm); 3, Aedeagus (bar = 0.04 mm). black hairs; aedeagal hood lightly sclero- tized or partly membranous; distal tubule long, membranous, often coiled. KEY TO NORTH AMERICAN JAPANAGROMYZA ADAPTED FROM SPENCER AND STEYSKAL (1986) 1. Dorsocentrals 3+1; frons reddish net Ri Vt RSRAG TD ae. ee a rutiliceps (Melander) - Dorsocentrals only 2 post sutural (0+ 2); frons blacksonibrowntere enact ree 2 2()serescutellarsyabsentssn- oreo cae oii 3 - Prescutellars present 3(2) Mesonotum distinctly greenish; cercus with numerous stout spines ...... perpetua Spencer - Mesonotum matt grayish black, at most only faintly greenish; cercus with few or no stout spines, but with long fine hairs ............ 4 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 4-8. Japanagromyza polygonivora Wieg- mann n. sp. male. 4, Hypandrium (bar = 0.04 mm); 5, sperm pump (bar = 0.015 mm); female 6, sperma- theca (bar = 0.02 mm); 7, cerci (bar = 0.03 mm); 8, egg guide, lateral view (bar = 0.03 mm). 4(3) Mesonotum primarily matt grayish black, only faintly greenish; cercus with several stout spines and unusually long black hairs ..... Si iaae ee SAR ae etn SARE desmodivora Spencer —- Mesonotum matt grayish black, never green- ish; cercus without stout spines but with many longelinesblackshainsi\.05 a.t9 eee eases bounties ts polygonivora Wiegmann n. sp. »@)eMesonotumboreenishts.. 2a. naeeeeo. 3. 6 - Mesonotum predominately black ......... BON Bee ree RRS i a4, ee viridula (Coquillett) 6(5) Upper orbital bristle shorter than lower ... EP Wa oi em AES HS te inaequalis (Malloch) - Upper orbital bristles equal in length ..... A sites Rae ree ee eee: aequalis Spencer Japanagromyza polygonivora Wiegmann, New SPECIES A small black fly, with light gray tomen- tum, 2.4-—2.7 mm long. Male.— Head: (Fig. 1) frons black, light gold tomentose, approximately equal to width of eye; parafrons black, bare; vertex black, bare, convex; lower orbitals 2 equal; upper orbitals 2 equal; orbital setulae sparse, erect; lunule light gold tomentose, concave, as high as basal width; antenna black, ped- icel with single black dorsoapical seta; Ist flagellomere subspherical, black, gray pi- lose; arista 2 as long as antenna, pubes- cent, with basal % swollen; gena black, gray tomentose, rectangular, 0.20 height of eye; palpus black, gray tomentose with short black hairs; proboscis brown with gold hairs. Thorax: mesonotum black, gray tomen- tose, dorsocentrals 0+2; prescutellars lack- ing; acrostichals irregularly in 9-10 rows, all except median two rows ending at pos- terior dorsocentral; setulae between supra- alar and dorsocentrals in 6—7 irregular rows; supra-alar as long as dorsocentrals; scutel- lum black, lightly gray tomentose; basal scu- tellars 2 x as long as apical scutellars; pleu- ron black, gray tomentose; postpronotal lobe with single long black seta, and several short black setae; notopleuron with 2 equal setae; anepisternum with posterior row of short black setae; katepisternum with single dor- sal row of 4 black setae; katepisternal seta 3x as long as anterior 3; proepisternum with single long black seta; epimeron black, gray tomentose. Legs: black with black setae; coxae with black setae anteriorly; foretibia with one lat- eral black bristle; midtibia with 2 lateral black bristles. Wing: 2.2-2.4 mm long, entirely micro- trichose; costa reaching M1+2 with three sections in proportion of 50:18:10; r-m be- fore middle of discal cell (2:3); ultimate sec- tion of M1+2 as long as penultimate; ul- timate section of M3+4 as long as penultimate (25:25); squama white with dense fringe of gold hairs; halter white. Abdomen: black, lightly gray tomentose, with dense black setae. Terminalia (Figs. 2-5): Epandrium black, with black hairs dorsally, without stout VOLUME 93, NUMBER 1 16KY is Benger cS ae art z nae é i WO: 18MM 65 $:66006 P: 66016 Ly Fig. 9. Japanagromyza polygonivora Wiegmann n. sp. puparium, posterior view SEM. spines; surstylus forked into 2 approxi- mately equal appendages, ventral incurved, dorsal downcurved, with sharp apical spine (Fig. 2); cercus extending to level of surstyli, black with long fine black hairs, with thin apical flange exteriorly, without stout spines; hypandrium (Fig. 4) slightly longer than phallapodeme, without hypandrial apo- deme; aedeagus as in Fig. 3, distiphallus 5 as long as phallapodeme, membranous; dis- tal tubule straight not coiled; endophallus heavily sclerotized dorsally. Female.—Identical to male in all char- acters except slightly larger in some dimen- sions such as frontal width and wing length. Female terminalia (Figs. 6-8): Ovipositor sheath trapezoidal, black, lightly gray to- mentose, as long as or slightly shorter than tergite 6; egg guides as in Fig. 8, with ap- proximately 20 rectangular serrations; cer- cus with 2 long tactile sensilla and scattered long hairs (Fig. 7); spermatheca spherical unmodified (Fig. 6). Host/early stages.—This species makes a blotch mine in the leaves of Polygonum sp. Subsequent collection of the host-plant in- dicates that it is a smartweed in the section Persicaria of the genus Polygonum probably belonging to the species P. densiflorum Meissner. This identification was based on poorly preserved vegetative material, how- ever, and should only be regarded as an ap- proximate species determination. Further collections are needed to obtain a more pre- cise host-plant determination. Puparium: 2.0-3.0 mm long, dark brown, visibly seg- mented; posterior spiracles ringed by 6-8 small projections on widely spaced conical protuberances (Fig. 9). Type material.—Holotype, male, Flori- da, Brevard Co., near Melbourne, 15-V-86, Hibbard-Smith colls. The holotype, 1 male, and 2 female paratypes deposited USNM Washington D.C.; the remainder, | male, 4 females and 5 pupae collected at the same locality, were returned to the Florida State Collection of Arthropods, Gainesville. The above description was made from the 66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON series of specimens due to damage from be- ing preserved in alcohol. Male paratypes were examined for characters missing 1n the holotype. The holotype is missing | upper orbital, 1 lower orbital, 1 flagellum, | dor- socentral, and 2 of 4 scutellars. Diagnosis: This species can be easily dis- tinguished from the other Japanagromyza species from the Caribbean area by the unique appendages of the surstylus (Fig. 2) and the absence of prescutellar bristles. Etymology: The name polygonivora is an adjective based on the larval feeding habit in the leaves of Polygonum sp. DISCUSSION The new species Japanagromyza polygo- nivora represents the first North American agromyzid attacking Polygonum sp. Further collections are needed to obtain an accurate species identification of the host-plant and to help determine the range of hosts in Po- lygonaceae. The close similarities between J. polygonivora and the other members in the genus illustrate the difficulties in making species level identifications in the Agro- myzidae. The male genitalia are crucial in distinguishing between closely related members of this family. J. polygonivora is no exception with its distinctive forked sur- stylus which is unique in the genus. ACKNOWLEDGMENTS I would like to thank Dr. H. V. Weems Jr. and G. C. Steyskal for providing me with the specimens, and D. Miller, F.C. Thomp- son, T. Henry and G. C. Steyskal of the Systematic Entomology Laboratory, USDA for their comments on the manuscript. I also gratefully acknowledge K. Hibbard of the Florida Department of Agriculture, Di- vision of Plant Industry for collecting spec- imens of the host-plant and Bryan Dutton, Department of Botany, University of Mary- land for his assistance with host-plant iden- tification. LITERATURE CITED Griffiths, G. C. D. 1972. The phylogenetic classifi- cation of Diptera Cyclorrhapha, with special ref- erence to the structure of the male postabdomen. Series Entomologica no. 8, 340 pp. Dr. W. Junk N.V., the Hague. McAlpine, J. F. 1981. Morphology and terminolo- gv—Adults. 2, pp. 9-63. Jn McAlpine, J. F. et al., eds., Manual of Nearctic Diptera. Research Branch, Agriculture Canada Monograph 27(1): 1-674. Sasakawa, M. 1954. Neue Agromyzidae aus Japan. Transactions of the Shikoku Entomological So- ciety 4: 35-49. 1958. The female terminalia of the Agro- myzidae with description of a new genus (1). Sci- entific Reports of the Saikyo (Kyoto) University of Agriculture no. 10: 133-145. 1963. Oniental Agromyzidae (Diptera) in Bishop Museum, |. Pacific Insects 5: 25—SO. Spencer, K. A. 1966. New and interesting Agromy- zidae from Florida. Stuttgarter Beitrage zur Na- turkunde 158, 20 pp. 1976. The Agromyzidae (Diptera) of Fen- noscandia and Denmark. Fauna Entomologica Scandinavica, vol. 5, part 1, Scandinavian Science Press, Klampenborg, Denmark, 304 pp. 1977. A revision of the Australian Agro- myzidae (Diptera). West Australian Museum Spe- cial Publication No. 8., 225 pp. Spencer, K. A. and C. E. Stegmaier. 1973. The Agro- myzidae of Florida with a supplement on species from the Caribbean. Arthropods of Florida, vol. 7, 205 pp. Spencer, K. A. and G. C. Steyskal. 1986. Manual of the Agromyzidae (Diptera) of the United States. U.S. Dept. of Agriculture, Agriculture Handbook No. 638, 478 pp. Wiegmann, B. M. 1989. A phylogenetic revision of the family Atelestidae (Diptera: Empidoidea) and its implications for the origin of the cyclorrha- phous Diptera. viii + 205 pp. University of Mary- land, M.Sc. thesis. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 67-69 PARASITE COLONIZATIONS AGAINST CRIOCERIS ASPARAGI (L.) AND C. DUODECIMPUNCTATA (L.) (COLEOPTERA: CHRYSOMELIDAE) IN NORTH AMERICA FROM 1983 TO 1988 R. M. HENDRICKSON, JR., F. GRUBER, G. MAILLOUX, AND J. J. DREA (RMH) Beneficial Insects Research Laboratory, Agricultural Research Service, U.S. Department of Agriculture, 501 South Chapel Street, Newark, Delaware 19713; (FG) European Parasite Laboratory, Agricultural Research Service, U.S. Department of Agri- culture, 13-17 rue de la Masse, 78910 Orgerus-Béhoust, France; (GM) Station de re- cherches de St-Bruno, 335 Est, Chemin des 25, St-Bruno de Montarville, P. Québec, Canada J3V 4P6; (JJD) Beneficial Insects Laboratory, Agricultural Research Service, U.S. Department of Agriculture, Bldg. 476, BARC-East, Beltsville, Maryland 2070S. Abstract. — Three species of parasites were colonized against Crioceris asparagi. Of these Lemophagus crioceritor Aubert (Hymenoptera: Ichneumonidae) is probably established in Canada. Two species of parasites were released against C. duodecimpunctata but none was recovered. Key Words: Insecta, asparagus, biological control, parasite, Lemophagus, Tetrastichus, Meigenia, Diaparsis, Paralispe, Mesochorus Two species of chrysomelid beetles, the asparagus beetle (AB), Crioceris asparagi (L.), and the spotted asparagus beetle (SAB), C. duodecimpunctata (L.), are serious pests of asparagus culture in North America. Both species are exotic; their origin is believed to be Europe. There are 83,000 acres of com- mercial asparagus in the United States with an annual value in excess of $82 million. The acreage is spread over 12 states with the highest production in California, Wash- ington, Oregon, Michigan, and Illinois. Cri- oceris species are serious pests of asparagus in all states except California. In three states, Washington, Michigan, and Illinois, the an- nual loss to beetle feeding damage, resultant market culling, and the cost of partially ef- fective chemicals used in an attempt to con- trol the beetles is between $1.4 and $1.6 million. In New Jersey, there are generally three spray applications per year at a total cost of about $200,000. Even so the beetles remain a problem. The USDA Agricultural Research Service conducted a biological control program against these beetles in North America in the 1980s. This is a summary of the project. MATERIALS AND METHODS To obtain parasites of the beetles, large numbers of late instar C. asparagi larvae were collected from asparagus fields in France, and then held in screen bottom cag- es at the European Parasite Laboratory, then at Sevres, at ambient room conditions. Bou- quets of asparagus ferns were supplied as food. Mature larvae dropped through the screen bottom of the cage and pupated in a mixture of moist sand and vermiculite. To obtain parasites of SAB, large numbers of asparagus berries infested with SAB larvae were collected and held under the same con- 68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ditions. Mature larvae of this species emerged from the berries, dropped to the substrate, and burrowed into it to pupate. Adult and pupal parasites which emerged were sent to the Beneficial Insects Research Laboratory, Newark, Del. Pupae were held in quarantine until emergence of the adult parasites. After clearance from quarantine, parasite species were divided among project researchers directly involved in the project in Delaware, Maryland, and Quebec, Can- ada, and cooperators in Delaware, Maine, Maryland, Missouri, New Jersey, Virginia, and Washington for study and colonization. Field recovery sampling of the coloniza- tion sites consisted of weekly collections of up to several hundred mature larvae, de- pending on the abundance of the beetle lar- vae. AB larvae were held in small cages in the laboratory with bouquets of asparagus ferns for food. Since SAB larvae develop in asparagus berries, 473 ml (1 pint) of fully formed berries, green or red in color, were collected weekly for recoveries of the intro- duced parasites. RESULTS AND DISCUSSION European collections: A total of 145,560 C. asparagi larvae and 136,560 C. duode- cimpunctata larvae were collected in France. From this material, five primary parasites were recovered. They were Diaparsis trun- catus (Gravenhorst) (Hymenoptera: Ich- neumonidae), Lemophagus crioceritor Au- bert (Hymenoptera: Ichneumonidae), Meigenia mutabilis (Fallen) (Diptera: Ta- chinidae), Tetrastichus asparagi Crawford and 7. crioceridis Graham (Hymenoptera: Eulophidae). Of these, two species were new to science: Lemophagus crioceritor was de- scribed by Aubert (1986) and Tetrastichus crioceridis by Graham (1983). The follow- ing is a summary of the parasite species composition and abundance from the total host collections. AB: Tetrastichus asparagi 10.3% Meigenia mutabilis 16.1% Lemophagus crioceritor 7.9% Total 34.3% The hyperparasite, Mesochorus testaceus Gravenhorst (Hymenoptera: Ichneumoni- dae), was recovered from 32.1% of the host insects containing L. crioceritor. SAB: Diaparsis truncatus 20.0% Tetrastichus crioceridis 17.8% Total 37.8% North American colonizations: The total numbers of parasites colonized in the Unit- ed States and Canada were 11,236 (50.2% 2) D. truncatus; 4481 (49.8% °) Lemophagus crioceritor (1600 additional pupae were used for laboratory studies); 8381 (56.8% 9°) M. mutabilis; 34,731 (96.5% 2) T. asparagi; and 86,018 (73.0% 2) T. crioceridis. Recovery of L. crioceritor: A total of 166 L. crioceritor was released at a field adjacent to the Beneficial Insects Research Labora- tory, Newark, Del. in 1984 and 1985. Re- covery attempts were made at this site from 1986 to 1989 and a total of 1566 AB larvae were collected. Of these, 65.0% produced adult beetles and 2.1% produced parasites of three species. A single specimen of L. crioceritor was recovered in 1986, three specimens in 1987, no specimens in 1988 or 1989, although a single live individual was observed on the asparagus plants in July 1988. In Maryland only one adult L. crioceritor was ever recovered from any of the colo- nization fields. That was from the Beltsville Agricultural Research Center in 1986, the year after the initial colonizations. Colonizations were also made by RMH and JJD at 25 other sites in Delaware, Maryland, and New Jersey. The sites were commercial fields subjected to regular in- secticidal treatments. AB larvae were col- lected from 19 of these fields for the next 2 years, but L. crioceritor was only recovered in the year of colonization. At L’Assomption, Quebec, a total of 312 adults of L. crioceritor was colonized in 1985 and 1986. In 1987 parasitization was 8.9% (n = 74). An administrative change in re- VOLUME 93, NUMBER 1 search objectives prevented recovery at- tempts in later years. Harmer et al. (1990), cooperators at the University of Guelph, Ontario, released L. crioceritor in 1987. They found a rate of parasitism of 15.7% for the first generation and 24.8% for the second generation in 1989, which suggests the parasite is probably es- tablished. The probable establishment of L. cri- oceritor in Canada and its probable failure to establish, at least in recoverable numbers, in the Middle Atlantic states suggest that the parasite is favored by cooler northern latitudes. A similar situation was noted in France. The overall percentage parasitiza- tion in the fields studied in France averaged 7.9%. However, in the most northern col- lection sites, the departments of Loiret and Loir-et-Cher, parasitization was 8.6%. In the most southern collection area, the depart- ment of Drome, the parasitization was only 0.6%. Recovery of other parasite species: The gregarious larval parasite, 7. asparagi, was recovered from AB collected from 1986- 1989. However, the overall parasitization was extremely low. Only 0.5% (n = 7) of the hosts from which live forms emerged were parasitized by the eulophid and 31 individ- uals emerged at the Beneficial Insects Re- search Laboratory. Although 7. asparagi was recorded in North America as early as 1863 (Clausen 1978), specimens were colonized to increase the genetic diversity of the par- asite and possibly increase its effectiveness in North America. FG found that 10.3% of the hosts were parasitized by 7. asparagi in France. At L’Assomption, Quebec, the average maximum parasitism by 7. asparagi was 39.1% in the first generation and 49.2% in the second generation for 1980-1987. These figures are very much higher than those found in Delaware. This suggests the par- asite has definite climatic preferences. 69 A native solitary parasite, Paralispe in- fernalis (Townsend) (Diptera: Tachinidae), was recovered from many samples in Mary- land and Delaware, but it produced only 2.1% parasitization (n = 22) at the Benefi- cial Insects Research Laboratory and ap- parently had no controlling impact on the populations of the host. Since no specimens of parasites released against SAB were recovered and only one species, L. crioceritor, is probably estab- lished against AB, the USDA has discon- tinued the introduction program. However, there is still the possibility that Lemophagus will continue to increase in the northern ar- eas of distribution of C. asparagi in North America. ACKNOWLEDGMENTS E. Rey assisted in collections and rearings in France. S. Barth participated in coloni- zations, collections, and rearings in north- eastern U.S. L. Ertle handled quarantine processing and provided sex ratio data. Initial identifications were provided by J. Aubert (1986; Lemophagus) and by M. Gra- ham (1983; 7. crioceridis). Other identifi- cations were provided by R. Carlson (Lem- ophagus and Mesochorus), M. Schauff (Tetrastichus), H. Townes (Diaparsis) and N. Woodley (Tachinidae). The paper was reviewed by M. Rose and D. Williams. LITERATURE CITED Aubert, J. F. 1986. Quatorze Ichneumonides Pétio- lées inédites ou synonymes. Bulletin de la Société Entomologique de Mulhouse, pp. 17-25. Clausen, C. P. 1978. Introduced parasites and pred- ators of arthropod pests and weeds: A world re- view. U.S.D.A. Agriculture Handbook No. 480. Graham, M. W. R. de V. 1983. A new species of Tetrastichus Haliday (Hymenoptera: Eulophidae), a parasite of the asparagus beetle, Crioceris duo- decimpunctata (L.). Entomological Gazette 34: 275-277. Harmer, C. S., M. K. Sears, and J. L. Laing. 1990. Parasitoid of the asparagus beetle. Biocontrol News 3: 34. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 70-85 REVIEW OF AFROTROPICAL BEACH FLIES OF THE TRIBE CANACINI AND SUBFAMILY NOCTICANACINAE (DIPTERA: CANACIDAE) WAYNE N. MATHIS AND AMNON FREIDBERG Department of Entomology, NHB 169, Smithsonian Institution, Washington, D.C. 20560, USA. Department of Zoology, The George S. Wise Faculty for Life Sciences, Tel- Aviv University, Tel-Aviv 69 978, Israel. Abstract. — Afrotropical beach flies of the tribe Canacini and subfamily Nocticanacinae are reviewed. Included are 3 species in Canace Haliday, 6 in Nocticanace Malloch, and 5 species in Procanace Hendel. Four new species are described: Canace zvuy (Limbe, Cameroon), Nocticanace littorea (Takaungu, 50 km N Mombasa, Kenya), Procanace can- zonerii (Limbe and Kribi, Cameroon), and Procanace pninae (Takaungu, 50 km N Mom- basa, Kenya). One synonymy is proposed: Procanace fluvialis Canzoneri, 1987 = Pro- canace grisescens Hendel, 1913. Diagnoses are provided for species groups and genera that include new species to facilitate their recognition. Key Words: taxonomic review Recent collecting in Cameroon and Ke- nya has resulted in specimens of the dip- terous family Canacidae that are new or lit- tle known from the Afrotropical Region. This paper reports the study of this material within the context of a review of the tribe Canacini (subfamily Canacinae) and the subfamily Nocticanacinae from the Afro- tropical Region. All of the new species de- scribed herein belong to genera that are in- cluded in Canacini or Nocticanacinae, and the latter has never been treated compre- hensively except for a listing of species in the recent catalog of Afrotropical Diptera (Cogan 1980). Since publication of this cat- alog, additional Afrotropical species have been added to Canacini (Canzoneri 1981, 1982) and Nocticanacinae (Canzoneri 1987, Canzoneri and Raffone 1987, Mathis 1988b). This treatment of Canacini and Nocticanacinae completes review of all known species of Afrotropical beach flies. Canacidae (Canacini, Nocticanacinae), beach flies, Afrotropical Region, Previously, Mathis (1982a) reviewed the tribe Dynomiellini (subfamily Canacinae), the only other tribe of beach flies known to occur in the Afrotropical Region. For each genus and species that is re- viewed, a synonymy, the known distribu- tion, including detailed information on the type locality, the depository of the primary types, and a remarks section, as appropriate, are provided. In the synonymies, only lit- erature that is pertinent to the Afrotropical Region or the taxons’ nomenclatural history is cited. For new species, a more complete description is provided, including diagnoses of appropriate species groups and genera. The species groups that are recognized for the genera Nocticanace and Procanace are not necessarily monophyletic assemblages and are used primarily to conveniently di- vide the genera into manageable units and to facilitate their identification. Their char- acterization and species composition may VOLUME 93, NUMBER 1 change as a result of revisionary studies. The descriptive format for the new species follows Mathis and Wirth (1979) and Ma- this (1982a, 1988a). More details concern- ing the morphology and higher classifica- tion of Canacidae are found in Mathis (1982a) and Wirth (1987). Two venational ratios are used in the de- scriptions. Costal Vein Ratio: The straight line distance between the apices of R,,, and R,,;/distance between the apices of R, and R,,;. M Vein Ratio: The straight line dis- tance along vein M between crossveins (dm- cu and r-m)/distance apicad of dm-cu. Amnon Freidberg and Fini Kaplan, Tel- Aviv University (TAU), Israel, collected the specimens from Cameroon, Kenya, and Ni- geria, and these collections are deposited in TAU and the National Museum of Natural History (USNM), Smithsonian Institution. Acronyms of other institutions, primarily those where primary types are deposited, are as follows: DCSA (Dipterorum Collec- tionis Strobl, Admont, Austria), BBM (Bernice P. Bishop Museum, Honolulu, Ha- wali), BMNH (British Museum (Natural History), London), HUS (Hokkaido Uni- versity, Sapporo, Japan), MCV (Museo Ci- vico di Storia Naturale di Venezia, Italy), MNHN (Muséum National d’Histoire Na- turelle, Paris, France), NMW (Naturhisto- risches Museum, Wien, Austria), TMC (Transvaal Museum Collection, Pretoria, South Africa), ZMA (Instituut voor Tax- onomische Zoologie, Zoologisch Museum, Universiteit van Amsterdam, Nether- lands). ANNOTATED KEY TO SUBFAMILIES, TRIBES, AND GENERA OF AFROTROPICAL CANACIDAE 1. Lateroclinate fronto-orbital setae 3 and katepi- sternal seta usually present (lacking in the gri- sescens group of Procanace); lamella of ¢ ter- minalia bearing 2 large setae, one apical the other subapical, each rather bluntly rounded (Subfamily Nocticanacinae) — Lateroclinate fronto-orbital setae either 4 or more, or 3 and with katepisternal seta lacking; 71 lamella of 2 terminalia with | large, apical seta, this usually acutely pointed (Subfamily Cana- CHINA C) eee eee ee ee a Wes ait ne aie 3 2. Intrafrontal setae absent, although anterior 13 of frons occasionally with scattered setulae . . BERN BOR TS ee RS Procanace Hendel [6 species in the Afrotropical Region] — One or 2 intrafrontal setae in addition to setu- SORE pote on Nocticanace Malloch [6 species in the Afrotropical Region] lae, if any 3. Lateroclinate fronto-orbital setae 3; katepister- nal seta lacking (Tribe Canacini) ........... EA Gee team en Hoe fed Seo eR Rad 8 eels Canace Haliday [3 species in the Afrotropical Region] — Lateroclinate fronto-orbital setae 4 or more; katepisternal seta sometimes present (Tribe Dynomiellini) 4. Katepisternal seta present, sometimes pale .. SEES Se Lae ake Isocanace Mathis [3 species in the Afrotropical Region; Mathis, 1982a: 11-20] Katepistemaliseta‘absenty...5.5nn ae eee 5 5. Anterior notopleural seta lacking; lateral scu- tellar setae l pain. 4425.20 Trichocanace Wirth [1 species in the Afrotropical Region, T. sinensis Wirth; Madagascar; Mathis, 1982a: 22] — Anterior notopleural seta present; lateral scu- tellarisetacapalls) ene eee ian 6 6. Vein M with last section arcuate; mesofrons uniformly and densely setulose; fronto-orbital setae 4-6; setae in general pale Be ln Ghee Eee a LES oe Xanthocanace Hendel [1 species in the Afrotropical Region, X. capensis Wirth; South Africa; Mathis, 1982a: 24] — Vein M with last section more or less straight, not distinctly arcuate; mesofrons with bare ar- eas, not densely setulose; fronto-orbital setae 4; setae in general dark colored ............ i ta NE he Dynomiella Giordani Soika [4 species in the Afrotropical Region (South Africa or Namibia); Mathis, 1982a: 9-11] SUBFAMILY CANACINAE TRIBE CANACINI GENUS CANACE HALIDAY Canace Haliday, in Curtis, 1837: 281 [pub- lished in synonymy; first made available by use in Haliday, 1839: 411]. Type spe- cies: Ephydra nasica Haliday, 1839, by subsequent monotypy (Haliday, 1839: 411).—Mathis, 1982b: 58 [revision, figs. of heads and ¢ and @ terminalia]. 72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Diagnosis.— Head: Mesofrons with 1-4 marginal, generally proclinate setae; post- ocellar setae, 1 pair, large (subequal in size to ocellar setae), divergent (with same or1- entation as ocellar setae); mesofrons densely microtomentose, usually appearing dull, vestiture not markedly differentiated from that of parafrons; large, lateroclinate, fron- to-orbital setae 3; arista with branching hairs short, not longer than basal aristal width, and extended nearly to tip; large, anaclinate, genal setae 2-4; large, anteroclinate, genal setae 1-2; eye longer than high. Thorax: Dorsocentral setae 4 (1+3); acrostichal setulae generally evident as 2 rows though setulae sometimes weakly de- veloped or with additional setae between acrostichal and dorsocentral setae anteri- orly; prescutellar acrostichal setae well de- veloped; supra-alar setae | (posterior); scu- tellum usually with sparse discal setulae and 2 pairs of lateral setae; notopleural setae 2; anepisternum usually with 2 large setae of subequal length along posterior margin, oc- casionally 3, also with a distinctive, anacli- nate seta inserted near posterodorsal angle; both proepisternum and proepimeron bear- ing a seta; katepisternum lacking a large seta, although usually with some setulae, es- pecially toward ventral margin. Apical sec- tion of vein M straight. Femora dark col- ored; fore femur subequal in size to middle and hind femora, bearing 3—5 long, slender, posteroventral setae, but lacking short, stout, anteroventral setae; hind tibia lacking an anteroventral, spinelike seta apically. Abdomen: Male terminalia: Surstylus in lateral view with fingerlike, ventral projec- tion and frequently with an enlarged, pos- teroventral process. Female genital lamellae slender, terminal setae long, slender, usually acutely pointed. Discussion. — Canace is an Old-World ge- nus and is only known to occur along the coasts of the North Sea and north Atlantic, the Mediterranean Sea, and west Africa (south Atlantic coast). We have collected larvae and reared adults of C. salonitana. The larvae occurred on the protected sides of large rocks along the coast of the Mediterranean Sea in Israel. The rocks, to varying degrees, were covered with algae, which is probably the food source for both larvae and adults. The key includes all known species of Canace, as all but two of these, C. saloni- tana (Mediterranean) and C. actites (Canary and Madeira islands), are now known from the Afrotropical Region. KEY TO SPECIES OF CANACE 1. Gena with 3-4 large, anaclinate setae below eye — Gena with 2 large, anaclinate setae below eye eee A ee ee ean es jatles) 2. Posterior process of surstylus robust, slightly enlarged subapically ........ C. nasica (Haliday) — Posterior process of surstylus generally thinner and gradually becoming narrower toward apex SO OEM oes celbn eae C. rossii Canzoneri 3. Mesofrons with 34 pairs of proclinate setae inserted along margin anterior of median ocel- lus; anterior acrostichal setae well developed, conspicuous; larger species, length 2.5 to 3.3 mm C. salonitana Strobl — Mesofrons with 1-2 pairs of proclinate setae inserted along margin; anterior acrostichal se- tae weak, inconspicuous, smaller species, length 1.5 to 3.1 mm 4. Mesofrons with 2 pairs of proclinate setae, pos- terior seta inserted about in line with posterior ocelli, anterior seta inserted in line with median ocellus; anterior margin of gena bearing 2 large, anteroclinate setae C. zvuy, new species — Mesofrons with | pair of proclinate setae, these inserted in line with median ocellus; anterior margin of gena bearing | large, anteroclinate SOLA) cy. jontuant aie ea ee eee: C. actites Mathis Canace nasica (Haliday) Fig. | Ephydra (Canace) nasica Haliday, 1839: 411. Canace nasica.—Loew, 1860: 29 [generic combination, review].—Frey, 1958: 48 [list, Cape Verde Islands].— Cogan, 1980: 694 [Afrotropical catalog].—Canzoneri, 1981: 216 [list, Senegal]. — Mathis, 1982b: 60 [revision, figs. of head, ¢ and @ ter- minalia]. VOLUME 93, NUMBER 1 Figs. 1, 2. nasica, lateral view; 2, Canace rossii, lateral view. Scale bar equals 0.1 mm. Posterior surstylar process. 1, Canace Distribution. — Afrotropical: Cape Verde Islands (Frey 1958), and Senegal (Dakar, N’Gor; Canzoneri 1981). Palearctic: Coast of the North Sea, Mediterranean, and is- lands of eastern Atlantic Ocean (Azores, Ca- nary Islands, and Madeira Islands). Type material. — Primary types apparent- ly lost (see Mathis 1982b: 60). Remarks.—Specimens from the two Af- rotropical localities noted above (Cape Verde Islands and coast of Senegal) have not been examined, and their identity re- mains unconfirmed. We would not be sur- prised to discover that the Afrotropical specimens are representatives of C. rossi. Comparison of C. nasica (two sites in En- gland) and a paratopotype of C. rossii re- vealed slight differences in the shape of the male terminalia, especially the posterior surstylar process (Figs. 1, 2). This process, in specimens from England (and presum- ably from Ireland, the type locality of this species), is slightly more robust than in spec- imens from Sierra Leone and has a slight, subapical swelling. We also studied speci- mens from Cameroon and found the pos- terior process to be very similar to that from Sierra Leone. Although the differences are slight, they are consistent in the sampling we were able to study, and we have elected to recognize the variation between popu- lations from northern European and those from western Africa as interspecific. Much better sampling from several sites between fs) these rather disjunct localities is needed to test this hypothesis further, specifically whether the variation is intraspecific, per- haps clinal, or whether the differences we found are consistent and represent indepen- dent lineages. Canace rossii Canzoneri Fig. 2 Canace rossii Canzoneri, 1982: 61. Distribution.—Sierra Leone, Western Area, Freetown (Lumley Beach di fronte a Juba.) and Cameroon (Kribi). Type material.—Holotype 6, MCV. Remarks.—See our comments under the ‘““Remarks” section of the previous species. Canace zvuy Mathis and Freidberg, NEw SPECIES Figs. 3-12 Description. — Small to moderately small beach flies, length 1.80 to 2.20 mm; mostly dark colored, brown to grayish brown. Head (Figs. 3-8): Eye with oblique ori- entation; anterodorsal portion of head, near juncture with antenna, less protrudent or angulate; frons with mesofrons and fronto- orbits similar, dark brown with some bronz- ish tinges, contrasted with charcoal-colored, dull parafrons; frons bearing 2 large, mar- ginal setae and | anterior setula (less than '’) length of larger setae), 2 larger setae in- serted more or less in line with posterior ocelli (posterior seta) and median ocellus (anterior seta), setula inserted between an- terior margin of frons and median ocellus; gena with 2 subequal, anaclinate setae just below the eye and 2 anteroclinate setae to- ward the anterior margin, one seta inserted at anterior margin, the other slightly pos- teroventrad; face and gena silvery gray. Thorax (Figs. 9-11): Acrostichal setulae sparse and mostly weakly developed, oc- casionally with a few larger setulae; disc of scutellum either bare of setulae or with |- 2. Wing with costal vein ratio averaging (0). 10; vein M ratio averaging 0.62. 74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 3-11. Scanning electron stereomicrographs of Canace zvuy. 3, Head, lateral view; 4, Gena and setae, lateral view; 5, Antenna, lateral view; 6, Frons, dorsoblique view; 7, Ocellar triangle and setae, dorsoblique view; 8, Fronto-orbits and setae, dorsal view; 9, Mesonotum, dorsal view; 10, Notopleuron and setae, lateral view; 11, Katepisternum and setulae, lateral view. Abdomen: Male terminalia (Fig. 12): Sur- stylus broadly L-shaped and with an an- teroventral, fingerlike process; anterior margin of surstylus just below midheight bearing several longer setulae. Type material—The holotype male is la- beled ““CAMEROON Limbe (shore) 14— 15.XI[Nov].1987 A. FREIDBERG.” The allotype female and paratypes (5 3, 14 2; TAU, USNM) bear the same locality data VOLUME 93, NUMBER 1 as the holotype; Fini Kaplan, Tel-Aviv Uni- versity, was the collector of six of the female paratypes. The holotype is double mounted (minute nadel in a polyporus block), is in excellent condition, and is deposited in the USNM. Natural history.— The type series was col- lected along the coast of Cameroon at a site just west of Limbe. The site was a polluted (tar), sandy beach with large, flat rocks among small, intertidal pools. The large rocks extended onto the sandy shore. The flies were collected by sweeping an aerial net against or just above the rocks. Etymology.—The specific epithet, zvuy, is a noun in apposition and is the transliter- ation of a Hebrew noun from the Bible meaning fly and is so named to recognize the collecting efforts of Israeli entomolo- gists, who collected the type series. Remarks.— Externally this species is sim- ilar to C. actites and C. salonitana but differs in the number and orientation of setae on the mesofrons and gena. The shape of the male terminalia also differs, especially that of the surstylus, which is broadly L-shaped as in C. salonitana but bears an anteroven- tral, rather sharply pointed process (the pro- cess is bluntly rounded in C. salonitana and has a more ventral orientation). SUBFAMILY NOCTICANACINAE GENUS NOCTICANACE MALLOCH Nocticanace Malloch, 1933: 4. Type spe- cies: N. peculiaris Malloch, by original designation.— Mathis, 1989: 594 [key to species groups]. Diagnosis.—Small to medium-sized beach flies, length 1.8 to 3.7 mm; general coloration grayish black. Head: Intrafrontal setae | pair; postocellar setae either absent or much reduced, less than % length of ocellar setae; ocelli arranged to form an isosceles triangle, distance between posterior ocelli greater than that between ei- ther posterior ocellus and the anterior ocel- lus. Lower facial margin sinuate; clypeus low, 75 Fig. 12. Canace zyuy. Male terminalia (epandrium, cerci, and surstylus), lateral view. Scale bar equals 0.1 mm. width subequal to length of antenna. Two large anaclinate genal setae; anteroclinate genal seta moderately well developed, at least > length of larger anaclinate genal setae. Pal- pus grayish black, bearing | to several long setae, each seta 2 to 3 times as long as greatest width of palpus. Thorax: Anepisternum with scattered setulae; proepisternal seta absent; katepis- ternal seta present, well developed. Legs en- tirely dark colored, grayish black; fore fe- mur bearing 4-6 long and evenly spaced setae along posteroventral margin, length of setae at least equal to and usually greater than width of femur. Discussion.—The species-group key, which follows, is essentially that of Mathis (1989) but with minor changes. In preparing the key, one of us (WNM) examined most species of the genus. The key, however, should still be considered preliminary, and we invite the critique and comment of users. The Afrotropical species all belong to the pacifica group, which is distinguished from other species groups by the characters out- lined in the species-group key and diagnosis that follow. ANNOTATED Key TO SPECIES GROUPS OF THE GENUS NOCTICANACE 1. Anterior notopleural seta absent — Anterior notopleural seta present ........... 3 76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2. Apical scutellar setae distinctly curved anacli- Nate aie See EI ee: the pacifica group {20 species; Old World, Pacific and Indian Oceans, especially Oceania] — Apical scutellar setae straight to very slightly curved the texensis group [3 species; Caribbean, Gulf of Mexico, Atlantic coast of southeastern United States; revised by Mathis 1989: 594-599] 3. Length of apical section of vein CuA, twice or more length of crossvein dm-cu ........... PLN See Pele ere es the galapagos group [8 species; Galapagos Islands and southwestern North America] — Length of apical section of vein CuA, subequal to length of crossvein dm-cu 4. Apical scutellar setae distinctly anaclinate REX seg Acie MON Ne ee oe ee cuteit de the ashlocki group [1 species, N. ashlocki Wirth; Galapagos Islands] — Apical scutellar setae not anaclinate ........ RS ST ee en wee x Rs the chilensis group [1 species, N. chilensis (Cresson); Chile (there are numerous undescribed species in this group)]} THE PACIFICA GROUP Diagnosis.—Coloration generally dark, grayish brown to grayish black but with ex- ceptions (N. flavipalpis and N. littorea) that are lighter, with some tan coloration on the body and legs extensively yellowish. Thorax: Acrostichal setulae absent; apical scutellar setae distinctly anaclinate; anterior notopleural seta absent; proepisternal seta(e) present; anepisternum with scattered setu- lae; katepisternal seta present. Legs usually entirely dark, grayish brown to black (N. flavipalpis and N. littorea are exceptions with legs that are mostly yellowish); fore femur with 4-6 long and evenly spaced setae along posteroventral margin, length greater than width of femur; midfemur of male lacking a comblike row of setae; hindtibia lacking spinelike setae apically. Wing with length of apical section of vein CuA, long, about twice length of crossvein dm-cu; vein M index 0.44. KEY TO AFROTROPICAL SPECIES OF THE PACIFICA GROUP OF NOCTICANACE 1. Tarsi, apices of tibiae, femora, and palpus yel- LOWS en See eee eee eee eee 2, — Legs and palpus entirely dark colored, mostly gravato blackishverayiy es seer 3 2. Posterior surstylar lobe less broadly developed, median portion narrower than digitiform apex Si SON OH N. flavipalpis Mathis and Wirth — Posterior surstylar lobe more broadly devel- oped (Fig. 13), median portion wider than dig- ItifOnMApe Xe ee ee N. littorea, new species 3. Granitic Seychelles N. mahensis (Lamb) — Malagasy subregion, Aldabra, Kenya, and South Africa 4. Dark coloration of scutum extended ventrally no farther than dorsal half of notopleuron, ven- tral: portion whitishorays eee eee + SAREE toh AO be? N. cyclura Mathis and Wirth — Dark grayish-brown coloration of scutum ex- tended ventrally at least to dorsal half of anepi- sternum 5. Surstylus emarginate ventrally, not deeply cleft, single lobe becoming wider ventrally to subapi- cal widest point, thereafter narrowed gradually to broad, emarginate, ventral margin N. actites Mathis and Wirth — Surstylus deeply cleft ventrally, posterior lobe slender, digitiform, width less than half ante- rior lobe, pointed apically; anterior lobe broad, parallel sided, broadly rounded apically N. caffraria (Cresson) Nocticanace actites Mathis and Wirth Nocticanace actites Mathis and Wirth, 1979: 790.— Mathis, 1988b: 24 [list, Aldabra (? identification)]. Distribution. — Madagascar. Fenerive (on beach), Est: dct Mananara, Ivontaka (15 m). Seychelles. Aldabra (Malabar: Passe Hou- areau; Picard: Settlement). Type material.— Holotype 6, MNHN. Nocticanace caffraria (Cresson) Canaceoides caffraria Cresson, 1934: 222. Nocticanace caffraria.—Wirth, 1951: 273 [revision, generic combination]; 1956: 51 [review].—Cogan, 1980: 694 [Afrotropi- cal catalog]. Distribution. —South Africa. Cape Prov- VOLUME 93, NUMBER 1 ince: East London, Plettenberg. Natal Prov- ince: Umkomaas (beach). Type material.— Holotype 6, TMC. Nocticanace cyclura Mathis and Wirth Nocticanace cyclura Mathis and Wirth, 1979: 791. Distribution. — Madagascar. Sud-Est: dct Fort-Dauphin, Sainte-Luce (10 m); Fene- rive (on beach). Type material.—Holotype 6, MNHN. Nocticanace flavipalpis Mathis and Wirth Nocticanace flavipalpis Mathis and Wirth, 1979: 792.—Mathis, 1988b: 23 [list, Al- dabra, Cosmoledo]. Distribution.— Madagascar. Est: dct Mananara, Ivontaka (15 m); Seychelles: Al- dabra (Grande Terre: Dune Jean-Luis), Cosmoledo (Menai Island: Station). Type material.— Holotype 6, MNHN. Nocticanace littorea Mathis and Freidberg, NEw SPECIES Fig. 13 Description.—A small, light-colored, beach-fly species of the pacifica group (see key to species groups), length 1.7 to 1.9 mm. Head: Grayish coloration of face and gena lighter, mostly whitish gray. Palpus yellow- ish. Thorax: Brown coloration of mesonotum extended laterally and ventrally to about dorsum of notopleuron, thereafter gradually becoming more whitish gray with some very faint greenish tinges. Pleural areas mostly whitish gray. Legs concolorous, mostly yel- lowish; dorsum of femur and to a lesser ex- tent tibia somewhat microtomentose, light- ly grayish. Abdomen: Dorsum mostly grayish; me- dian portion of each tergum with some brownish-purple coloration, lateral margins often faintly bluish gray. Male terminalia 77 Fig. 13. Nocticanace littorea. Male terminalia (epandrium, cerci, and surstylus), lateral view. Scale bar equals 0.1 mm. (Fig. 13): Surstylus deeply cleft ventrally, with a distinct anterior and posterior lobe; anterior lobe moderately slender and long, medial surface bearing numerous, short, stout setae on ventral half; posterior lobe with median portion conspicuously swollen before much more narrowed apex, antero- ventral margin, especially medial aspect, bearing 5-6 stout setae. Type material.—Holotype ¢ is labeled “KENYA Takaungu, 50 Km North Mom- basa 3. XII. 1989 [3 Dec 1989] A. FREID- BERG & FINI KAPLAN.” Five paratypes (2 6, 3 2; USNM) bear the same label data as the holotype. The holotype is double mounted (minute nadel in a plastic block), is in good condition (slightly teneral), and is deposited in the USNM. Natural history.—The type locality is a wooded estuary, known by the inhabitants of the village as ‘““The Creek,” which is drained of sea water during low tide. During high tide the shore is a mixture of sand with small stones. Beach and shore flies were abundant along the high-water line and on an artificial wall, which was nearby. Etymology.—The species epithet, /itto- 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rea, 1s a Latin adjective, meaning “‘pertain- ing to the seashore,” and its usage here al- ludes to the habitat where this species was collected. Remarks.—This species 1s very similar to N. flavipalpis and the two are distinguished from other Afrotropical congeners by the yellowish palpus, front trochanter, tarsi and apex of femora, particularly the ventral sur- face of the latter; brown coloration of meso- notum extended lateroventrally no farther than dorsal angle of notopleuron; surstylus deeply cleft ventrally, forming a distinct an- terior and posterior lobe; posterior lobe, in lateral view with apex digitiform; anterior lobe in lateral view more or less parallel sided ventrally, median surface bearing nu- merous spinelike setulae. This species is dis- tinguished from C. flavipalpis by the shape of the surstylar lobes: posterior lobe more broadly developed in lateral view, appear- ing conspicuously swollen subapically; an- terior lobe more slender, appearing longer. Nocticanace mahensis (Lamb) Canace mahensis Lamb, 1912: 328. Nocticanace mahensis.—Wirth, 1951: 274 [generic combination].—Cogan, 1980: 694 [catalog].—Mathis, 1982c: 423 [re- view, figs. of 6 terminalia].— Mathis, 1988b: 23 [list, Mahé]. Distribution.—Seychelles. Mahé: Long Island; Anse aux Pins, Anse Royale Bay, Anse Takamaka, Bel Ombre, Coevity, Machabee, Port Launay. Type material.— Holotype °, BMNH. Genus Procanace Hendel Procanace Hendel, 1913: 93. Type species: Procanace grisescens Hendel, by original designation].— Mathis and Wirth, 1979: 794 [diagnosis and discussion].— Mathis, 1988a: 330 [diagnosis and key to species groups]. Diagnosis.—General coloration whitish gray, olivaceous, to blackish brown. Head: \ntrafrontal setae absent, but with a few setulae inserted anteriorly; fronto-or- bital setae 3; ocelli arranged to form equi- lateral or isosceles triangle, if isosceles, the greater distance is between posterior ocelli. Arista bearing tiny hairs over entire length, appearing macropubescent. Large anacli- nate genal setae 2; anteroclinate genal seta moderately well developed. Palpus not bearing long setae. Lower facial margin, in lateral view, more or less horizontal. Thorax: Acrostichal setae usually lacking (setulae present in species of the williamsi group); scutellar disc lacking setae (1-2 pairs of scutellar disc setulae occur in P. naka- zatoi Miyagi of the williamsi group); mar- ginal scutellar setae 2, apical pair not ana- clinate; supra-alar seta | (posterior); anterior and posterior notopleural setae present, length subequal; anepisternum with scat- tered setulae and | large seta along the pos- terior margin, lacking a prominent, anacli- nate seta inserted near posterodorsal angle. Katepisternal seta usually present (lacking in species of the grisescens group). Apical section of vein M straight. Femora pale col- ored (dark colored in some species groups); fore femur subequal in size to middle and hind femora, lacking posteroventral setae; hind tibia lacking spinelike setae apically. Abdomen: Male terminalia as follows: Epandrium in posterior view wider than high; cerci reduced, poorly sclerotized; sur- stylus with an anterior and posterior lobe, the latter larger, sometimes markedly so and shape unique to species. Discussion.— The Afrotropical species of Procanace are assigned to two species groups: the fu/va group, with two species, and the grisescens group, with three species. These two groups are distinguished by the characters outlined in the species-group key and diagnosis, which follow. ANNOTATED KEY TO SPECIES GROUPS OF PROCANACE HENDEL 1. Katepisternal seta absent ....the grisescens group [5 species; Oriental, eastern Palearctic, Oceanian, Malagasy, Kenya, Seychelles] VOLUME 93, NUMBER | —aukatepistemalisetaspresent pane ae ace 2, 2. Clypeus high, width about twice height; palpus blackish brown; proepisternal seta absent ... Sa oe hg Hee eee eens ee eae the nigroviridis group [7 species; Hawaiian Islands] — Clypeus low, width at least 4 times height; pal- pus yellowish; proepisternal seta(e) present... 3 3. Acrostichal setulae present, in 2 irregular rows Sty ete ea StI CRA treo the williamsi group [4 species; Hawaiian and Ryukyu Islands] — Acrostichal setulae absent .................. 4 4. Postocellar setae either absent or much re- GUCCU RR rsa ost i ee the fulva group [11 species; Oriental, eastern Palearctic, and Afrotropical] — Postocellar setae present, subequal to length of OCEIIATESEAN s.r onickontk eee the cressoni group [3 species; Oriental, Nearctic] THE FULVA GROUP Diagnosis.—Generally whitish oliva- ceous to brown. Head: Postocellar setae either absent or much reduced; ocelli arranged to form equi- lateral triangle. Palpus yellow. Clypeus low, height '4 to %4 width. Thorax: Acrostichal setulae absent; pro- episternal seta(e) present; anepisternum with scattered setulae; katepisternal seta present. Femora and tibiae mostly yellowish but with considerable grayish microtomentum; tarsi yellowish, becoming darker apically; fore fe- mur lacking long, evenly spaced setae along posteroventral margin, length of setae that are present equal or less than width of fe- mur; middle femur of ¢ bearing comblike sparse row of setae. Wing with apical section of vein CuA, short, subequal to or slightly longer than crossvein dm-cu; vein M index averaging 0.57. Discussion.—The fu/va group, as pres- ently characterized, comprises 11 species that occur on beaches around and within the Pacific and now the Indian and Atlantic Oceans. The specimens from Cameroon and Kenya represent the first known departures from the Australasian and Oceanian Regions. Disjunct distributions sometimes indicate that a taxon may be paraphyletic, a possibility that should be considered when this group is studied further. 79 KEY TO AFROTROPICAL SPECIES OF THE FULVA GROUP 1. Legs entirely yellow except for apical 1-2 tar- someres; male terminalia as in Figs. 25-26 .. BA ec MUIR, eee P. pninae, new species — Legs mostly yellowish but with some oliva- ceous coloration; male terminalia as in Figs. DSH) Ae et eee te P. canzonerii, new species Procanace canzonerii Mathis and Freidberg, New SPECIES Figs. 14-24 Description.—Small to moderately small beach flies, length 1.90 to 2.40 mm; thorax and head generally brown to grayish brown dorsally, becoming lighter ventrally, abdo- men mostly gray to blackish gray. Head (Figs. 14-19): Frons lacking a dif- ferentiated mesofrons and parafrons, most- ly appearing dull, grayish charcoal, with some brownish coloration around anterior margin of ocellar triangle and at anterior margin; 2—6 setulae inserted between ocellar triangle and anterior margin; postocellar setae much reduced, at most 4 length of ocellar setae; fronto-orbits slightly more grayish, becoming whitish anterior of fron- to-orbital setae. Scape and pedicel blackish; Ist flagellomere mostly blackish but dis- tinctly yellowish orange basomedially. Face and gena mostly whitish gray; parafacials with some brownish to yellowish brown col- oration adjacent to eye. Gena with 2 large, anaclinate setae and | large, anteroclinate seta; eye-to-cheek ratio averaging 0.35. Thorax (Figs. 20-22): Third dorsocentral seta inserted slightly medially from align- ment of 2nd and 4th setae. Legs mostly yel- lowish; femora and tibiae with whitish gray microtomentum invested on anterior and dorsal surfaces; apical 2—3 tarsomeres be- coming darker brown; middle femur of male with 4—5 black setae inserted posteroven- trally about '4 from apex. Wing with costal vein ratio averaging 0.21; vein M ratio av- eraging 0.56. Abdomen: Male terminalia (Figs. 23-24): 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 14-22. Scanning electron micrographs of Procanace canzonerii. 14, Head, lateral view, 15, Gena and setae, lateral view; 16, Antenna, lateral view; 17, Frons, dorsoblique view; 18, Ocellar triangle and setae, dorsoblique view; 19, Fronto-orbits and setae, dorsal view; 20, Mesonotum, dorsal view; 21, Notopleuron and setae, lateral view; 22, Katepisternum and setulae, lateral view. Posterior surstylar lobe much larger than width of posterior lobe, but slightly more anterior lobe, bluntly rounded ventrally and than '2 length, apex bearing several setulae. bearing numerous setulae, many of which, Type material.— The holotype male is la- including those at distal half of lobe, longer beled ““CAMEROON Limbe (shore) 14— than lobe’s width; anterior lobe less than % 15.XI.[NOV]1987 A. FREIDBERG.” The VOLUME 93, NUMBER 1 81 Figs. 23, 24. Male terminalia of Procanace canzonerii. 23, Epandrium, cerci, and surstyli, posterior view; 24, Epandrium, cercus, and anterior and posterior surstylar processes, lateral view. Scale bar equals 0.1 mm. allotype female and 66 paratypes (25 4, 41 2; TAU, USNM) bear the same locality data as the holotype (12 specimens were collect- ed by Fini Kaplan). An additional 32 para- types (15 8, 17 2; TAU, USNM) are from: Cameroon. Kribi (beach), route N7, 28-29 Nov 1987, A. Freidberg, F. Kaplan. The holotype is double mounted (minute nadel in polyporus block), is in excellent condi- tion, and is deposited in the Smithsonian Institution (USNM). Natural history.—The specific habitat at the type locality comprised fist-sized cobble at the mouth of a river, which was in the middle of Limbe. The cobble was largely bare, perhaps scoured by floods, but some algae was found on an adjacent site down- stream. Although the cobble habitat ap- peared relatively sterile, the flies were abun- dant. At Kribi, the habitat comprised a narrow sand beach, which was margined by J[po- moea sp. (Ipomoeaceae), with occasional concentrations of intertidal rocks, which was where most of the flies were collected. Etymology.—The specific epithet, can- zonerii, is a patronym to honor Mr. Silvano Canzoneri, who has contributed much to our understanding of beach flies from the Afrotropical Region. Remarks.—The only other Afrotropical species of the fu/va group is the next species, which is also new. This species is distin- guished by the slightly darker legs and the shape of the male terminalia, especially the more bluntly rounded posterior surstylar process (lateral and posterior views). Procanace pninae Mathis and Freidberg, NEw SPECIES Figs; 25,26 Description.—Small beach flies, length 1.65 mm; thorax and head generally brown to grayish brown dorsally, becoming lighter ventrally, abdomen mostly gray to blackish gray. Head: Frons lacking a well-differentiated mesofrons and parafrons, mostly microto- mentose, appearing dull, gray, with some yellow to brownish coloration at anterior 4; 2-6 setulae inserted between oceliar triangle and anterior margin; postocellar setae much reduced, at most '4 length of ocellar setae; fronto-orbits slightly lighter, more whitish, becoming orangish at level of ptilinal su- ture. Scape and pedicel blackish; Ist flagel- lomere mostly blackish but distinctly yel- lowish orange basomedially. Face and gena mostly whitish gray; parafacial with consid- 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs: 25,, 26. Male terminalia of Procanace pninae. 25, Epandrium, cerci, and surstyli, posterior view; 26, Epandrium, cercus, and anterior and posterior surstylar processes, lateral view. Figs. 27, 28. Male terminalia of Procanace cogani. 27, Epandrium, cerci, and surstyli, posterior view; 28, Epandrium, cercus, and anterior and posterior surstylar processes, lateral view. Scale bar equals 0.1 mm. erable orangish to yellowish brown colora- tion adjacent to eye. Gena with 2 large, ana- clinate setae and | large, anteroclinate seta; eye-to-cheek ratio averaging 0.32. Thorax: Third dorsocentral seta inserted slightly mediad from alignment of 2nd and 4th setae. Legs mostly yellowish; femora and tibiae with whitish microtomentum thinly invested on anterior and dorsal surfaces; apical 2—3 tarsomeres becoming darker brown; middle femur of male with 3—4 black setae inserted posteroventrally about '3 from apex. Wing with costal vein ratio 0.22; vein M ratio 0.60. Abdomen: Male terminalia (Figs. 25, 26): Posterior surstylar lobe much larger than anterior lobe, narrowed towards venter and bearing numerous setulae; anterior lobe less than ' width of posterior lobe, but slightly more than 34 length in lateral view, apex slightly swollen and bearing several setulae. Type material. — The holotype male is la- beled ‘““KENYA Takaungu, 50 Km North Mombasa 3. XII. 1989 [3 Dec 1989] A. FREIDBERG & FINI KAPLAN.” The ho- lotype is double mounted (minute nadel in plastic block), is in good condition (the ab- domen has been removed and dissected, the parts are in an attached microvial, the legs and wing on the right side have been re- VOLUME 93, NUMBER | moved and are glued to the plastic block), and is deposited in the Smithsonian Insti- tution (USNM). Natural history.—See comments under Nocticanace littorea. Etymology.—The species epithet, pninae, is a genitive patronym to recognize the sup- port of Pnina Freidberg to her husband’s study of Diptera, especially his field work in Africa and elsewhere. Remarks.—The lighter colored legs, en- tirely yellowish with a thin investment of whitish microtomentum on the dorsum of the femora, and the shape of the male ter- minalia, especially the posterior surstylar process (lateral and posterior views), distin- guish this species from congeners. THE GRISESCENS GROUP Remarks.—Three species of the grises- cens group are known to occur in the Af- rotropical Region, and the characters to dis- tinguish them are primarily the shape of the male terminalia. These differences are often subtle and difficult to use without access to validly determined specimens for direct comparison. KEY TO THE AFROTROPICAL SPECIES OF THE GRISESCENS GROUP (MALES ONLY) 1. Anterior surstylar process distinctly curved posteriorly in lateral view; posterior surstylar process with apical 3 conspicuously tapered, pointed apically; median margin of epandrium bearing several stout setae .... P. cogani Mathis — Anterior surstylar process straight; posterior surstylar process broadly produced apically, if apical 3 tapered, gradual; median margin of epandrium with at most small setulae, no large SOLACE Ce ee aceon fears aishopetameres <3 2 2. Posterior surstylar process with apex distinctly curved mediad (best seen in posterior view); in lateral view this process is slightly angulate sub=basally 9 995.26 eee P. grisescens Hendel — Posterior surstylar process with general ori- entation of apex a continuation of basal por- tion, not distinctly curved medially, in lateral view this process tapered to pointed apex ... Sieh eee A RINE LOND EP: P. pauliani Mathis and Wirth 83 Procanace cogani Mathis Figs. 27, 28 Procanace cogani Mathis, 1988b: 24 [figs. of 6 terminalia]. Distribution. —Seychelles. Mahé: Anse Aux Pins (beach). Type material.— Holotype ¢, USNM. Procanace grisescens Hendel Figs. 29, 30 Procanace grisescens Hendel, 1913: 93.— Wirth, 1951: 258 [revision, figs. of ¢ ter- minalia].—Muiyagi, 1965: 96 [revision, figs. of 6 and 2 terminalia]; 1973: 82 [list]. —Delfinado, 1970: 528 [list, New Guinea, figs. of and 2 terminalia].—Can- zoneri and Raffone, 1987: 75; Kenya. Ukunda. Procanace fluvialis Canzoneri, 1987: 95 [habitus fig.]. NEw SYNONYM. Distribution.— Widespread in the Old World tropics. Afrotropical: Kenya, Liberia (Suakoko), Nigeria (Ile-Ife, Zaria), Sierra Leone (Bathurst), Sudan (Khartoum, Nile River), Zaire (Yangambi). Australasian/ Oceanian: Papua New Guinea, Yap Islands. Oriental: Bangladesh, Japan (Ryukyu Is- lands), Malaysia, Nepal, Taiwan (Anping, Tainan), Thailand. Type material.—Syntypes (many) 42 are from Taiwan: Anping (September), and are deposited in NMW. The holotype ¢ of P. fluvialis is from Su- dan: Khartoum, Nile River, 2 December 1980, Rallo and is deposited in MCV. Natural history.—This is one of the few species of Canacidae that occurs in fresh- water habitats, perhaps exclusively (all re- corded localities for this species have not been checked). Remarks.— We directly compared speci- mens of P. grisescens that were collected on the Sauter expedition to Taiwan with the holotype and other specimens of P. fluvialis and could not find differences to distinguish them. We thus feel that the specimens stud- 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 29, 30. Male terminalia of Procanace grisescens. 29, Epandrium, cerci, and surstyli, posterior view; 30, Epandrium, cercus, and anterior and posterior surstylar processes, lateral view. Figs. 31, 32. Male terminalia of Procanace pauliani. 31, Epandrium, cerci, and surstyli, posterior view; 32, Epandrium, cercus, and anterior and posterior surstylar processes, lateral view. Scale bar equals 0.1 mm. ied are conspecific, hence our synonymy of the two names. Procanace pauliani Mathis and Wirth Figs. 31, 32 Procanace pauliani Mathis and Wirth, 1979: 794. Distribution.— Madagascar. Sud-Ouest: dct Tuléar, Saint-Augustin (6 m). Type material.— Holotype 6, MNHN. ACKNOWLEDGMENTS Special thanks are extended to Fini Kap- lan who made special effort to collect beach flies from Cameroon, Kenya, and Nigeria and graciously made the specimens avail- able to our study. We are grateful to Silvano Canzoneri (MCV) who kindly exchanged or loaned specimens, especially of species that he has described from the Afrotropical Re- gion. We also thank Jon K. Gelhaus and Allen Norrbom for critically reviewing an earlier draft of this manuscript and sug- gesting improvements. The illustrations were skillfully inked by Ms. Elaine R. S. Hodges. Many specimens were collected as part of field work on the Seychelles and Al- dabra, and we gratefully acknowledge fund- ing from the Smithsonian Institution and VOLUME 93, NUMBER 1 the guidance of Brian Kensley, who led our expedition there. We are also grateful for the cooperation of the Seychelles Islands Foundation for facilitating our field work. LITERATURE CITED Canzoneri, S. 1981. Ricerche condotte dal prof. A. Giordani Soika nel Senegal ed in Gambia (Diptera: Ephydridae e Canaceidae). Bollettino del Museo civico di Storia Naturale di Venezia 31[1980]: 201- DON 1982. Ephydridae e Canaceidae della Sierra Leone (Diptera). 7n Ricerche biologiche in Sierra Leone. Academia Nazionale dei Lincei 255: 53- 62. 1987. Sugli Ephydridae e Canacidae del Su- dan (Diptera, Cyclorrhapha. Bollettino del Museo civico di Storia Naturale di Venezia 37: 79-97. Canzoneri, S. and G. Raffone. 1987. Ditteri raccolti dal Dr. Walter Rossi in Kenya (Ephydridae, Can- acidae). Bollettino del Museo civico di Storia Na- turale di Venezia 37: 57-76. Cogan, B. H. 1980. 79. Family Canacidae, p. 694. Jn Crosskey, R. W., ed., Catalog of the Diptera of the Afrotropical Region. British Museum (Natural History). London. Cresson, E. T., Jr. 1934. Descriptions of New Genera and Species of the Dipterous Family Ephydridae, XI. Transactions of the American Entomological Society 60: 199-222. Curtis, J. 1837. A guide to an arrangement of British insects; being a catalogue of all the named species hitherto discovered in Great Britain and Ireland. 2nd ed., 294 pp. London. Delfinado, M.D. 1970. The species of the genus Pro- canace in New Guinea (Diptera: Canaceidae). Pro- ceedings of the Hawaiian Entomological Society 20(3): 527-531. Frey, R. 1958. Zur Kenntnis der Diptera brachycera p. p. der Kapverdischen Inseln. Societas Scientia- rum Fennica. Commentationes Biologicae 18(4): 1-61. Haliday, A. H. 1839. Remarks on the generic distri- bution of the British Hydromyzidae (Diptera). An- nals of Natural History 3: 217-224, 401-411. Hendel, F. 1913. H. Sauter’s Formosa-Ausbeute: Acalyptrate Musciden (Dipt.). II. Supplementa Entomologica 2: 77-112. Lamb, C. G. 1912. The Percy Sladen Trust Expedi- tion to the Indian Ocean in 1905, under the lead- ership of Mr. J. Stanley Gardiner, M.A. Vol. IV. Number XIX. Diptera: Lonchaeidae, Sapromy- 85 zidae, Ephydridae, Chloropidae, Agromyzidae. Transaction of the Linnean Society of London 15: 303-348. Loew, H. 1860. Neue Beitrage zur Kenntniss der Dip- teren. Siebenter Beitrag, pp. 1-46. Berlin. Malloch, J. R. 1933. Some acalyptrate Diptera from the Marquesas Islands. Bernice P. Bishop Muse- um, Bulletin 114: 3-31. Mathis, Wayne N. 1982a. Studies of Canacidae (Dip- tera), I: Suprageneric revision of the family, with revisions of new tribe Dynomiellini and new genus Isocanace. Smithsonian Contributions to Zoology 347: 1-29. 1982b. Canacidae of Israel, with a review of the palaearctic species of the genus Canace Hali- day (Diptera). Entomologica Scandinavica 13: 57- 66. 1982c. Description of a new species of Noc- ticanace Malloch (Diptera: Canacidae) from Sri Lanka, with notes on two related species. Pro- ceedings of the Entomological Society of Wash- ington 84(3): 421-425. 1988a. First record of the genus Procanace Hendel from North America, with the description of a new species (Diptera: Canacidae). Proceedings of the Entomological Society of Washington 90(3): 329-333. 1988b. Beach flies of the Republic of Sey- chelles (Diptera: Canacidae). Bulletin of the Bio- logical Society of Washington 8: 22-29. 1989. A review of the beach flies of the Ca- ribbean and Gulf of Mexico (Diptera: Canacidae). Proceedings of the Biological Society of Washing- ton 102(3): 590-608. Mathis, Wayne N., and W. W. Wirth. 1979. Beach flies of Madagascar (Diptera: Canacidae). Annals of the Natal Museum 23(3): 785-796. Miyagi, I. 1965. Onthe marine shore flies of the genus Nocticanace from Japan (Diptera: Canaceidae). Kontya 33(3): 299-303. 1973. Occurrence of three species of Cana- ceidae in the Philippines (Diptera: Canaceidae). Kontyt 41(1): 82. Wirth, W. W. 1951. A revision of the dipterous fam- ily Canaceidae. Occasional Papers of the Bernice P. Bishop Museum 20(14): 245-275. 1956. New species and records of South Af- rican Canaceidae (Diptera). Journal of the Ento- mological Society of South Africa 19(1): 47-51. 1987. Canacidae. 102 [chapter], pp. 1079- 1083. In McAlpine, J. F., ed., Manual of Nearctic Diptera. Vol. 2. Monograph No. 28, 1332 pp., Research Branch, Agriculture Canada. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 86-91 KARYOTYPE AND NOTES ON THE MALE REPRODUCTIVE SYSTEM AND NATURAL HISTORY OF THE HARVESTMAN VONONES SA YI (SIMON) (OPILIONES, COSMETIDAE) JAMES C. COKENDOLPHER AND STANLEY R. JONES (JCC) 2007 29th Street, Lubbock, Texas 79411; (SRJ) 102 Cathy Street, Elizabethtown, Kentucky 42701. Abstract.—The Texas Platycynorta transversalis is regarded as a junior synonym of V. sayl. Karyotypes of male and female V. sayi from central Texas reveal 2n = 78. This number is almost twice that previously reported for any other harvestman. The male reproductive system of V. sayi is described and illustrated. Natural history data are provided for egg laying and developmental times. The supercooling points ranged from —2.5 to —7.2°C, with poor recovery and death following thawing. Key Words: tering The identification of the Cosmetidae of the U.S.A. presents a difficult problem. How much variation in a character constitutes a different species? As presented in the liter- ature, there is a maximum of five species in the U.S.A.: Vonones sayi (Simon) in central and eastern U.S.A. (type from Texas), Von- ones bimaculata (Banks) from California (correct locality?), Vonones ornata (Say) from Florida and Georgia, Vonones dentica (Walker) from Ohio, and Platycynorta transversalis Roewer from Texas. Based on the original description, Roewer’s species is a synonym of V. sayi. Goodnight and Good- night (1953, 1973) treated all the eastern U.S.A. populations as one species. From our examination, we find that the abdom- inal spines of V. ornata are always present and consistently larger than the similarly placed low tubercles in V. sayi. Even so, doubt remains about the range and extent of variation in the U.S.A. cosmetids. To help resolve this problem we attempted to karyotype Vonones from numerous locali- ties in the U.S.A. Karyotype, reproductive system, natural history, supercooling, overwin- In addition to the karyological study, we obtained biological data from captive spec- imens. Little natural history data have been published on the members of the exclu- sively American family Cosmetidae. Only a single note has been published on biology of Vonones (Goodnight 1958): V. ““ornata”’ in Indiana. Three other reports have ap- peared on biology of cosmetids: Juberthie (1972) on Cynorta cubana (Banks) from Cuba, Canals (1936) on Gyne orensis (S6- rensen) and Metalibitia paraguayensis (S6- rensen) from Argentina, and Goodnight and Goodnight (1976) on Erginulus clavotibialis (Pickard-Cambridge) from Belize. RESULTS AND DISCUSSION Karyology.— Parthasarathy and Good- night (1958) reported observations on the genital morphology and karyotype of “Vo- nones ornata (Wood)” from Indiana. Ob- taining countable spreads of V. sayi chro- mosomes proved quite difficult, a point not noted by Parthasarathy and Goodnight. Be- cause juveniles are very rarely seen, adults Bans eK aw BX ber wee SH Sm 2K BE SR CO KB LMR ES SS WH SE GR KA ss ts gw 2% R3e “ak ne te Ge ch oF BU CA Lee sz Ke xe x 32 Fig. 1. Karyotype of male Vonones sayi from Kerr Co., Texas (2n = 78). were the sole source of material for karyo- typing. During 1982 and 1983, V. ornata were obtained from Florida whereas V. sayi were collected from numerous localities in Texas, Oklahoma, Nebraska (Jefferson Co.), and Tennessee (Davidson Co.). Ovaries and tes- tes were prepared for chromosomal obser- vation during January—March, May, July, October, and November. From these, only seven cells from one male collected during May resulted in observable (but not count- able) individual chromosomes. Not until 1984 were we able to obtain living material collected in the proper season, from which we successfully karyotyped two males and one female from central Texas, collected in April. Karyotypes using the gonads as the source of cells are best obtained from subadult har- vestmen. In our case, active gonadial cells in adults were sought. From a series of an- imals collected 6.4 km E of Kerrville, Kerr Co., Texas, we prepared (following the air- drying method of Cokendolpher and Brown 1985) five males and 11 females on 5 April. A second series of animals from the same locality was karyotyped on 17 May. All an- imals had been dipped or injected with a 0.005% colchicine solution 24-40 hours prior to karyotyping. Although several cells from many slides were dividing, clear, countable spreads were only obtained from two males and a female from 5 April with a 24 hour colchicine treatment. Good spreads were only counted for one cell in the female (2n = 78) and four cells for males (n = 39, 2n = 78). Chromosomes were often overlapping in spreads with only slightly condensed chromosomes. Despite these dif- ficulties, these spreads were used to verify the chromosome morphologies observed in more highly condensed plates. Altogether, there were 38 pairs of submeta- and meta- centric chromosomes and one pair (small- est) in which the centromere could not be detected (Fig. 1). No obvious sex chromo- some could be determined by either mor- phology or activity. Our results are totally unlike those of Par- thasarathy and Goodnight (1958). They re- ported finding 2n = 24 + X in males. Be- cause they used squash and sectioning techniques, we assume they are in error. Not 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Male genital system (minus penis) of Vonones sayl, preserved in 80% ethanol, critical point dried, gold coated, and then examined with a Scanning Elec- tron Microscope (Hitachi model s-570). See Fig. 3 for abbreviations. only is their count much lower than ours, but an XO-XX (male heterogametic) sex determining system has never been verified in any opilionid (Tsurusaki and Cokendol- pher in press). Karyotyping additional males from Indiana with an air-drying method will be required to be certain. The highest chromosome number pre- viously recorded for Opiliones is 2n = 40 in Pseudobiantes japonicus Hirst (Tsurusaki 1986, Tsurusaki and Cokendolpher in press). Although in different families, Vonones and Pseudobiantes (Phalangodidae) are the only members of the suborder Laniatores that have been karyotyped. Although we could not obtain chromo- some counts for numerous populations of Fig. 3. Diagrammatic view of Vonones sayi male genital system. T = testis, VE = vasa efferens, VD = vas deferens, SR = seminal reservoir, PO = propulsion organ, P = penis, DE = ductus ejaculatorius, TR = truncus, G = glans, AT = apophysis of truncus, AG = apophysis of glans. V. sayi, we did obtain what is probably the first accurate count. Male genital morphology.—While dis- secting numerous males for this study and that of Jones and Cokendolpher (1985), we found the description and illustration of the male reproductive system of Vonones pre- sented by Parthasarathy and Goodnight (1958) to be incorrect. They reported find- ing a single testicle connected at one end by a sperm duct to the vas deferens. As their drawing has served as the standard for the Cosmetidae (Suzuki 1966), we feel it desir- able to re-illustrate the system in greater detail. After careful dissection of living an- imals, we find the reproductive system to be similar to other harvestmen. Although the vasa efferentia are small and difficult to see, they are attached to both ends of the VOLUME 93, NUMBER 1 testis (Fig. 2). Other details are as in Fig. 3 (morphological nomenclature as in Juber- thie 1965, Martens 1976, and Silhavy 1966). Natural history.—We obtained data on natural history while maintaining animals in culture over several years. Based on our studies, V. sayi spermatogenesis is most ac- tive in April and May in central Texas, and is the time period during which specimens were collected for our previous paper on spermatogenesis (Jones and Cokendolpher 1985). Similarly, peak egg laying occurs dur- ing April. Based on cytological studies, Goodnight (1958) reported V. ‘“‘ornata” in Indiana must breed and lay eggs throughout the summer months. Egg laying in Tropical species is different with C. cubana laying eggs year-round (Juberthie 1972), and E. clavotibialis (Goodnight and Goodnight 1976) laying eggs during the rainy seasons (May and December). The juvenile stages of C. cubana last 4.5- 7 months at 20-25°C and E. clavotibialis requires 121 days post-hatch (temperature not reported, presumably 20-—26°C as used for eggs). The lengths of the juvenile stages in V. sayi are unknown. Individual V. sayi collected as adults have been held in cap- tivity for over three years on a diet of dead insects, live fruit flies, and fruits. Cynorta cubana is recorded (Juberthie 1972) to live 2.5 years as an adult. Because so little is known about juveniles of the Cosmetidae, we here present our data. Females of V. sayi, in captivity, lay eggs during the winter. Several females collected in Wichita Falls, north-central Texas, pro- duced the following numbers of eggs: De- cember (1 egg), February (2), March (32), and April (50). These females were collected during May of the previous year. Eggs are laid singularly (usually only one per day) and packed into crevices of moist wood. The eggs are covered with small pieces of wood and soil, presumably to hide them. Covering eggs with debris is also reported in C. cubana by Juberthie (1972) and as- sorted cosmetids and gonyleptids by Canals 89 (1936). The actual egg laying process was never observed. Egg development took 20- 38 days (n = 17). This range is obviously influenced by temperature, which varied greatly over the egg-laying period in an un- heated room (about 5—20°C). Cynorta cu- bana egg development lasts 16-27.5 days at 28-20°C, whereas E. clavotibialis lasts 13 and 23-27 days at 26 and 20°C. Because the V. sayi eggs were camouflaged, some shorter incubation times may be due to the fact that they went undetected for some time. Of the 17 individuals which hatched only two pre- sumably molted to the third instar (2nd nymphal stage) (16 and 25 days) and the 16 day 3rd instar presumably molted to the 4th instar (25 days later). As these individuals died while still juveniles, the number of molts to adulthood is unknown. Although never observed in V. sayi, it is assumed that the first instar (emerging lar- va) molts immediately after leaving the egg shell, as do C. cubana and E. clavotibialis. No exuviae were recovered and instars were determined by uneven jumps in body size. Molting and hatching were not observed, so the chewing (‘eating’) of the exuviae could not be verified. Such behavior is known for several cosmetids and gonyleptids (Canals 1936, Juberthie 1972) and postulated for E. clavotibialis by Goodnight and Goodnight (1976). The first instar or larva does not have a tooth-like structure to aid in hatch- ing. The absence of this egg-tooth is also reported in the Gonyleptidae (Munoz Cue- vas 1971). Such a tooth is also missing in a drawing of the larva of E. clavotibialis (Goodnight and Goodnight 1976). Supercooling. —Cold-hardiness in arthro- pods can be enhanced by: (1) cold-accli- mation, (2) supercooling, and (3) freezing tolerance. Preparation to avoid injury at temperatures too low for continued growth is referred to as cold-acclimation. Super- cooling is the adaptation in which the freez- ing point of body fluids is lowered. If an animal can withstand freezing of body tis- sues, it is considered to be freeze-tolerant. 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Because the supercooling point is the freez- ing point, it is the lowest temperature sur- vived by a freeze-intolerant species. Vonones sayi is a member of the Cos- metidae, a primarily Tropical American family. Besides being the cosmetid with the most northern distribution (reported as V. “ornata” from Indiana by Goodnight 1958), it has the distinction of having the largest range (Indiana south to Florida, west to Ne- braska and Texas in the U.S.A., and north- ern Mexico). Because adults are collected throughout the year and juveniles are pres- ent during winter in Texas, overwintering for this species poses several problems. To overwinter, these animals must either avoid freezing (by cold-acclimation or supercool- ing) or be freeze-tolerant. Several animals from Kerr and Wichita counties (central and northern), Texas, were supercooled during 1981 (methods follow Francke et al. 1986, except probe was at- tached to abdomen with Vaseline on juve- niles and taped to adults). Field observa- tions revealed that these animals gather under wood and logs helping to insulate the animals from the winter cold. Two males tested during early May had supercooling poimts, of =3:4-and —5.5°@: Immediate thawing resulted in poor recovery, a lack of ambulatory ability, and death by the third day. Two second instars and one third instar reared at room temperature were super- cooled three days post-feeding. The second instars supercooled to —7.0 and —7.2°C, whereas the 3rd instar supercooled at —3.5°C. Like the adults, poor recovery was noted with death by the third day. Although the sample sizes are small and our warming periods were not timed (see Baust and Rojas 1985), the values obtained suggest that this species supercools to avoid bodily freezing. In February 1990, winter-collected adults from Shelby County, eastern Texas, were allowed to live in the laboratory at 24—26°C for a week prior to further experiments. One group of six adults (males and females) was cooled (average rate 0.1°C/min) to —3.5°C and a second group of six adults was cooled (average rate 1.25°C) to —26°C for 36 hours. Warming the animals (at approximately the same rate as cooling) revealed: (1) none sur- vived freezing to — 26°C, (2) two (male and female) survived cooling to —3.5°C. The survivors moved and fed normally for one week, at which time their supercooling points were determined to be —2.5 and —5.0°C. Two females which had not been previously frozen were also tested. Their su- percooling points were —3.2 and —3.8°C. As a large sample of animals was not avail- able, the animals could not have been tested at a variety of cooling and warming rates to determine if there was an optimal rate (see Baust and Rojas 1985). While the super- cooling points were sufficient to protect an- imals in Texas, another means would be required for overwintering populations in the northern U.S.A. Goodnight (1958) re- ported that this species aggregates under piles of logs or brush heaps with the coming of fall and apparently disappears into the ground to avoid winter cold. It is unknown whether these animals remain relatively ac- tive or diapause and to what depth they retreat. If animals retreat deep into earth cracks and holes, their supercooling points would not have to be lower than the south- ern populations because of the warmer tem- peratures. ACKNOWLEDGMENTS We express our sincere gratitude to Dr. Jerry Berlin for suggestions and permission to use the electron microscope and labora- tory facilities at the Department of Biolog- ical Sciences, Texas Tech University. Dr. Nobuo Tsurusaki, Tottori University, kind- ly commented on a draft of the manuscript. His ever helpful comments over the years have been greatly appreciated and respect- ed. Mr. William F. Rapp is thanked for helping the senior author collect Vonones in Nebraska. Mr. Kari J. McWest, Dr. W. David Sissom, and Dr. James C. Trager kindly provided living Vonones from east- VOLUME 93, NUMBER 1 ern Texas, Tennessee, and Florida, respec- tively. LITERATURE CITED Baust, J. G. and R. R. Rojas. 1985. Review—insect cold hardiness: Facts and fancy. Journal of Insect Physiology 31(10): 755-759. Canals, J. 1936. Observaciones bioldgicas en arac- nidos del orden opiliones. Revista Chilena de Hi- storia Natural 40: 61-63. Cokendolpher, J. C. and J. D. Brown. 1985. Air-dry method for studying chromosomes of insects and arachnids. Entomological News 96(3): 114-118. Francke, O. F., J. C. Cokendolpher, and L. R. Potts. 1986. Supercooling studies on North American fire ants (Hymenoptera: Formicidae). Southwest- ern Naturalist 31(1): 87-94. Goodnight, C.J. 1958. Two representatives ofa trop- ical suborder of opilionids (Arachnida) found in Indiana. Indiana Academy of Science 67: 322- B23. Goodnight, C. J. and M. L. Goodnight. 1953. Tax- onomic recognition of variation in Opiliones. Sys- tematic Zoology 2(4): 173-180. . 1973. Opilionids (Phalangida) from Mexican caves. Association of Mexican Cave Studies, Bul- letin 5, pp. 83-96. Goodnight, M. L. and C. J. Goodnight. 1976. Ob- servations on the systematics, development, and habits of Erginulus clavotibialis (Opiliones: Cos- metidae). Transactions of the American Micro- scopical Society 95(4): 654-664. Jones, S. R. and J. C. Cokendolpher. 1985. Sper- matogenesis in the harvestman Vonones sayi (Simon) (Opiliones: Laniatores: Cosmetidae). Bul- 91 letin of the British Arachnological Society 6(9): 403-413. Juberthie, C. 1965. Données sur l’écologie, la déve- loppement et la reproduction des Opilions. Revue d’Ecologie et de Biologie du Sol 2(3): 377-396. 1972. Reproduction et développement d’un opilion Cosmetidae, Cynorta cubana (Banks), de Cuba. Annales de Speleologie 27(4): 773-785. Martens, J. 1976. Genitalmorphologie, System und Phylogenie der Weberknechte (Arachnida: Opilio- nes). Entomologica Germanica 3(1/2): 51-68. Mufioz Cuevas, A. 1971. Etude du développement embryonnaire de Pachylus quinamavidensis (Arachnida, Opilions, Laniatores). Bulletin du Museum National d’Historie Naturelle, 2e Sér. 42(6): 1238-1250. Parthasarathy, M. D. and C. J. Goodnight. 1958. The chromosomal patterns of some Opiliones (Arach- nida). Transactions of the American Microscopi- cal Society 77(4): 353-364. Silhavy, V. 1966. Okologische und genitaliomor- phologische Bemerkungen iiber einige Arten der Familie Cosmetidae Simon aus Kuba (Arachnoi- dea, Opilionidea). Deutsche Entomologische Zeit- schrift, n.s. 13(1/3): 263-266. Suzuki, S. 1966. (Phalangida, Opiliones), pp. 90-139. In T. Uchida, Débutsu Keitébunrui-gaku [Sys- tematic Zoology]. Nakayama-shoten, Tokyo, 7(2A) (in Japanese). Tsurusaki, N. 1986. Chromosomes of harvestmen (Opiliones, Arachnida): A review of ongoing re- search and method of chromosome observation. Seibutsu Ky6zai, No. 21, pp. 33-49 (in Japanese). Tsurusaki, N. and J. C. Cokendolpher. (In press.) Chromosomes of sixteen species of harvestmen (Arachnida, Opiliones, Caddidae and Phalangi- idae). Journal of Arachnology. —— PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 92-95 DESCRIPTION OF A PHYTOPHAGOUS DORYCTINE BRACONID FROM BRASIL (HYMENOPTERA: BRACONIDAE) PAUL M. MARSH Systematic Entomology Laboratory, U.S. Department of Agriculture, % U.S. National Museum of Natural History, NHB-168, Washington, D.C. 20560. Abstract. —A new species of the doryctine braconid genus Allorhogas is described from Brasil. Biological studies of this species suggest that it is phytophagous on tissue of legume seeds. Key Words: In 1986 several specimens of a doryctine braconid were submitted for identification by Margarete de Macédo and Richard Mon- teiro, Universidade Federal do Rio de Ja- neiro, Brasil. Biological information with the specimens stated that they were reared from legume seeds and were apparently feeding on the seed tissue. I identified the braconids as an undescribed species of A/- lorhogas Gahan. Although many specimens of this genus have been reared from galls or stems (Marsh 1979) or seeds (Whitehead 1975), it has usually been suspected they attacked some sort of insect larva in these plant tissues. Phytophagy had never been reported, or even suspected, in Braconidae. However, these researchers seemed to have good evidence that the braconids were in- deed feeding on the seed tissue and have subsequently published preliminary results of their studies on the biology of these wasps (de Macédo and Monteiro 1989). Most of the specimens of the genus A/- lorhogas in the USNM collection, both de- scribed and undescribed species, have long Ovipositors characteristic of the subfamily Doryctinae and suggesting the need to pen- etrate plant tissue to find host larvae. How- ever, both A. muesebecki Guimaraes and Braconidae, Doryctinae, phytophagous the new species described here have ex- tremely short ovipositors which could in- dicate they do not need to find a host deep in plant tissue. Guimaraes (1957) states that A, muesebecki was reared from plant galls and concluded that, since this was the only insect reared from the galls, the galls were possibly induced by the wasp. Although phytophagy has never been found before in the Braconidae, it does oc- cur sporadically in the Chalcidoidea (some Eurytomidae, Torymidae and Eulophidae, and all Tanaostigmatidae and Agaonidae). Seeds are highly nutritious, and doryctine braconids, as ectoparasites which usually paralyze their hosts (idiobionts), could be considered little more than specialized Fig. 1. wings. Allorhogas dyspistus, n. sp., fore and hind VOLUME 93, NUMBER | 98 Figs. 2-8. Allorhogas dyspistus, n. sp.: 2, head and thorax, lateral view; 3, fore tibia; 4, propodeum, dorsal view; 5, metasoma, lateral view of apex; 6, thorax, dorsal view; 7, mesonotum, dorsal view; 8, metasoma, dorsal view. 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON predators, so why not evolve to eat seed tissue when insect hosts are not present? We know very little about the biologies of Bra- conidae which attack cryptic hosts in plant tissue, particularly in the tropics, and al- though it seems incredible at first, perhaps the occurence of phytophagy is not that un- reasonable. There are other doryctine gen- era with species that have been reared from seeds, e.g. Stenocorse and Heterospilus. The two species of these genera associated with seeds plus several undescribed neotropical species of Heterospilis all have very short Ovipositors similar to the A//orhogas species mentioned here. Clearly there need to be more detailed studies of this phenomenon and I hope that this paper and the study by de Macédo and Monteiro will stimulate such research. Allorhogas dyspistus Marsh, NEw SPECIES Female.— Body length, 3.5—4.0 mm; fore wing length, 3.0 mm. Body color: honey yellow; antennal flagellum, lateral faces of scutellum, metanotum and basal-lateral ar- eas of propodeum, ovipositor sheaths, hind tarsus, spots at apex of first tergum, and transverse stripe across middle of tergum 2+3 black; wing veins brown, stigma trans- lucent. Head: antenna with 27-28 flagel- lomeres; face weakly granulo-rugose with smooth raised median area; frons, vertex and temple granular; frons concave; malar space % eye height; ocelli small, diameter of lateral ocellus about 4 ocell-ocular dis- tance. Thorax: propleuron (Fig. 2) rugose with median longitudinal scrobiculate groove; mesonotum (Fig. 2) sharply decli- vous anteriorly, mesonotal lobes (Figs. 6, 7) granular, middle lobe with scrobiculate groove posteriorly which sometimes ex- tends nearly to anterior edge, notauli scro- biculate and meeting posteriorly in wide ru- gose area; scutellum granular; mesopleuron (Fig. 2) granular, subalar groove strongly scrobiculate, sternaulus smooth; propo- deum (Figs. 4, 6) rugose, with two weakly granular or coriaceous semicircular areas laterally at base. Metasoma: first tergum (Fig. 8) wider at apex than long, arched at base, longitudinally striate, with two basal cari- nae connected by cross carina near base; tergum 2+3 longitudinally striate; fourth tergum striate on basal half; remainder of terga granular; ovipositor (Fig. 5) barely vis- ible dorsally. Legs: fore tibia with row of 10-12 stout spines on anterior edge (Fig. 3). Wings (Fig. 1): first and second segment of radius in fore wing about equal in length; postnervellus in hind wing strongly curved toward wing apex. Male.—Essentially as in female except femora swollen, hind femur length only slightly more than twice greatest width. Holotype.— Female: BRASIL, Marica, Rio de Janeiro; January 1986; ex seeds of Pithecellobium tortum Mart. Deposited in the U.S. National Museum, Washington, |DM Ge Paratypes.— 3 females, 8 males, same data as holotype. Deposited in the U.S. National Museum, the Canadian National Collec- tion, Ottawa, and the Universidade Federal do Rio de Janeiro, Brasil. Etymology.—The specific name is from the Greek dyspistos meaning “‘hard to be- lieve, incredible,” which refers to the phy- tophagous behavior reported for this spe- cies. This species is similar to A. muesebecki Guimaraes, also from Brasil, but is distin- guished by its larger eyes and shorter malar space; in dyspistus the malar space is % of the eye height, in muesebecki it is nearly equal to the eye height. ACKNOWLEDGMENTS I thank Margarete de Macédo for provid- ing specimens for this description. Scott Shaw (University of Wyoming) and Robert Wharton (Texas A&M University) offered many helpful suggestions on the manu- script. VOLUME 93, NUMBER 1 95 LITERATURE CITED Marsh, P.M. 1979. Family Braconidae, pp. 144-295. In Krombein, K. V. et al., eds., Catalog of Hy- Guimaraes, J. A. 1957. Contribuicaéo ao estudo de menoptera in America north of Mexico. Smith- Cecidilogia Brasiliana. Thesis, Esc. Nac. Agron., sonian Institution Press, Washington, D.C. Univ. Rural, Rio de Janeiro, p. 25. Whitehead, D. R. 1975. Parasitic Hymenoptera as- Macédo, M. V. de and R. T. Monteiro. 1989. Seed sociated with bruchid-infested fruits in Costa Rica. predation by a braconid wasp, A/lorhogas sp. (Hy- J. Wash. Acad. Sci. 65: 108-116. menoptera). J. New York Entomol. Soc. 97: 358- 362. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 96-100 LIFE HISTORY STUDIES, HOST RECORDS, AND MORPHOLOGICAL DESCRIPTION OF GENITALIA OF EURYTOMA TYLODERMATIS ASHM. (HYMENOPTERA: EURYTOMIDAE) FROM SOUTH DAKOTA B. McDANIEL AND A. BOE Plant Science Department, South Dakota State University, Brookings, South Dakota 57007. Abstract. —Eurytoma tylodermatis Ashm. was found to be parasitic on Acanthoscelides perforatus (Horn) in seed pods of Canada milk-vetch (Astragalus canadensis L.). Obser- vations on larval behavior and frequency of parasitism are discussed. Male and female genitalia of E. tylodermatis are illustrated. Key Words: Eurytoma tylodermatis Ashm. parasitizes bruchid beetles and numerous other Cole- optera throughout the eastern and central United States (Burks 1979). Bugbee (1967) listed 56 host species for E. tylodermatis, many of which are of economic importance, but cautioned that more exact determina- tions of host relationships are needed. He stated that the species can act as either a primary or secondary parasite on beetles and moths. Pierce (1908) regarded it as an im- portant parasite of the cotton boll weevil (Anthonomus grandis Boheman) in Texas, and Ritcher (1936) recorded its parasitism of the larger apple curculio (7achypterellus quadrigibbus magnus List) in Wisconsin. This paper records a new host for E. ty- lodermatis, presents descriptions of E. ty- lodermatis male and female genitalia, and describes observations and data regarding larvae and adults of E. tylodermatis asso- ciated with Canada milk-vetch in eastern South Dakota. MATERIALS AND METHODS In August 1988 and 1989 mature seed pods (Fig. 1) of Canada milk-vetch were Acanthoscelides perforatus, Dinarmus acutus, Astragalus canadensis collected from a field nursery at Brookings, South Dakota and placed in 0.9-liter glass jars (five to ten racemes per Jar in ten jars) in 1988 and 1.8-liter paper cartons (ten to twenty racemes per carton in five cartons) in 1989. Jars were sealed and maintained at room temperature. Cartons were wrapped with aluminum foil and fitted with a 5-ml vial to serve as a trap (Fig. 2). Cartons were placed on a shaded bench in a greenhouse. Chalcids that emerged from pods in the jars during fall of 1988 and bruchids that emerged during spring of 1989 were re- moved, identified, and counted in July 1989. Cartons were checked daily through Octo- ber 1989 and chalcids trapped in the vials were identified and counted. In June 1989, green pods containing de- veloping seeds were dissected in order to make observations on chalcids parasitizing A. perforatus larvae. Chalcid larvae found parasitizing beetle larvae were placed 1n 5-ml vials and observed daily until adults emerged. Genitalia were removed from ten females and ten males of FE. tylodermatis. Ilustra- tions were made for both sexes and data VOLUME 93, NUMBER | Fig. 1. Pods of Canada milk-vetch (Astragalus can- adensis L.). were recorded for number of rami spines and eighth tergite setae of the females. Mor- phological terminology is that of Snodgrass (1941). RESULTS AND DISCUSSION The following is a description of female and male reproductive structures based on the 20 adults obtained from laboratory rear- ings. Female (Fig. 3): 2nd Valvifers (Vf2) (semicircular sheaths) with two setae near apodemes of laminated bridge (Lam.Br.); ramus spines (Ra.Sp.) range from 23 to 35 on right valvifer (mean of 29.6 for 10 spec- imens), and from 25 to 36 on left valvifer 97 BPS. S oF Fig. 2. Paper carton rearing container. (mean of 30.5 for 10 specimens); spines widely spaced in laminated bridge region, close together near fulcral plate region; ful- cral plate (Ful.Pl.) attachment near notched area (Ful.Pl.Not.) of fulcral plate; outer ovi- positor plate (Ops.Ot.PI.) fused with 8th ter- gite (8-Teg.); 8th tergite with row of setae (8-Teg.Set.) ranging from 41 to 51 (mean of 46.3 for 10 specimens), setae count begins with first seta anterior to cercus (Ce.); setae single adjacent to cercus becoming paired toward attachment of outer ovipositor plate and fulcral plate; setal region plated, bor- dered by dark line that divides fused outer ovipositor plate into light and dark pig- mented areas; apex of 8th tergite with series of long setae similar in structure to other 8th tergite setae [these setae associated with epipygium (Ep.)]; cercus with 5 setae of dif- ferent sizes and shapes. Inner ovipositor plate (Ops.In.Pl.) separated from 2nd val- vifers (Vf2) by darkened region that con- nects apodemes of outer and inner rami and the groove in which the fulcral plate fits along with monitoring spines (Ful.PI.Sp.); inner Ovipositor plate with plated region ex- tending to region of fused ovipositor sheaths (Ops.Sh.); ovipositor sheath not articulated, containing the ovipositor sheath ligament (Ops.Sh.Lg.) which connects sheaths in which the ovipositor is held; ovipositor sheaths lightly striated with typical series of eurytomid setae at apex. 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON PL _ Ful Pl. Sp —_Ful PINot Fig. 3. Ovipositor of Eurytoma tylodermatis Ashm. Abbreviations: (Lam.Br.) Laminated bridge; (Ra.) Ra- mus; (Ra.Sp.) Ramus spines; (Vf2) 2nd Valvifer (Semi- circular sheet); (Ful.Pl.Sp.) Fulcral plate spines; (Ful.P1.Not.) Fulcral plate notch; (Ful.Pl.) Fulcral plate; (Ops.In.Pl.) Ovipositor inner plate; (Ops.Ot.Pl.) Ovi- positor outer plate; (Ops.Sh.Lg.) Ovipositor sheath lig- ament; (8-Teg.) 8th tergite; (8-Teg.Set.) 8th tergite se- tae; (Ops.Sh.) Ovipositor sheath; (Ce.) Cercus; (Ep.) Epipygium. Male (Figs. 4, 5): reproductive apparatus with 4 dorsal aedeagal sensory pores (Aeg.S.Por.); aedeagal striae (Aeg.Str.) ex- tend from posterior aedeagal sensory pores to beyond anterior aedeagal sensory pores; aedeagal sensory pores may be paired or staggered on dorsal surface of aedeagus (Fig. 4); ventrally apex of aedeagus with five sen- sory pores on each side; these smaller than dorsal pores; aedeagus (Aed.) elongated ca- pable of extending beyond digiti (Dgi.); par- ameres with three setae; apex setae normally hidden between digiti and aedeagus; para- mere plate setae (Par. P1.Set.) located on out- er margin of curved paramere plate (Par.PI.); posterior paramere plate setae longer and larger than anterior paramere plate setae; anterior paramere plate setae located adja- cent to digiti apodemes (Dig.Apd.); apex se- tae similar in size to latter paramere setae; digiti with two finger-like teeth and a round- ed projection; each digitus with two pores on base; digital apodemes short (Fig. 5), fused to aedeagal apodemes (Aeg.Apd.); ae- deagal apodemes protrude from caulis (Ca.); paramere plates (Par.Pl.) and Volsellar plates (Vos.Pl.) fused to caulis with pigmentation being darker on the sides of reproductive apparatus and lighter in the center; caulis forms opening below digiti in which aedea- gal apodemes are connected to muscles of the 8th tergite. There is a narrow nonpig- mented membrane connecting the aedeagus to the epipygium and 8th tergite. This mem- brane is beset with small spine-like setae. The membranous portion of this structure surrounds the aedeagus and contains two acorn-like setae. This structure is attached to the aedeagus and apparently plays a role in exsertion and retrieval of the entire male reproductive apparatus as well as the ae- deagus. PARASITIC BEHAVIOR Information on behavior of E. tyloder- matis larvae was obtained by splitting green, well-developed seed pods along their septa to expose seeds in the chambers of each of the two locules. On July 18, 1988, we observed a pinkish-white larva that was feeding on an A. perforatus larva. A grayish- black, pubescent hatched egg, which resem- bled E. tylodermatis eggs (Pierce 1908), was attached to the bruchid larva. The pod loc- ule containing the parasitic larva and its host was placed in a 5-ml vial and stored at room temperature. Daily observations were made and on July 22 the parasitic larva crawled out of the locule, defecated, and pupated. By July 26 the pupa had become solid black except for prominent reddish- VOLUME 93, NUMBER 1 SS — Parelset: Sein Aeg.S.Por. Vos. Pl. Figs. 4, 5. 99 F Par Pl Male reproductive apparatus of Eurytoma tylodermatis Ashm., dorsal and ventral, respectively. Abbreviations: (Aeg.) Aedeagus; (Par.Pl.Set.) Paramere plate setae; (Par.Pl.) Paramere plate; (Aeg.S.Por.) Ae- deagal sensory pores; (Aeg.Str.) Aedeagal striae; (F.Par.P1.) Fused paramere plate; (Ca.) Caulis; (Vos.PI.) Volsellar plate; (Aeg.Apd.) Aedeagal apodemes; (Dgi.) Digiti; (Dgi.Apd.) Digiti apodemes. brown compound eyes. On July 30 an adult female of E. tylodermatis emerged and was one of the ten females studied for the struc- ture of the genitalia. These observations made it possible to recognize E. tyloder- matis larvae in subsequent pod dissections. Numbers of adult E. tylodermatis and the pteromalid Dinarmus acutus Thomson reared from inflorescences of Canada milk- vetch in 1988 and 1989 are presented in Table 1. Total numbers recorded of each species were 68 and 112 for E. tylodermatis and D. acutus, respectively. Male D. acutus outnumbered females by 67% while females of E. tylodermatis outnumbered males by 52%. The number of A. perforatus adults reared in 1988 was 70, indicating that ap- proximately 35 and 24% of the beetles were parasitized by D. acutus and E. tyloder- matis, respectively. Since D. acutus out- numbered E. tylodermatis by approximate- ly 50% in 1989 rearings, the pteromalid was the more predominant parasite of A. per- foratus in this study. These data agree close- ly with a previous study on the effects of parasitism by D. acutus on Canada milk- 100 Table 1. Numbers of parasitic chalcids reared from Canada milk-vetch pods infested with the bruchid Acanthoscelides perforatus. Dinarmus acutus Eurytoma tylodermatis Year Male Female Male Female encnewenennccccenceccecececsanesccccsssesee= no. ee 1988 39 2 18 22 1989 31 2 9 19 vetch seed predation by Acanthoscelides perforatus (Horn) (Boe et al. 1989). In that study, D. acutus parasitized over 45% of A. perforatus larvae. We have reared E. tylodermatis from seed pods of American licorice (Glycyrrhiza lep- idota Pursh) and false indigo (Amorpha fru- ticosa L.) containing Acanthoscelides aureo- lus (Horn) and A. submuticus (Sharp), respectively. It is likely that E. tylodermatis also parasitizes bruchid beetles in seeds of these two legumes. However, we have not dissected pods to observe E. tylodermatis larvae feeding behavior in these species. ACKNOWLEDGMENTS The authors extend their gratitude to Dr. E. E. Grissell, Systematic Entomology Lab- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON oratory, ARS-USDA for providing identi- fication of Eurytoma tylodermatis and to Kathy Robbins for assistance in data ac- quisition. This research was supported by the South Dakota Agricultural Experiment Station, SDSU, Brookings, project numbers H-277 and H-388, contribution no. 2457. LITERATURE CITED Boe, A., B. McDaniel, and K. Robbins. 1989. Direct effect of parasitism by Dinarmus acutus Thomson on seed predation by Acanthoscelides perforatus (Horn) in Canada milk-vetch. J. Range Manage. 42: 514-515. Bugbee, R. E. 1967. Revision of chalcid wasps of the genus Eurytoma in America north of Mexico. Proc. U.S. Nat. Mus. 118: 433-552. Burks, B. D. 1979. Eurytomidae. Jn Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and B. D. Burks, eds., Catalog of Hymenoptera in America north of Mexico. Vol. 1. Symphyta and Apocrita (Para- sitica). Smithsonian Institution Press, Washing- ton, D.C. 1198 pp. Pierce, W. D. 1908. The economic bearing of recent studies of the parasites of the cotton boll weevil. J. Econ. Entomol. 1: 119-121. Ritcher, P. O. 1936. Larger apple curculio in Wis- consin. J. Econ. Entomol. 29: 698-703. Snodgrass, R. E. 1941. The male genitalia of Hy- menoptera. Smithsonian Miscellaneous Collec- tion 99(14): 36-37. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 101-124 THE SPECIES OF ANASTREPHA (DIPTERA: TEPHRITIDAE) WITH A GRANDIS-TYPE WING PATTERN ALLEN L. NORRBOM Systematic Entomology Laboratory, USDA, ARS, PSI, % U.S. National Museum, NHB 168, Washington, D.C. 20560. Abstract.—A diagnostic key, illustrations, and descriptions are provided for grandis (Macquart) and the five other species of Anastrepha with similar wing patterns: atrigona Hendel, bezzii Lima (= balloui Stone, n. syn.), castilloi, n. sp., grandicula, n. sp., and shannoni Stone. Phylogenetic relationships among these species are discussed. Anastrepha bezzii is removed from the grandis species group, and bivittata (Macquart) and fumipennis Lima are tentatively included. A lectotype is designated for grandis. The egg of shannoni is briefly described. Host plant and distribution data are summarized. Key Words: Anastrepha, grandis, descriptions, phylogeny Resumen. —En este trabajo se presenta una clave, descripciones e ilustraciones de gran- dis (Macquart) y otras cinco especies de Anastrepha con patrones de coloracion alar semejantes: atrigona Hendel, bezzii Lima (= balloui Stone, syn. n.), castilloi, sp. n., gran- dicula, sp. n., y shannoni Stone. Relacciones filogenéticas entre estas especies son discu- tidas. Anastrepha bezzii se transfiere del grupo de especies ““grandis”’, y bivittata (Macquart) y fumipennis Lima se incluyen tentativamente. Se designa un lectotipo de grandis. El huevo de shannoni se describe brevemente. Los datos de plantas de alimentacion y de distribucion son presentados. Anastrepha grandis (Macquart) is a pest of the fruits of various native and intro- duced species of Cucurbitaceae in many ar- eas of South America (Norrbom and Kim 1988b). It has recently received consider- able attention in regard to the extent to which it attacks honeydew, a type of melon (Cucu- mis melo L.) that is grown commercially in a number of Latin American countries (Harper 1987). In this paper, I describe two new species of Anastrepha with wing pat- terns similar to grandis, and that might be confused with it. I also discuss the relation- ships among these species and those that Steyskal (1977) placed in the grandis species group. MATERIALS AND METHODS I follow the morphological terminology of McAlpine (1981) and White (1988), ex- cept as noted in Norrbom and Kim (1988a). Acronyms for institutions used in the text are as follows: AMNH—American Muse- um of Natural History, New York; BMNH— British Museum (Natural History), London; CNC—Canadian National Collection, Ot- tawa; CMP—Carnegie Museum of Natural History, Pittsburgh; CUI—Cornell Univer- sity, Ithaca, FMNH—Field Museum of Natural History, Chicago; FSCA—Florida State Collection of Arthropods, Gainesville; DEI—Institut fiir Pflanzenschutzforschung der Akademie der Landwirtswissenschaf- 102 ten, Eberswalde; ICA—Instituto Colombi- ano Agropecuario, Palmira and Medellin; INPA—Instituto Nacional de Pesquisas da Amazonia, Manaus; INTA—Instituto Na- cional de Technologia Agropecuaria, Con- cordia; LACM-—-Los Angeles County Mu- seum of Natural History; MCZ—Museum of Comparative Zoology, Harvard Univer- sity, Cambridge; NMW-— Naturhistorisches Museum Wien; PAN—Polish Academy of Sciences, Warsaw; PMMT—Programa Mosca del Mediterraneo, Sanidad Vegetal, SARH, Tapachula, Mexico; SMT—Staat- liches Museum fiir Tierkunde, Dresden; TMB-—Termeszettudomanyi Muzeum, Bu- dapest,; UCV—Universidad Central de Venezuela, Maracay; UCRSJ—Universi- dad de Costa Rica, San José; UMO— Uni- versity Museum, Oxford; USNM—Nation- al Museum of Natural History, Smithsonian Institution, Washington, D.C.; USU—Utah State University, Logan; WSU — Washing- ton State University; ZIL— Zoological In- stitute, University of Lund. The dendrogram (Fig. 1) showing the pos- sible phylogenetic relationships among the species in the grandis group has not been rigorously tested. Some of the characters used are highly variable within Anastrepha and resolution of their polarity within the grandis group will require a thorough cla- distic analysis of the entire genus. An ad- mittedly subjective hypothesis is presented here, which may be useful until a more com- prehensive analysis can be completed. DIAGNOSIS AND RELATIONSHIPS OF THE GRANDIS GROUP Steyskal (1977) defined the grandis spe- cies group in one of the couplets of his up- dated version of Stone’s (1942) key to the species of Anastrepha. He included those species that have what might be termed the “grandis-type”’ wing pattern. In this pattern 1) cell r, is entirely infuscated, lacking the normal marginal hyaline spot or band im- mediately apical to the apex of vein R,, 2) the S-band is complete, and 3) only the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON castilloi fumipennis grandicula ic ) ” [je O [= = oO o eS — 77) fo) atrigona bivittata Fig. 1. Possible phylogenetic relationships among the species of the grandis group. The following are possible synapomorphies: |) cell r, without marginal hyaline spot (note homoplasy in #7); 2) body color relatively dark; 3) cell r,., entirely infuscated; 4) cell bm at least weakly infuscated; 5) second section of vein M more than 2.9 times as long as third section; 6) aculeus tip broadly rounded; 7) cell r,; with marginal hyaline spot (reversal of #1). proximal arm of the V-band is present. In some specimens of grandis, the infuscation at the base of the marginal part of cell r, is faint, but there is never a distinctly bordered band or triangular hyaline spot as in most species of Anastrepha. A number of Anas- trepha species might be confused with spe- cies with the grandis-type pattern because they also lack the hyaline spot in r,. Most of the species of the daciformis group lack this r, spot, but they differ in having the S-band interrupted in cell dm, in having the basal half of the scutellum darker than the apex, and in a number of genitalic charac- ters. Some specimens of cordata Aldrich also lack the r, spot, but these have a complete V-band with the proximal arm blackish and broadly expanded. Six species of Anastrepha have the gran- dis-type wing pattern. The keys of Stone (1942) and Steyskal (1977) include grandis, VOLUME 93, NUMBER 1 atrigona Hendel, shannoni Stone, and bezzii Lima, as well as ba/loui Stone, here consid- ered a synonym of bezzii. Two new species, castilloi Norrbom and grandicula Norr- bom, also have this pattern. Despite the similarity of their wing patterns, the mo- nophyly of these six species is doubtful. Anastrepha bezzii is probably more closely related to species such as mucronota Stone, crebra Stone, and a number of others that have the endophallic sclerite of the disti- phallus narrowed and not distinctly con- voluted (Fig. 8A, B). The plesiomorphic shape of this sclerite (occurring in Toxotryp- ana Gerstaecker and most other Anastre- pha) is broad, with a distinct internal tube (Fig. 8C-E). The undulation of vein R, ,; is another apomorphic character found in many of these species, although it is fre- quently variable intraspecifically as in bez- zil. Although connected in cell r,, the sep- aration of the C- and S-bands along vein R,,; in bezzii is another similarity shared with these species, which usually have the bands completely separated. The connec- tion of the C- and S-bands is a highly vari- able character in Anastrepha, one that has certainly been affected by homoplasy, but it is noteworthy that in the other species with the grandis-type wing pattern, these bands are connected along R,.;. Like mucronota, crebra, and related species, bezzii also re- tains the plesiomorphic absence of lateral creases on the proctiger (Fig. 7C). They are present (Figs. 7E, 11A) in the other species with the grandis-type wing pattern as in the majority of Anastrepha species. The shape of the aculeus tip, the elongate pattern of dorsal scales on the eversible membrane, and the weakly sclerotized spermathecae, although distinctive in bezzii, are autapo- morphic and indicate little about its rela- tionships. One character that contradicts the hypothesis of its relationship to mucronota and related species is the posterior orbital seta, which is absent in bezzii and some- times weak or absent in grandis, shannoni, and grandicula. The variability of this char- 103 acter in the latter three species and the dis- tribution of the other character states dis- cussed above suggest that this is the result of homoplasy, however, like the similarity in wing pattern. On this basis I transferred bezzii from the grandis group to the mu- cronota group (Norrbom and Kim 1988b). Anastrepha grandis, shannoni, grandicu- la, and atrigona are probably closely relat- ed, although their monophyly is difficult to clearly demonstrate. Except for the shape of the aculeus tip, their female and male ter- minalia are very similar, as is their color- ation. The absence of the hyaline spot from r, and the basal part of r,,; and the absence of the apical arm of the V-band are probable synapomorphies, although neither of these character states (especially the latter) is unique to these species within Anastrepha. A monophyletic group including them should possibly also include two poorly known species, fumipennis Lima and bi- vittata (Macquart), which I have not ex- amined. Anastrepha fumipennis is known only from two females from Rio de Janeiro (Lima 1934) and a third from Espirito Santo (R. A. Zucchi, pers. comm.), whereas biv- ittata is currently unrecognized because the type material, from an unknown locality, has not been located (I did not find it in the UMO or the Museum National d’Histoire Naturelle, Paris). From Macquart’s (1843) description it appears to be closely related to or perhaps even conspecific with fumi- pennis, although Lima (1934) noted that the terminalia are shorter in fumipennis than is shown in Macquart’s illustration of bivit- tata. The scutal color pattern may also dif- ferentiate the two taxa, although neither au- thor thoroughly described this character. Both fumipennis and bivittata have a mar- ginal hyaline spot in cell r,, but their wings otherwise closely resemble atrigona; cell bm is infuscated, the apical arm of the V-band is absent, the proximal arm is close to the S-band in cell dm (at least in fumipennis), the pattern is brown, and the ratio of the second and third sections of vein M is great- 104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON er than 2.9. The aculeus tip in fumipennis is also almost identical to that of atrigona (its shape is unknown in bivittata). Except perhaps for the wing pattern color, these are all probable synapomorphies. One possible hypothesis of relationship among these three species and grandis, grandicula, and shan- noni to explain these character state distri- butions is that the loss of the r, hyaline spot is a Synapomorphy for the entire group, but it reappeared due to reversal in fumipennis and bivittata (Fig. 1). A more thorough study of the latter two species is needed to better understand their relationships. The relationship of castilloi to the other species with the grandis-type wing pattern is less certain. The presence of the proctiger creases indicates that it is not closely related to bezzii, but this is a synapomorphy shared with many Anastrepha species. The wing pattern (absence of r, hyaline spot and api- cal arm of V-band) and generally dark col- oration may be synapomorphies with the rest of the grandis group, but the much dif- ferent shape of the surstylus, the pattern of dorsal scales on the eversible membrane, and the scutal color pattern suggest that the similarities in wing pattern could be the re- sult of convergence. The latter three char- acter states might also be autapomorphic for castilloi, however. KEY TO THE SPECIES OF ANASTREPHA WITH THE GRANDIS-TYPE WING PATTERN The following key may be used in place of couplets 2—5 in the key of Stone (1942) or in place of that on page 7 in Steyskal (1977). 1. Wing (Fig. 2A, B) with a hyaline area in cell r,,, between crossveins bm-cu and r-m; section of vein M between bm-cu and r-m less than 1.85 times as long as section between r-m and dm-cu; vein R,,, usually undulant .. bezzii Lima — Wing (Fig. 2C-G) with cell r,,, entirely infus- cated basal to r-m; section of vein M between bm-cu and r-m more than 1.85 times as long as section between r-m and dm-cu; vein R,,, almost/Stralshtep rere ene eee nek 2 2. Cell r,,, entirely infuscated (Fig. 2E, G); tho- racic pleura with well differentiated yellowish and redbrown areas or mostly yellowish with brownish medial spot on anepimeron ....... Cell r,,, (Fig. 2C-D, F) with large subapical hyaline area; thoracic pleura without well dif- ferentiated dark areas except sometimes on greater ampulla, laterotergite, or metapleuron . Abdomen with medial yellowish stripe; cell br (Fig. 2G) entirely infuscated; dark areas of scu- tum more or less evenly redbrown (similar to Fig. 3A); syntergosternite 7 less than 4.0 mm long; aculeus tip (Fig. 13G) serrate, with pair of dorsal lobes; surstylus (Fig. 11C) elongate, slightly acute; aedeagus less than 5.0 mm long 5 ere ats 35 Feat ee shannoni Stone Abdomen entirely yellowish or orange; cell br (Fig. 2E) with large hyaline area apical to bm- cu; scutum (Fig. 3C) with irregular brown stripe, darker than orange or redbrown areas, slightly lateral to dorsocentral seta; syntergosternite 7 more than 4.75 mm long; aculeus tip (Fig. 13E, F) non-serrate, slender, with dorsal and ventral ridges; surstylus (Fig. 11B) moderately long, blunt; aedeagus more than 6.0 mm long .... RP sd PORN ed a a re grandis (Macquart) . Wing (Fig. 2C) with section of vein M between bm-cu and r-m more than 2.8 times as long as section between r-m and dm-cu; cell bm in- fuscated, at least weakly; proximal arm of V-band narrowly separated from or partially fused with S-band in cell dm; aculeus tip (Fig. 13A) non-serrate, at least 0.12 mm wide .... Se ee ate ne ood eak atrigona Hendel Wing (Fig. 2D, F) with section of vein M be- tween bm-cu and r-m less than 2.5 times as long as section between r-m and dm-cu; cell bm hyaline; proximal arm of V-band separated from S-band in cell dm by hyaline area at least as wide as arm of V-band; aculeus tip serrate Onless than! 0:07 mmiwid ele . Greater ampulla with distinct dark brown spot; scutum (Fig. 3B) with medial yellowish stripe broadly expanded posteriorly, fusing with nar- row Stripe in line with dorsocentral seta; syn- tergosternite 7 greater than 4.0 mm long; ever- sible membrane with single row of large dorsal scales (Fig. 10C); aculeus tip (Fig. 13C) non- serrate, lessithan O!O/smmbwides sass WU elt Seed Pe eee castilloi Norrbom, n. sp. Greater ampulla yellowish; scutum similar to Fig. 3A, with medial yellowish stripe narrow posteriorly, not extending laterally beyond ac- rostichal seta, narrow stripes in line with dor- socentral seta absent; syntergosternite 7 less than 4.0 mm long; eversible membrane with VOLUME 93, NUMBER 1 105 Fig. 2. Wing. A, A. bezzii, male. B, A. bezzii, female. C, A. atrigona. D, A. grandicula. E, A. grandis. F, A. castilloi. G, A. shannoni. numerous large scales in triangular pattern (Fig. 10D); aculeus tip (Fig. 13D) serrate, approxi- matelyOAldomim wider. -sasaaceeaaaseine Anastrepha atrigona Hendel (Figs. 2C, 3A, 4, 7A, B, LOA, 13A, 14A) Anastrepha atrigona Hendel 1914a: 70 (Su- rinam), 1914b: 15 (in key, catalog), 20 (description again as n. sp.); Lima 1934: 499, 555 (in key); Greene 1934: 132, 138 (in keys), 146 (Brazil); Stone 1942: 12 (in key), 21 (2 terminalia; Brazil); Fernandez 1953: 12 (in key), 18 (Venezuela); Foote 1967: 7 (in catalog); Steyskal 1977: 7 (in key); Zucchi 1978: 17 (in key), 27; Cara- ballo 1981: 30 (in key), 44 (Venezuela). Type data.—Holotype ¢ (NMW), “aus Surinam, Firma Staudinger & Bang-Haas.” It bears a label with “Surinam V.-IX.,” a label with “‘Anastrepha atrigona H. det. Hendel” in Hendel’s writing, and a red “Type” label. I have added a holotype label. Hendel apparently removed the right wing for illustration and it may be lost. I have 106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON TBG West Longitude Smithsonian Institution 1983 Prepared by Theoph* **'*' Griswold Fig. 4. Distributions of A. atrigona (dots = examined specimens; squares = literature records) and A. fu- mipennis (triangles). VOLUME 93, NUMBER | 107 2.8 13.0 bast Camp) 12.0 2.6 * camp 11.0 2.4 ce E E E =] : re) £ 10.0 5 3 = Steere a i e rr) i b o -_- Es So 7) a A A 2 5 ee Sai ° 8.0 = A . pp 7) g °, i) Os °, °. @, 1.8 e m 7.0 gv vi Vi . : e °, 6.0 1.6 A | 3.6 3.8 4.0 4.2 4.4 4.6 4.8 5.0 B 3.6 3.8 4.0 4.2 4.4 4.6 4.8 5.0 Mesonotum In. (in mm) Mesonotum In. (in mm) Fig. 5. A. bezzii: A, Syntergosternite 7 length (in mm) versus mesonotum length (in mm); B, Relative syntergosternite 7 length (syntergosternite 7 length/mesonotum length) versus mesonotum length. For data points, b = specimens from Brazil, g = Guatemala, m = Mexico, p = Panama, v = Venezuela. Line A = correlation based all specimens (except Campinas 8, see text), line B = correlation for specimens from Vicosa, Brazil only, line P = correlation based on specimens from Mexico-Panama only, and line V = correlation for specimens from Venezuela only. dissected the abdomen which is now stored in a microvial. Description.— Mostly orange to red- brown. Setae dark redbrown to black. Head: Concolorous except dark brown ocellar tu- bercle; 4—5 frontal setae; 2 orbital setae, pos- terior one always strong. Antenna extends 0.70-0.78 distance to lower facial margin. Thorax: Mesonotum (Fig. 3A) 3.30-4.08 mm long. Scutum entirely microtrichose; mostly orange to dark redbrown; single dis- tinct yellowish medial stripe from anterior margin to slightly posterior to acrostichal seta, slightly expanded posteriorly but usu- ally not extending laterally beyond acros- tichal seta; no narrow yellowish stripe in line with dorsocentral seta; rarely with poorly differentiated brown stripe slightly lateral to dorsocentral seta, but never as dark as in grandis; distinct yellowish sublateral stripe from transverse suture to posterior margin, crossing intra-alar seta; scuto-scu- tellar suture rarely with irregular dark brown medial spot. Subscutellum and medioter- gite redbrown, broadly dark brown laterally. Pleura with typical dark areas at most weak- ly differentiated. Katepisternal seta undif- ferentiated or weak, yellowish, no longer than postocellar seta. Wing (Fig. 2C): Length 7.50-9.22 mm. Vein R,,; almost straight. Vein M strongly curved apically; section be- tween bm-cu and r-m 2.91-—3.43 times as long as section between r-m and dm-cu. Pat- tern dark orange brown to dark brown, its margins distinct. Cell r,,; with large sub- apical hyaline area; entirely infuscated ba- sally. Cell br and base of cell dm entirely infuscated or with small hyaline area; if present in br just apical to bm-cu, this spot does not reach vein R,,; and extends no more than half way to dm-cu. Cell bm in- fuscated, often darkly, but sometimes only faintly yellowish. Proximal arm of V-band extends anteriorly to R,,;; fused to S-band 108 at R,,; and often also in cell dm. Abdomen: Tergites unicolorous orange to redbrown. Male terminalia: Outer surstylus (Fig. 7A, B) broad basally, tapering apically to narrow lobe; moderately long, section apical to apex of inner surstylus |.20-1.70 times as long as basal part; in lateral view similar to gran- dis. Proctiger with distinct lateral fold; scler- otized part narrowly divided into 3 sections. Aedeagus 5.10-5.76 mm long; 1.38-1.49 times as long as mesonotum. Distiphallus 0.58-0.63 mm long, relatively stout; en- dophallic sclerite similar to grandis, strong, stout and distinctly convoluted apically. Fe- male terminalia: Syntergosternite 7 3.70- 4.45 mm long; 1.08-1.18 times as long as mesonotum. Eversible membrane (Fig. 10A) with dorsobasal scales in triangular pattern; 10-11 large hooklike scales in broadest row; largest scales 0.20 mm long. Aculeus 3.55— 4.40 mm long; tip (Fig. 13A) 0.37-0.42 mm long, 0.13-0.14 mm wide, non-serrate, sides broadly convex. Spermathecae (Fig. 14A) ovoid to elongate ovoid. Distribution (Fig. 4).— Probably through- out the humid forests of Amazonia, the Guianas, and Venezuela. The exact locality within Surinam where the holotype was col- lected is unknown. It is represented on the distribution map by a dot in the center of the country. I examined four specimens with labels with “4385” and ‘‘Dem.,” one of which also has a label with “‘Demerara, Af- rica.” I am assuming that this is a mistake and that the site is the Demerara River in Guyana. No species of Anastrepha are known to be established in Africa. The distribution of atrigona appears dis- junct from that of its possible sister species, fumipennis (and/or bivittata), from south- ern Brazil (Fig. 4). It overlaps that of grandis only in northern Venezuela. Specimens examined.—Holotype (see Type data). BRAZIL: Amazon., H. W. Bates, 1 2 (BMNH). Amazonas: Amazon River, Atary to Manaus, 20-21.1X.1930, Holt, Blake & Agostini, 1 @ (USNM); | km E Campinas field station, Km 60 N Manaus, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 221979, L.EEewin, 16) (NBA lee (USNM); Manaus, Campus Univ., various dates, IT, XI—XII. 1975-1978, J. A. Rafael, 1 6 2 2 (INPA) 1 6 1 @ (USNM); Manaus, 2.VII.1986, 22.VIII.1985, B. Klein, 1 2 (INPA) 1 @ (USNM). Para: Santarem, H. W. Bates, 1 ¢ 1 2 (BMNH); Santarem, | 2 (CMP); Santarem, IX, 1 ?(USNM); Obidos, 1 6(AMNH); Dampfer [steamer] Prainha— Mte. Alegre, 30.V.1927, Zerny, 1 2 (NMW) 1 ¢ (USNM). Rio Branco: Mt. Roraima, 1932, J. G. Myers, 1 6 (MCZ) 1 2? (USNM). GUYANA: “Demerara, Africa, 4385,” O. S. Westcott, 2 6 (INHS) 1 6 1 2 (USNM). VENEZUELA: Amazonas: Cerro Duida, 6.111.1929, 1 2(AMNH). Aragua: El Limon, 450 m, 6.VII.1966, J. R. Dedordy, 1 2 (UCV); Rancho Grande, Portachuelo, 1100 m, 28.V.1980, J. A. Clavijo, 1 6 (UCV). Anastrepha bezzii Lima (Figs. ZA. 5B),5, 6,,7C, Db) SAB: 10B, 13B, 14B) Anastrepha Bezzii Lima 1934: 498 (descrip- tion, 6 genitalia; Brazil), 555 (in key); Lima 1937: 63 (description of ?; Brazil). Anastrepha bezzii; Stone 1942: 12 (in key), 20; Foote 1967: 8 (in catalog); Steyskal 1977: 7 Gn key); Zucchi 1978: 17 (in key), 33; Zucchi 1984: 561 (2 terminalia; Bra- zil). Anastrepha balloui Stone 1942: 12 (in key), 20 (description, 2° terminalia; Venezuela, Panama); Fernandez 1953: 12 (in key), 17 (host; Venezuela); Guagliumi 1966: 193, 197, 232 (hosts); Foote 1967: 7 (in catalog); Korytkowski and Ojeda 1968: 43 (6 terminalia), 1969: 79 (Peru); Stey- skal 1977: 7 (in key); Caraballo 1981: 30 (in key), 41 (taxonomy, hosts; Venezue- la); Aluja et al. 1987 (Mexico); Jiron et al. 1988: 132 (Costa Rica); Norrbom and Kim 1988b: 13 (hosts). New SYNONYMY. Type data.—bezzii: holotype 4 (Instituto Oswaldo Cruz, no. 1796), “Brazil, Rio de Janeiro, Manguinhos, 24.II.1902.”’ Zucchi (1978) reexamined it and reported its date VOLUME 93, NUMBER 1 109 800 1000 TBG. West Longitude Fig. 6. Distribution of 4. bezzii (dots = examined specimens; squares = literature records). Smithsonian Institution 1983 Prepared by Theophilus Brit! Grivwold 110 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON | | A Ne a i ey) He = z eS (\ NY \ .y e B Fig. 7. A, B, A. atrigona. C, D, A. bezzii. E, F, A. castilloi. A, B, D, F, epandrium and surstyli (posterior view). C,E, epandrium, surstyli, and proctiger (lateral view). Bar = 0.1 mm. Fig. 8. Distiphallus (lateral view except B, ventral). A, B, A. bezzii. C, A. castilloi. D, A. grandis. E, A. shannoni. Bar = 0.1 mm. VOLUME 93, NUMBER 1 a TBG. 30 West Longitude Smithsonian Institution 1983 pared by Theophilus Brit! Griswold Fig. 9. Distributions of A. castilloi (dot), A. grandicula (square), and A. shannoni (triangles); solid figures = examined specimens; hollow = literature records. of collection as 24.III.1917. Lima dissected and mounted the abdomen and aswing on slides 1876 and 1877. balloui: holotype ? (USNM), “Venezuela: San Juan de los Mo- rros, on Terminalia catappa, 12.1V.1938, C. H. Ballou.” It bears a label with “S. J. los Morros, Venez, 249 C. H. Ballou, 4-12- 1938” and ‘‘on Terminalia catappa,” a red “Type No. 51652 U.S.N.M.” label, and a Stone determination label with “‘Anastre- pha balloui Stone.” Stone dissected and mounted the terminalia on slide 39.I1V.17°. Description.—Mostly yellowish to or- ange. Setae redbrown to dark redbrown. Head: Concolorous except dark brown ocel- lar tubercle; 2-4 frontal setae; | (rarely 2) orbital seta (posterior seta present, only on left side, in only | of 38 specimens exam- ined). Antenna extends 0.50 distance to lower facial margin. Thorax: Mesonotum 112 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 10. Dorsal scales of eversible membrane. A, A. atrigona. B, A. bezzii (redrawn from Stone 1942, at slight lateral view). C, A. castilloi. D, A. grandicula. E, A. grandis. F, A. shannoni. ok Fig. 11. A,B, A. grandis. C, A. shannoni. A, epandrium, surstyli, and proctiger (lateral view). B, C, epandrium and surstyli (posterior view). VOLUME 93, NUMBER 1 113 West Longitude Smithsonian Institution 1983 . «Brit Griswold Fig. 12. Distribution of A. grandis (dots = examined specimens; squares = literature records). 3.66-4.52 mm long (up to 5.00 in Brazil (Zucchi 1984)). Scutum entirely microtrich- ose; mostly yellowish to orange with indis- tinct whitish or translucent areas; pattern difficult to see, similar to Fig. 3B, but trans- lucent medial stripe hardly expanded until about halfway between transverse suture and posterior margin, and then abruptly so; pos- terior part of medial stripe larger and more rectangular than in castilloi, fusing with nar- row translucent stripe extending from an- terior margin in line with dorsocentral seta; scuto-scutellar suture without distinct me- dial dark brown spot. Subscutellum and me- diotergite entirely yellow to orange. Pleura with typical dark areas poorly differentiat- ed. Katepisternal seta undifferentiated or weak, yellowish, no longer than postocellar seta. Wing (Fig. 2A, B): Length 8.39-10.29 mm (up to 10.75 in Brazil (Lima (1937)). Vein R,,; almost straight to strongly un- dulant. Vein M strongly curved apically, 114 Eig 3% grandicula. E, F, A. grandis. G, A. shannoni. Bar = 0.10 mm. reaching apex of S-band; section between bm-cu and r-m 1.48-1.79 times as long as section between r-m and dm-cu. Pattern faint yellow to faint orange brown, its mar- gins often diffuse. Cells dm, br and r,,,; crossed by hyaline area between bm-cu and r-m, rarely narrowly entering cell r,. Cell r,,, also with large subapical hyaline area. Cell bm hyaline. Proximal arm of V-band strong until vein M, usually extending faint- ly to R,,5; separate from S-band. Abdomen: Tergites unicolorous yellowish to orange. Male terminalia: Outer surstylus (Fig. 7D) strongly delimited from epandrium, broad basally, gradually tapering to rounded apex; relatively short, section apical to apex of inner surstylus 0.60-0.70 times as long as basal part; posterior side slightly concave (Fig. 7C). Proctiger (Fig. 7C) without dis- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Aculeus tip (ventral view except E, lateral). A, A. atrigona. B, A. bezzii. C, A. castilloi. D, A. tinct lateral fold; sclerotized part undivided. Aedeagus 9.71-12.10 mm long; 2.56-2.83 times as long as mesonotum. Distiphallus (Fig. 8A, B) 0.65—0.80 mm long, relatively stout; endophallic sclerite strong, moder- ately broad, but not distinctly convoluted apically, a subapical hooklike structure usu- ally apparent. Female terminalia: Synter- gosternite 7 6.28—9.55 mm long (as short as 5.56 mm in Venezuela (Caraballo 1981), up to 12.50 mm in Brazil (Lima 1937)); 1.63- 2.16 times as long as mesonotum (up to 2.63 times in Brazil (Lima 1937)). Eversible membrane (Fig. 10B) with dorsobasal scales in elongate pattern 1.85-—3.33 mm long (up to 4.00 in Brazil (Zucchi 1984)); largest scales at basal third, 0.08—0.09 mm long. Aculeus 5.62-9.13 mm long (up to 10.00 in Brazil (Zucchi 1984)); tip (Fig. 13B)0.25-0.35 mm VOLUME 93, NUMBER 1 115 Fig. 14. A-F, spermathecae (2 of 3 shown). G, eggs. A, A. atrigona. B, A. bezzii. C, A. castillot. D, A. grandicula. E, A. grandis. F, G, A. shannoni. long, 0.09-0.10 mm wide, non-serrate, sides slightly convex, ventral surface transversely concave with apical 4 slightly convex. Sper- mathecae (Fig. 14B) weakly sclerotized, elongate ovoid. Remarks.—There is considerable varia- tion in terminalia length among specimens that are recognized here as bezzii (Table 1, Fig. 5). Stone (1942) and subsequent au- thors recognized populations from Mexico to Venezuela and Peru as a distinct species, A. balloui, because they have syntergoster- nite 7, the aculeus, the part of the eversible membrane with dorsal scales (the “‘rasper”’ of Stone 1942), and the aedeagus shorter than in the few Brazilian specimens previ- ously known. The lengths of all of these structures are correlated, however, and these differences are all the result of variation in terminalia length. The distinctive shape of the aculeus tip, the pattern of the scales of the eversible membrane, the shape of the surstyli, and the wing pattern and venation are consistent in the Brazilian and non-Bra- zilian specimens. Table 1 and Fig. 5 present an analysis of Table 1. Anastrepha bezzii: mesonotum length (in mm), syntergosternite 7 length (in mm), and their ratio in specimens from Mexico—Panama, Venezuela, and Brazil. Mexico—Panama (n = 10) Venezuela (n = 8) Brazil (n = 6) Mesonotum length 3.66-4.53 Mean and std. dev. 4.09 + 0.33 Syntergost. 7 In. 6.67-8.48 Mean and std. dev. TO O69 Synt. 7/meson. In. 1.80-1.96 Mean and std. dev. 1.90 + 0.05 3.70-4.48 4.08-5.00 By CW9) ae (078) 4.54 + 0.32 6.28-7.30 8.07-10.00 6.70 + 0.29 9.09 + 0.72 163=1-75 1.90-2.17 1.68 + 0.04 2.01 + 0.09 SS ee ——___ nnn ——_—-” 116 the variation in the length and relative length (ratio to mesonotum length) of syntergo- sternite 7 between samples from Mexico- Panama, Venezuela, and Brazil. All measurements were taken from examined specimens with the addition of those for single females from Vicosa, Brazil (Zucchi 1984) and Venezuela (Caraballo 1981). The measurements of the female reported by Lima (1937) from Campinas, Goias, Brazil, which unfortunately has been lost (Zucchi 1984), are plotted for comparison, but are not included in the analysis. Caraballo (1981) found greater variation in synter- gosternite 7 length in a larger sample of fe- males from Venezuela (5.56-—7.30 mm; n = 75). She did not report the relative length, but her range of mesonotal lengths suggests that it is consistent with that in the speci- mens I examined. Syntergosternite 7 length and relative length are significantly different (¢ test, P < 0.01) between all samples, as is the arcsine transformation of the inverse of the latter ratio. Mesonotum length is significantly dif- ferent between Brazil and Venezuela (P < 0.01) and Brazil and Mexico-Panama (P < 0.02), but not between Venezuela and Mex- ico-Panama. Thus, although the Brazilian females have longer terminalia, their rela- tive terminalia length is closer to that of the Mexico-Panama sample than to that of the Venezuela sample or the Campinas, Brazil female of Lima (1937) (Fig. 5B). Similarly, the regression lines for these two samples are closer than that for the Venezuela sam- ple (Fig. SA). There does not appear to be an allometric relationship, at least within samples (Fig. 5B). I have not examined the male holotype of bezzii which is assumed to be conspecific with the Vicosa sample. The type locality of Rio de Janeiro is much closer to Vicosa than to Campinas. At least the Mexico-Pan- ama populations seem to be conspecific, and although the relative terminalia length of the Venezuelan sample is significantly dif- ferent statistically, whether there is any bi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ological significance is questionable. The status of the Campinas female is less certain since Lima’s measurements cannot be ver- ified, but if they are accurate, it is at least as different from the other Brazilian speci- mens as the Venezuela population. I do not believe there is enough evidence at the present time to justify the recognition of the Venezuelan population as a distinct species, and I therefore regard balloui Stone a synonym of bezzii. Continued use of this name is not justifiable without also regard- ing the Campinas female as representative of another separate species. Unless other characters or behavioral or biological stud- ies demonstrate otherwise, I believe that all of these specimens should be considered conspecific. Additional collecting, an anal- ysis of larger sample sizes, and biochemical or behavioral studies to test the statistical and biological significance of the differences among them would certainly be useful. The scutal color pattern in bezzii is faint and difficult to see. It was obscured by un- derlying tissues in most of the specimens I examined, most of which were preserved in fluid prior to pinning. Biology.—The only well confirmed hosts are Sterculia apetala (Jacq.) Karst. (Cara- ballo 1981) and S. chicha St. Hil. (R. A. Zucchi, pers. comm.), although Guagliumi (1966) also listed Terminalia catappa L. as a host. The latter record may be erroneously based on Stone’s (1942) report that some of the types of ba/loui were collected ‘‘on” T. catappa, but Guagliumi’s listing (p. 232) of two Opius species as “‘Parassita di Anastre- pha balloui Stone s/Terminalia” suggests that he may have seen reared specimens. The larvae reared from S. chicha in Brazil fed inside the seed rather than in the pulp of the fruit (R. A. Zucchi, pers. comm.). Distribution. — Southern Brazil, Peru and Venezuela to Mexico (Fig. 7). Specimens examined.— Holotype of bal- loui (see Type data). BRAZIL: Minas Ger- ais: Vicosa, “ex. Sterculia chicha (seed),” 20.VII.1987, G. P. Santos, 5 65 2@(USNM). VOLUME 93, NUMBER 1 COSTA RICA: Alajuela: Orotina, L. F. Ji- ron, 1 @ (UCRSJ). Puntarenas: Lepanto, 8.1.1986, L. F. Jiron & J. Soto, 1 ¢(USNM). GUATEMALA: Guazacapan, Sta. Rosa, XI.1986, E. Muniz, 1 2 (USNM). MEXI- CO: Chiapas: Soconusco Region, | 4 1 2 (PMMT); Soconusco Region, I.1984, M. Aluja, 1 6 (USNM). PANAMA: Barro Col- orado Isld., V-VI.1943, J. Zetek, 1 6 1 9 (USNM); same, 4.I]I.1967, R. D. Akre, 1 ° (WSU); Arraiyan, [HI.1950, 1951, J. Zetek, 1 6 1 2(USNM); El Cermeno, various dates I, VI, VI-VU, [X—X.1939-1942, J. Zetek, 6 6321 2 paratype (USNM) | 2 (NMW),. VENEZUELA: Aragua: Gonzalito, 3.11. 1950, F. Fernandez-Yepez, | 2 (BMNH); El Limon, 450 m, 5.VIII.1981, G. Yepez, | @ (UCV); Maracay, 17.VII.1945, F. Fernan- dez-Yepez, 1 2 (CUI). Guarico: San Juan de los Morros, ‘‘on Terminalia catappa,” 12- 28.1V.1938, C. H. Ballou, 6 ¢ 3 2 paratypes (USMN) 1 ¢ 1 2 paratypes (BMNH) | ¢ paratype (NMW). Anastrepha castilloi Norrbom, NEw SPECIES (rigs. ZE. 3B: 7B. EAs C 40: 10C, 13C, 14C) Type data.—Holotype 2 (UCV), VENE- ZUELA: Bolivar: Auyan-tepui, campamen- to A, 5°44.8’N, 62°29.5’'W, 2200 m, 10- 13.11.1988, L. J. Joly & A. Chacon. I have dissected the terminalia which are now stored in a microvial. Paratypes: same data as holotype, 7 6 1 ¢(UCV), 442 2(USNM). Description.— Mostly orange to dark brown. Setae dark redbrown to black. Head: Concolorous except dark brown ocellar tu- bercle; 4-6 frontal setae; 2 orbital setae, pos- terior one strong. Antenna extends 0.75- 0.86 distance to lower facial margin. Tho- rax: Mesonotum (Fig. 3B) 3.33-4.20 mm long. Scutum entirely microtrichose; mostly moderate to dark brown; single distinct yel- lowish medial stripe from anterior margin to slightly posterior to acrostichal seta, broadly expanded and rounded (not rect- angular) posteriorly, extending laterally to 17 or almost to dorsocentral seta, fusing with narrow, faint orange stripe extending from anterior margin to dorsocentral seta; dis- tinct yellowish sublateral stripe from trans- verse suture to posterior margin, crossing intra-alar seta; distinct yellowish mark ex- tending posteriorly from postpronotal lobe to presutural supra-alar seta, almost joining sublateral stripe on transverse suture; scuto- scutellar suture without distinct medial dark brown spot. Subscutellum and mediotergite light brown, broadly dark brown laterally. Pleura with typical dark areas weakly to moderately differentiated except spots on laterotergite and metapleuron dark brown; greater ampulla mostly dark brown. Kat- episternal seta weak, orange, smaller than postocellar seta. Wing (Fig. 2F): Length 7.65-9.22 mm. Vein R,,,; almost straight. Vein M weakly to moderately curved api- cally; section between bm-cu and r-m 1.96- 2.34 times as long as section between r-m and dm-cu. Pattern orange brown, its mar- gins distinct. Cell r,,, with large subapical hyaline area; entirely infuscated basally. Cell br with large hyaline spot apical to dm- cu, extending more than halfway to r-m. Cell dm with large basal hyaline spot. Cell bm hyaline. Proximal arm of V-band ends at M, broadly separated from S-band. Ab- domen: Tergites unicolorous yellowish to light brown. Male terminalia: Outer sursty- lus (Fig. 7F) rapidly tapering basally, blunt apically; short, section apical to apex of in- ner surstylus 0.45—0.55 times as long as bas- al section; in lateral view (Fig. 7E), posterior margin almost straight. Proctiger (Fig. 7E) setose dorsally, with distinct lateral fold; sclerotized part narrowly divided into 3 sec- tions. Aedeagus 5.47—5.95 mm long; 1.33- 1.54 times as long as mesonotum. Disti- phallus (Fig. 8C) 0.59-0.71 mm long, slen- der; endophallic sclerite weakly sclerotized and slender, but similar in shape to grandis, stout and distinctly convoluted apically. Fe- male terminalia: Syntergosternite 7 4.49- 4.95 mm long; 1.21—1.35 times as long as mesonotum. Eversible membrane (Fig. 10C) 118 with dorsobasal scales minute except 10-11 large hooklike scales in single row, separat- ed from minute scales by membranous area; largest scales 0.25 mm long. Aculeus 3.95- 4.53 mm long; shaft 0.05—0.06 mm wide at midpoint, sides parallel; tip (Fig. 13C) 0.40- 0.43 mm long, 0.05—0.06 mm wide, non- serrate, sides slightly convex. Spermathecae (Fig. 14C) globose. Distribution (Fig. 9).—Known only from the type locality. Etymology.—At the suggestion of one of the collectors of the type series, Prof. Luis Joly, this species is named for Dr. Carlos Castillo, an enthusiastic conservationist and the President of Maraven, the company which organized the expedition to Auyan- tepul. Anastrepha grandicula Norrbom, NEw SPECIES (Figs: 2D? 9. 10D; 13D; 14D) Type data.—Holotype ¢ (FSCA), CO- LOMBIA: Amazonas: Rio Putumayo, Puerto Arica, 8.VII.1978, M. A. Tidwell. I have slide mounted the right wing and dis- sected the abdomen which is now stored in a microvial. Description.— Mostly orange brown. Se- tae dark redbrown to black. Head: Concol- orous except dark brown ocellar tubercle; 3 frontal setae; 2 orbital setae, posterior one weak. Antenna extends 0.65 distance to lower facial margin. Thorax: Mesonotum 3.74 mm long. Scutum entirely microtrich- ose; mostly orange brown; pattern similar to Fig. 3A, single distinct yellowish medial stripe from anterior margin to slightly pos- terior to acrostichal seta, slightly expanded posteriorly but not extending laterally be- yond acrostichal seta; no narrow yellowish stripe in line with dorsocentral seta or dark brown stripe slightly lateral to dorsocentral seta; distinct yellowish sublateral stripe from transverse suture to posterior margin, cross- ing intra-alar seta; scuto-scutellar suture without distinct medial dark brown spot. Subscutellum and mediotergite orange PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON brown, broadly dark brown laterally. Pleura with typical dark areas weakly differenti- ated. Katepisternal seta weak, yellowish, subequal to postocellar seta. Wing (Fig. 2D): Length 7.98 mm. Vein R,,, almost straight. Vein M strongly curved apically; section be- tween bm-cu and r-m 2.29 times as long as section between r-m and dm-cu. Pattern or- ange brown, its margins distinct. Cell r,,; with large subapical hyaline area; entirely infuscated basally. Cell br with large hyaline spot apical to dm-cu, extending more than halfway to r-m. Cell dm with large basal hyaline spot. Cell bm hyaline. Proximal arm of V-band extends anteriorly to R,,;, but not joined to S-band. Abdomen: Tergites unicolorous yellowish. Female terminalia: Syntergosternite 7 3.62 mm long; 0.97 times as long as mesonotum. Eversible membrane (Fig. 10D) with dorsobasal scales in trian- gular pattern; 8 large hooklike scales in broadest row; largest scales 0.20 mm long. Aculeus 3.29 mm long; shaft 0.14 mm wide at midpoint, sides parallel; tip (Fig. 13D) 0.41 mm long, 0.14 mm wide, apical half serrate, sides very slightly concave in basal half, evenly tapering, straight in apical half. Spermathecae (Fig. 14D) ovoid. Distribution (Fig. 9).— Known only from the type locality. Etymology.—The epithet, which means “rather large’ in Latin, was selected because of the similarity of this species with grandis and its slightly smaller female terminalia. Anastrepha grandis (Macquart) (Figs. 2E, 3C, 8D, 10E, 11A, B, 12, 13E, F, 14E) Tephritis grandis Macquart 1846: 340 (de- scription; Colombia or Venezuela). Trypeta (Acrotoxa) grandis; Loew 1873: 231. Anastrepha grandis; Bezzi 1909: 284 (type data), 286 (in key); Hendel 1914a: 69, 1914b: 14 (in key), 15 (in catalog); Bezzi 1919a: 4 (in key), 6 (description; Para- guay, Brazil); Bezzi 1919b: 374; Lima 1930: 21 (hosts; Brazil); Fischer 1932: 302 (description, host, larva; Brazil), 1934: 18 VOLUME 93, NUMBER 1 (description, hosts; Brazil); Lima 1934: 496 (6 terminalia, hosts; Brazil), 555 (in key); Greene 1934: 132, 138 (in keys), 145 (hosts; Paraguay, Brazil; see also shan- noni); Blanchard 1937: 41 (Argentina); Stone 1942: 12 (in key), 22 (2 terminalia, hosts; Panama); Fernandez 1953: 12 (in key), 19; Blanchard 1961: 288 (in key), 297 (Argentina); Foote 1967: 11 (in cat- alog); Korytkowski and Ojeda 1968: 37 (hosts), 43, 1969: 78-79 (Peru); Steyskal 1977: 7 (in key); Zucchi 1978: 17 (in key), 56 (Brazil); Malavasi et al. 1980: 11 (host; Brazil); Caraballo 1981: 30 (in key), 46 (description, hosts; Venezuela); Boscan et al. 1980: 55 (host; Venezuela); Norrbom and Kim 1988b: 28 (hosts); Steck and Wharton 1988: 999 (larvae, egg); Nasci- mento et al. 1988: 54 (biology). Anastrepha schineri Hendel 1914a: 69 (Bo- livia), 1914b: 15 (in key), 16 (in catalog), 19 (description again as n. sp.); Fischer 1932: 303 (synonymy); Greene 1934: 145; Stone 1942: 23; Foote 1967: 11; Koryt- kowski and Ojeda 1968: 43; Zucchi 1978: 56: Anastrepha latifasciata Hering: 1935: 227 (description; Brazil); Lima 1937: 63 (syn- onymy); Stone 1942: 23; Foote 1967: 11; Korytkowski and Ojeda 1968: 43; Zucchi 1978: 56. Type data.—T. grandis: Lectotype (here designated) 2? (UMO), “De la Nouvelle-Gre- nade. Collection de M. Bigot.” The lecto- type was the only putative syntype of T. grandis 1 found in the exotic cabinet of the Bigot Collection (UMO). It bears a label with “Tephritis grandis n. sp.”” in Mac- quart’s writing and “°, Nov. Granata, Macq. D. Exot. nom.” in Bigot’s writing. It is miss- ing both wings, but is clearly recognizable as this species by its body color and ter- minalia, especially the tip of the aculeus, which is slightly projecting as is common in this species and as is shown in Macquart’s illustration (tab. 18, fig. 14). Nueva Gra- nada was the Spanish colony including Co- IS lombia and Venezuela. 4. schineri: Holo- type 6 (TMB), “aus Bolivia, Coroica.” It has a label with “‘Bolivia Coroico,” a label with ‘*Anastrepha schineri H. typus det. Hendel” in Hendel’s writing, and an orange bordered “typus” label. A. /atifasciata: Holotype 2 (PAN), “‘von Sta. Catharina, Sammler Li- derwaldt.”’ The type locality is the state or perhaps the island of Santa Catarina, Brazil. The holotype has the following labels: “‘S. Catarina, Liiderwaldt” (green), ““Type”’ (or- ange), ““Anastrepha 2 det. Dr. Enderlein,” and “Anastrepha latifasciata m. Type, det. M. Hering 1935.” I added holotype and de- termination labels. Description.— Mostly orange to red- brown. Setae usually moderate redbrown. Head: Concolorous except dark brown ocel- lar tubercle; 3-5 frontal setae; 1-2 orbital setae, posterior one often weak when pres- ent. Antenna extends 0.60-0.72 distance to lower facial margin. Thorax: Mesonotum (Fig. 3C) 2.88-4.22 mm long. Scutum en- tirely microtrichose; mostly orange to red- brown; single distinct yellowish medial stripe from anterior margin to slightly posterior to acrostichal seta, moderately expanded posteriorly, extending laterally beyond acrostichal seta but not reaching dorsocen- tral seta; no narrow yellowish stripe in line with dorsocentral seta, but dark brown, sometimes poorly differentiated stripe slightly lateral to dorsocentral seta; distinct yellowish sublateral stripe from transverse suture to posterior margin, crossing intra- alar seta; scuto-scutellar suture usually with irregular medial dark brown spot. Subscu- tellum and mediotergite orange brown, broadly dark brown to black laterally. Pleu- ra with typical dark areas weakly differen- tiated except anepimeron usually with dark brown spot. Katepisternal seta undifferen- tiated or weak, yellowish, no longer than postocellar seta. Wing (Fig. 2E): Length 7.95-10.30 mm. Vein R,,,; almost straight. Vein M moderately curved apically; section between bm-cu and r-m 2.18—2.69 times as long as section between r-m and dm-cu. Pat- —_ 120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tern faint yellow brown to orange brown, its margins often diffuse. Cell r,,, entirely infuscated. Cell br with large hyaline area apical to bm-cu, extending more than half- way to r-m. Cell dm with large basal hyaline area. Cell bm hyaline. Proximal arm of V- band ending at M or extending anteriorly, often faintly, to R,,; to fuse with S-band. Abdomen: Tergites unicolorous yellowish to orange. Male terminalia: Outer surstylus (Fig. 11B) broad basally, tapering apically to short blunt lobe; moderately long, section apical to apex of inner surstylus 0.95-1.33 times as long as basal section; posterior side distinctly concave (Fig. 1 1A). Proctiger with distinct lateral fold; sclerotized part narrow- ly divided into 3 sections or only weakly connected (Fig. 11A). Aedeagus 6.70-7.40 mm long; 1.70-2.33 times as long as me- sonotum. Distiphallus (Fig. 8D) 0.55-0.68 mm long, relatively stout; endophallic scler- ite strong, stout and distinctly convoluted apically. Female terminalia: Syntergoster- nite 7 4.99-6.28 mm long; 1.40-1.59 times as long as mesonotum. Eversible membrane (Fig. 1OE) with dorsobasal scales in broad subtriangular pattern; 14-16 large hooklike scales in broadest row; largest scales 0.30 mm long. Aculeus 5.27-6.18 mm long, of- ten as long or longer than syntergosternite 7; tip (Fig. 13E, F) 0.58-0.66 mm long, 0.16- 0.18 mm wide, non-serrate, sides slightly convex, ventral surface with V-shaped ridge subbasally, dorsal surface with V-shaped ridge at midlength. Spermathecae (Fig. 14E) ovoid to elongate ovoid. Remarks.— The aculeus was broken in the female with the shortest syntergosternite 7 and was probably no more than 5.00 mm long. Distribution (Fig. 12).—Southern Brazil, northern Argentina, Paraguay, and along the Andean Cordillera from Bolivia to Vene- zuela. Apparently grandis does not occur in the Guianas, Amazonia, and northeastern Brazil. In the latter country it has been col- lected in Bahia (Bondar 1950; exact locality not reported and not shown in Fig. 12), Mato Grosso, and Goias (Zucchi 1988), but it has not been detected north of these states, in- cluding in Rio Grande do Norte, where an extensive monitoring program has been op- erating since 1986 (Nascimento et al. 1988, A. Malavasi, pers. comm.). In Bolivia and northward, grandis appears restricted to lower elevations of the Andes and bordering areas. It has not been detected in the low- land coastal melon-growing region of Ec- uador (Harper 1987). The record from Pan- ama (Stone 1942) is questionable. The single female in the USNM on which it was prob- ably based was intercepted from a plane or ship from Panama, but it is not certain that the cargo in which it was found originated in that country. Biology.—The only normal hosts native to the Neotropics are Cucurbita pepo L., C. moschata (Duchesne) Poiret, and C. max- ima Duchesne (Fischer 1932, Malavasi et al. 1980, Caraballo 1981, Boscan et al. 1980). In addition, grandis has been reported to attack introduced cucurbits such as water- melon (Citrullus lanatus var. lanatus (Thunb.) Matsum. & Nakai) (Lima 1934, Stone 1942), melon (Cucumis melo L.) (Korytkowski & Ojeda 1968), cucumber (Cucumis sativus L.) (Lima 1934, Stone 1942), and calabash gourd (Lagenaria si- ceraria (Mol.) Standl.) (Caraballo 1981). Fi- scher (1934) reared one specimen from a guava fruit, but considered it an abnormal record because the fruit came from a tree in the middle of a field planted with Cu- curbita. Despite the comments of Koryt- kowski and Ojeda (1969), records from Citrus are doubtful. Host records are sum- marized by Norrbom and Kim (1988b). Nascimento et al. (1988) report various bi- ological parameters involving oviposition and duration of life stages based on studies with laboratory cultures in Sao Paulo. Specimens examined.—Lectotype of grandis; holotypes of latifasciata and schi- neri (see Type data). ARGENTINA: Mi- siones: Montecarlo, 4. VI.1982, G. Putruele, 3622(INTA) 1 61 2(USNM). BOLIVIA: VOLUME 93, NUMBER 1 Mapiri, Sarampioni, 700 m, 1.III.1903, 1} (SMT); Sucre, 17.VI.1966, J. C. Ballard, 1 @ (USU); Yungas von Coroico, 1800 m, 7.XII.1906, 1 2 (SMT). BRAZIL: Espirito Santo: Espirito Santo, Fruhstorfer, 1 ¢ (DEI). Goias: Anapolis, various dates V- VII,XI1.1936, G. Fairchild, 4 6 (MCZ) 2 ¢ (USNM); 24 km E Formoso, 25.V.1956, F. S. Truxal, 2 ?(LACM). Mato Grosso: Cam- po Grande, X.1947, A. Maller, 1 6 1 @ (USNM); Chapada [probably Chapada dos Guimaraes], XI, 1 6 (AMNH); Maracaju, V.1937, 2 6 2 2 (USNM). Minas Gerais: Vicosa, “reared from squash,” XI.1928, M. Kisliuk & C. E. Cooley, 1 ¢ 2 2 (USNM). Parana: Rolandia (Caviuna), II,III,V- VII,X.1946-1947, A. Maller, 5 4 6 2 (AMNH) | ¢ 1 2 (USNM). Rio de Janeiro: Nilopolis, 18.X1.1931, M. Kisliuk & C. E. Cooley, 1 6 (NMW) 1 2° (USNM); Nova Iguacu, 16.XI.1931, M. Kishiuk & C. E. Cooley, | 2 (NMW); Rio de Janeiro, “bred in fruits of Cucurbitaceae,” 1927, 2462 2 (MCZ); Rio de Janeiro, reared ex water- melon, P. 22.XI, Em. 11.XII.1931, M. Kis- liuk & C. E. Cooley, 2 ¢ 3 2 (USNM); Tin- gua, III.1940, R. C. Shannon, | e (USNM). Rio Grande do Sul, VII.1930, M. Kisliuk & C. E. Cooley, 1 specimen without abdo- men (USNM). Santa Catarina: Corupa (Hansa Humboldt), various. dates I, VI, VII.1946, A. Maller, 2 6 1 ? (AMNH) 1 2(USNM); Nova Teutonia, various dates, all months, various years 1936-1973, F. Plaumann, 6 6 22 2? (CNC) 6 ¢ 6 2 (USNM) 24642 (AMNH) | ¢ 3 2? (MCZ) 241 2 (BMNH) 2 ¢ 2 2 (FMNH). Sao Paulo: Sto. Amaro, “larva colhida de abobora, crysal. 24.1V.1933, eclosao 5.VI.1933,”’ Fischer, 1 6 2 2 (DEI); Sao Paulo, 26.XII.1931, M. Kisliuk & C. E. Cooley, 1 ¢(USNM); Univ. Sao Paulo, laboratory culture of A. Mala- vasi, reared ex “pumpkin,” XI.1986, G. J. Steck, 1 6 1 2 (USNM). COLOMBIA: An- tioquia: Santa Barbara, 5.II.1988, A. Bus- tillo, 1 2 (USNM). Boyaca: Muzo, 900 m, J. Bequaert, | ¢ (MCZ). Cauca: Santander, 26.1.1989, A. Trochez, 1 6 1 @ (USNM). {21 Santander: Bucaramango, 20.XII1.1988, A. Trochez, | ¢ 1 2 (USNM). Valle: Florida, 18.1V.1988, A. Trochez, 1 2 (USNM). EC- UADOR: Loja: Malacatos, 26.XII.1980, C. Pacheco, 1 2 (USNM). PANAMA, inter- cepted at San Francisco, California, 18.11.1937, 1 2 (USNM). PARAGUAY: Alto Parana: Hohenan, IV.1938, F. Schade, 1 6 1 2 (USNM); Bella Vista, VI.1938, F. Schade, 1 6 (USNM); Bella Vista, I, 1 2 (MCZ). Concepcion: Horqueta, A. Schulze, 1 2(USNM). Guaira: Villarrica, V.1940, F. Schade, 1 @ (USNM). La Cordillera: San Bernardino, K. Fiebrig, 1 2(USNM). PERU: Cuzco: Quillabamba, 23-27.XII.1983, L. Huggert, 1 @ (ZIL); [Rio] Mishagua—Uru- bamba fl. [junction], 13.X.1903, 1 ¢(SMT). VENEZUELA: Aragua: Rancho Grande, 1100 m, 22—24.VI.1984, D. S. Bogar, 1 ¢ (USNM). Carabobo: Las Trincheras, 700 m, SOMMELSS ED: Oterosioi(WiEv): Anastrepha shannoni Stone (Figs: 2G) 8E, 9. 1T0F NIG MIsG. I4EG) (part) Anastrepha grandis; Greene 1934: 145 (misidentification). Anastrepha shannoni Stone 1942: 12 (in key), 23 (description, Peru); Foote 1967: 16 (in catalog); Steyskal 1977: 7 (in key). (?)Anastrepha shannoni; Korytkowski and Ojeda 1968: 49 (2 terminalia), 1969: 78 (Berm): Type data.— Holotype 6 (USNM). It bears a label with ““Chimbotes, Amaz. Peru,” a label with “R. C. Shannon Coll. Mar.20.31,” a red “Type No. 51653 U.S.N.M.”’ label, and a Stone determination label with ‘‘Anastrepha shannoni Stone.” Stone dis- sected and mounted the left wing and ab- domen on slides 39.VIII.4b and 39.II1.9b, respectively. He reported the type locality as ““Chimbotes, Amazonas, Peru,” but it is probably the town of Chimbote, at 3°52’S 70°45'W in Loreto Department. The ‘**Amaz.” on the label probably refers to the Amazon River, along which this town is located, rather than Amazonas Depart- ment. 122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Description.— Mostly redbrown to dark redbrown. Setae dark redbrown to black. Head: Concolorous except dark brown ocel- lar tubercle; 3-4 frontal setae; 2 orbital se- tae, posterior one usually weak. Antenna extends 0.70-0.80 distance to lower facial margin. Thorax: Mesonotum 3.65-3.90 mm long. Scutum entirely microtrichose; mostly redbrown to dark redbrown; single distinct yellowish medial stripe from anterior mar- gin to slightly posterior to acrostichal seta, moderately expanded posteriorly, extend- ing laterally beyond acrostichal seta but not reaching dorsocentral seta; narrow, faint or- ange stripe extending from anterior margin, crossing dorsocentral seta; distinct yellow- ish sublateral stripe from transverse suture to posterior margin, crossing intra-alar seta; distinct yellowish mark extending posteri- orly from postpronotal lobe to presutural supra-alar seta, almost joining sublateral stripe on transverse suture; scuto-scutellar suture often entirely dark brown, but with- out distinct medial spot. Subscutellum and mediotergite narrowly orange medially, broadly redbrown to dark brown laterally. Pleura with typical dark areas of anepister- num, katepisternum, anepimeron, meron, and laterotergite redbrown to dark brown, strongly contrasting with lighter areas. Ka- tepisternal seta undifferentiated or weak, yellowish, no longer than postocellar seta. Wing (Fig. 2G): Length 8.10-8.60 mm. Vein R,,; almost straight. Vein M strongly curved apically; section between bm-cu and r-m 2.45-2.90 times as long as section between r-m and dm-cu. Pattern light yellow brown to dark orange brown, its margins often dif- fuse. Cells br and r,,, entirely infuscated. Cell dm with large basal hyaline area. Cell bm hyaline. Proximal arm of V-band not extending anteriorly beyond M, not con- nected to S-band. Abdomen: Tergites 1-5 with even-sided medial yellowish stripe, lat- eral third redbrown. Male terminalia: Outer surstylus (Fig. 11C) broad basally, gradually tapering to slightly acute apex; elongate, sec- tion apical to apex of inner surstylus 1.45- 1.80 times as long as basal section; similar to grandis in lateral view. Proctiger with distinct lateral fold; sclerotized part narrow- ly divided into 3 sections. Aedeagus 4.90 mm long; 1.26 times as long as mesonotum. Distiphallus (Fig. 8E) 0.60—0.62 mm long, relatively stout; endophallic sclerite strong, stout and distinctly convoluted apically. Fe- male terminalia: Syntergosternite 7 3.30- 3.95 mm long; 0.90-1.03 times as long as mesonotum. Eversible membrane (Fig. 10F) with dorsobasal scales in triangular pattern; 6-7 large hooklike scales in broadest row; largest scales 0.12 mm long. Aculeus 3.00 mm long; shaft 0.13-0.14 mm wide at mid- point, sides parallel; tip (Fig. 13G) 0.23 mm long, 0.09-0.10 mm wide, apical three-fifths serrate, sides slightly concave, dorsal sur- face with large basal lobe on each side. Sper- mathecae (Fig. 14F) ovoid. Egg (Fig. 14G): Length 1.90-2.02 mm. Broadest diameter 0.22-0.26 mm. Elongate, distinctly curved; slender, especially apical (non-micropyle) half, broadest at basal third. Remarks.—As noted by Stone (1942), Greene (1934) misidentified the holotype as grandis. Korytkowski & Ojeda (1968) de- scribed five males and one female from Ja- yanca, Lambayeque, Peru that they deter- mined as shannoni. According to their description and illustrations, the wing pat- tern and the scutal color pattern of these specimens match those of the holotype ex- cept that the yellowish medial scutal stripe is broader posteriorly, extending laterally beyond the dorsocentral seta. The shapes of the aculeus tip, the scales of the eversible membrane, and the surstylus are similar to A. grandis, however, differing markedly from those of the holotype and other specimens of shannoni described in this paper, and the lengths they reported for syntergosternite 7 (4.58 mm) and the aculeus (4.83 mm) are intermediate between grandis and the Ven- ezuelan females of shannoni. It is possible that Korytkowski and Ojeda confused the terminalia of these specimens with those of specimens of grandis, or these specimens VOLUME 93, NUMBER 1 may represent a different, undescribed spe- cies. I was unable to borrow them for more careful study. The description of the egg is based on several dissected from the abdomen of one of the Venezuelan females. They are ex- tremely similar to the eggs of grandis (Steck and Wharton 1988), and those of both spe- cies are notable for their large size and slen- der shape. Distribution (Fig. 9).—Western Amazo- nia and southern Venezuela; also western Peru if the specimens reported by Koryt- kowski and Ojeda (1968) were correctly de- termined. Specimens examined.—Holotype (see Type data). BRAZIL: Amazonas: Rio Pu- rus, ““Hyutanahan” [= Huitanaa], II.1922, S. M. Klages, 1 6 (CMP). VENEZUELA: Bolivar: Caura Val., C. W. Johnson Collec- tion, 1 2 (MCZ) 1 ¢ (USNM). ACKNOWLEDGMENTS The following individuals kindly loaned or donated the specimens examined in this study: D. A. Grimaldi and D. D. Judd (AMNH); B. R. Pitkin and N. P. Wyatt (BMNH); H. J. Teskey (CNC); C. W. Young (CMP); J. K. Liebherr (CUI); J. Kethley and P. P. Parillo (FMNH); H. V. Weems, Jr. (FSCA); H. J. Miiller and D. Gaedike (DEI); A. Trochez and A. E. Bustillo (ICA); J. A. Raphael (INPA); G. Putruele (INTA); C. L. Hogue (LACM); S. R. Shaw (MCZ); R. Con- treras-Lichtenberg (NMW); E. Kierych (PAN); W. R. Enkerlin and J. Guillen (PMMT); R. Krause (SMT); L. Papp (TMB); C. J. Rosales and L. Joly (UCV); L. F. Jiron (UCRSJ); G. C. McGavin (UMO); W. J. Hanson (USU); R. S. Zack (WSU); R. Dan- ielsson (ZIL). I am grateful to A. Malavasi (Univ. Sao Paulo), V. Hernandez-Ortiz (Inst. de Ecolo- gia, Jalapa), J. D. Harper (N. Carolina State Univ.), J. Fowler (USDA, APHIS), R. A. Zucchi (Univ. Sao Paulo, Piracicaba), A. Freidberg (Tel Aviv Univ.), and N. E. Woodley, G. J. Steck and J. M. Kingsolver 123 (SEL) for their valuable reviews of this pa- per. I also thank Dr. Zucchi for the gift to the USNM of the specimens of bezzii from Brazil. Linda Lawrence (SEL) drew the fig- ures of the mesonota and Brit Griswold pre- pared the wing illustrations. I thank R. Rob- bins for permission to use the distribution maps, which were prepared for the Smith- sonian Institution. LITERATURE CITED Aluja, M., M. Cabrera, E. Rios, J. Guillén, H. Cele- donio, J. Hendrichs, and P. Liedo. 1987. A sur- vey of the economically important fruit flies (Dip- tera: Tephritidae) present in Chiapas and a few other fruit growing regions in Mexico. Fla. Ento- mol. 70: 320-329. Bezzi, M. 1909. Le specie dei generi Ceratitis, Anas- trephae Dacus. Bol. R. Scuola Super. d’Agr., Lab. Zool. Gen. e Agr. Portici 3: 273-313. 1919a. Una nuova specie brasiliana del ge- nere Anastrepha (Dipt.) Bol. R. Scuola Super. d’Agr., Lab. Zool. Gen. e Agr. Portici 13: 3-14. 1919b. Descoberta de uma nova mosca das fructas no Brazil. Chacaras e Quintaes (1919): 373- 374. Blanchard, E. E. 1937. Dipteros argentinos nuevos o poco conocidos. Rev. Soc. Entomol. Argentina 9: 35-58. 1961. Especies argentinas de género Anas- trepha Schiner (sens. lat.) (Diptera, Trypetidae). Revista de Investigaciones Agricolas (Buenos Aires) 15: 281-342. Bondar, G. 1950. Moscas de frutas na Bahia. Bol. do Campo 34: 13-15. Boscan de Martinez, N., J. R. Dedordy, and J. R. Re- quena. 1980. Estado actual de la distribucion geografica y hospederas de Anastrepha spp. (Dip- tera—Trypetidae) en Venezuela. Agron. Trop. 30: 55-63. Caraballo de Valdivieso, J. 1981. Las moscas de fru- tas del genero Anastrepha Schiner, 1868 (Diptera: Tephritidae) de Venezuela. M.S. thesis, Univer- sidad Central de Venezuela, Maracay. 210 pp. Fernandez Yepez, F. 1953. Contribucion al estudio de las moscas de las frutas del genero Anastrepha Schiner (Diptera: Trypetidae) de Venezuela. II Congreso de Ciencias Naturales y Afines (Caracas) Cuaderno no. 7: 1-42. Fischer, C. R. 1932. Nota taxonomica e biologica sobre Anastrepha grandis Macq. (Dipt., Trypeti- dae). Rev. Entomol. (Sao Paulo) 2: 302-310. 1934. Variacdo das cerdas frontaes e outras motas sobre duas especies de Anastrepha (Dipt. Trypetidae). Rev. Entomol. (Sao Paulo) 4: 18-22. 124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Foote, R. H. 1967. Family Tephritidae (Trypetidae, Trupaneidae). Jn P. E. Vanzolini and N. Papavero (eds.), A Catalogue of the Diptera of the Americas south of the United States. Dept. Zool., Sec. Agric., Sao Paulo, Fasc. 57: 1-91. Greene, C. T. 1934. A revision of the genus Anas- trepha based on a study of the wings and on the length of the ovipositor sheath (Diptera: Trypet- idae). Proc. Entomol. Soc. Wash. 36: 127-179. Guagliumi, P. 1966. Insetti e Aracnidi delle piante comuni del Venezuela segnalati nel periodo 1938— 1963. Rel. Monogr. Agric. Subtrop. Trop. (Inst. Agron. L’Oltremare, Firenze) (n.s.) 86: 1-391. Harper, J. D. 1987. Final report to ASOFRUT on the Anastrepha grandis project on honey dew in western Ecuador. Unpublished. 8 pp. Hendel, F. 1914a. Analytische Ubersicht tiber die Anastrepha-Arten (Dipt.). Wien Entomol. Z. 33: 66-70. [January] . 1914b. Die Bohrfliegen Siidamerikas. K. Zool. Anthrop.-Ethnogr. Mus. Abhandl. Ber. (1912) 14: 1-84. [June] Hering, E. M. 1935. Neue siidamerikanische Bohr- fliegen aus dem Stettiner Museum. (9. Beitrag zur Kenntnis der 7rypetidae.) (Dipt.). Stettiner En- tomologische Zeitung 96: 225-229. Jiron, L. F., J. Soto-Manitiu, and A. L. Norrbom. 1988. A preliminary list of the fruit flies of the genus Anastrepha (Diptera: Tephritidae) in Costa Rica. Fla. Entomol. 71: 130-137. Korytkowski, C. and D. Ojeda Pena. 1968. Especies del genero Anastrepha Schiner 1868 en el nor- oeste peruano. Rev. Peru. Entomol. 11: 32-70. 1969. Distribucion ecologia de especies del genero Anastrepha Schiner en el nor-oeste Peru- ano. Rev. Peru. Entomol. 12: 71-95. Lima, A. da Costa. 1930. Sobre as moscas de frutas que vivem no Brasil. Chacaras e Quintaes 34: 21- 24. . 1934. Moscas de frutas do genero Anastrepha Schiner, 1868 (Diptera: Trypetidae). Mem. Inst. Oswaldo Cruz 28: 487-575. 1937. Novas moscas de frutas do genero Anastrepha (Diptera: Trypetidae). O Campo 8: 34— 38, 60-64. Loew, H. 1873. Monographs of the Diptera of North America, pt. III. Smithsn. Misc. Collect. 11 (no. 256): 1-351. Macquart, J. 1843. Diptéres exotiques nouveaux ou peu connus. Mem. Soc. Roy. des Sci., de l’Agr. et des Arts, Lille, 1842: 162-460. 1846. Diptéres exotiques nouveaux ou peu connus. I** Supplément. Mem. Soc. Roy. des Sci., de l’Agr. et des arts, Lille, 1844: 133-364. Malavasi, A., J.S. Morgante, and R. A. Zucchi. 1980. Biologia de “moscas-das-frutas” (Diptera, Te- phritidae). I. Lista de hospedeiros e ocorrencia. Rev. Bras. Biol. 40: 9-16. McAlpine, J. F. 1981. Morphology and terminolo- gy—adults, pp. 9-63. In J. F. McAlpine, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vocke- roth, and D. M. Wood (coords.), Manual of Ne- arctic Diptera, vol. 1. Agriculture Canada, Mono- graph No. 27. Ottawa. Nascimento, A. S. do, A. Malavasi, J. S. Morgante. 1988. Programa de monitoramento de Anastre- pha grandis (Macquart, 1845) (Dip.; Tephritidae) e aspectos da sua biologia, pp. 54-63. Jn H. M. L. de Souza (coord.), Moscas-das-Frutas no Brasil, ANAIS. Fundacao Cargill, Campinas, SP, Brazil. Norrbom, A. L. and K. C. Kim. 1988a. Revision of the schausi group of Anastrepha Schiner (Diptera: Tephritidae), with a discussion of the terminology of the female terminalia in the Tephritoidea. Ann. Entomol. Soc. Am. 81: 164-173. 1988b. A list of the reported host plants of the species of Anastrepha (Diptera: Tephritidae). U.S. Dept. Agric., Animal and Plant Health In- spection Service, Plant Protection and Quaran- tine, Hyattsville, MD. 114 pp. Steck, G. J. and R. A. Wharton. 1988. Description of immature stages of Anastrepha interrupta, A. limae, and A. grandis (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 81: 994-1003. Steyskal, G. C. 1977. Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae). Ento- mological Society of America, Washington, D.C. 35 pp. Stone, A. 1942. The fruitflies of the genus Anastre- pha. U.S. Dept. Agric. Misc. Publ. No. 439, 112 pp. White, I. M. 1988. Tephritid flies, Diptera: Tephriti- dae. Handbooks for the Identification of British Insects 10 (Sa): 1-134. Zucchi, R. A. 1978. Taxonomia das especies de Anas- trepha Schiner, 1868 (Diptera: Tephritidae) assi- naladas no Brasil. Ph.D. dissertation, Universi- dade de Sao Paulo, Piracicaba, Brazil. 105 pp. 1984. Redescricéo da fémea de Anastrepha bezzii Lima, 1934 (Diptera: Tephritidae). Revta. bras. Ent. 28: 561-562. 1988. Moscas-das-frutas (Dip.; Tephritidae) no Brasil: Taxonomia, distribuicao geografica e hospedeiros, pp. 1-10. Jn H. M. L. de Souza (coord.), Moscas-das-Frutas no Brasil, ANAIS. Fundagao Cargill, Campinas, SP, Brazil. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 125-135 MORPHOLOGICAL DIFFERENCES IN GENITALIA OF BRUCHOPHAGUS (HYMENOPTERA: CHALCIDAE) THAT INFEST ALFALFA, RED CLOVER, AND BIRDSFOOT TREFOIL SEEDS (HYMENOPTERA: EURYTOMIDAE) B. McDANIEL AND A. BOE Plant Science Department, South Dakota State University, Brookings, South Dakota 57007. Abstract. —Morphology of female and male genitalia of Bruchophagus gibbus (Bohe- man), B. roddi Gussaskovskii, and B. platypterus (Walker) was studied in specimens col- lected in Arizona, California, Maryland, South Dakota, and Wisconsin from red clover (Trifolium pratense L.), alfalfa (Medicago sativa L.), and birdsfoot trefoil (Lotus cornic- ulatus L.), respectively. Key Words: The name Bruchophagus funebris How- ard was applied by Urbahns (1920) and So- renson (1930) to eurytomids reared from alfalfa (Medicago sativa L.) and red clover (Trifolium pratense L.) seeds in the United States during the early 1900s. After 1930 and until 1952, as pointed out by Strong (1962), the name Bruchophagus gibbus (Boheman) was indiscriminately applied to eurytomids reared from seeds of legumes in North America. However, as early as 1933 Russian entomologists reported a new spe- cies, Bruchophagus roddi Gussakovskil, from alfalfa seeds (Strong 1962). Fedoseeva (1956) described Bruchophagus kolobovae Fedoseeva from seeds of birdsfoot trefoil (Lotus corniculatus L.) in Russia, and Neun- zig and Gyrisco (1958) concluded that eur- ytomids reared from birdsfoot trefoil seeds in North America were B. kolobovae. Strong (1962) studied male and female genitalia of eurytomids reared from alfalfa, red clover, and birdsfoot trefoil seeds and concluded that three distinct species, B. roddi, B. gib- bus, and B. kolobovae, respectively, infested these legumes in North America. He used Bruchophagus, alfalfa, red clover, birdsfoot trefoil, genitalia the relative angle formed by the second rami and the second valvifer to separate the three species. Graham (1970) examined lecto- types of eurytomids from red clover and birdsfoot trefoil and concurred with Fedo- seeva’s (1956) and Strong’s (1962) conclu- sion that species from red clover and birds- foot trefoil were not conspecific. Graham (1970) compared lectotype females of Sys- tole platyptera Walker with reared material of B. kolobovae and considered them to be conspecific. Strong (1962) reported the male genitalia of the three species were almost morphologically identical, and Graham (1970) found less evident differences be- tween males than females of B. gibbus, B. roddi, and B. platypterus. The objectives of this research were to describe the morphology of the female and male genitalia of B. gibbus, B. roddi, and B. platypterus collected in the United States and to identify quantitative genitalia char- acteristics, in addition to those described by Strong (1962), that have taxonomic signif- icance. MATERIALS AND METHODS Genitalia from 30 females and 20 males of B. gibbus, B. roddi, and B. platypterus were mounted on slides in Hoyer’s solution and observed at 100-1200 magnifications. Data obtained for females were ovipositor sheath length, number of ovipositor sheath setae, number of right and left ramus spines, and number of eighth tergite setae. The length of the reproductive apparatus was determined for males. Origins of the specimens studied were as follows. For B. gibbus, 25 females and 10 males were from three different counties in South Dakota, and five females and 10 males were from Woodsboro, Maryland (Freder- ick Co.). For B. roddi, 23 females and 11 males were from 10 different counties in South Dakota, 5 females and one male were from Fresno, California (Fresno Co.), 2 fe- males were from Chandler, Arizona (Mari- copa Co.) and eight males were from Mad- ison, Wisconsin (Dane Co.). For B. platypterus, 17 females and 11 males were from Brookings, South Dakota (Brookings Co.), and 13 females and 9 males were from Madison, Wisconsin (Dane Co.). Quantitative data were subjected to anal- ysis of variance and means were compared by the least significant difference at the 0.01 level. RESULTS AND DISCUSSION DESCRIPTION OF GENITALIA MORPHOLOGY Bruchophagus gibbus Female (Fig. 1): Second valvifers (Vf2) with two setae near apodemes of laminated bridge (Lam.Br.); rami spines (Ra.Sp.) range from 30 to 36 on right valvifers (mean of 33.0 for 30 specimens) and from 24 to 36 on left valvifers (mean of 32.3 for 30 spec- imens), spines widely spaced in laminated bridge region, close together near fulcral plate region; fulcral plate (Ful.Pl.) attach- ment of ovipositor outer plate (Ops.Ot.PI.) near fulcral plate notch (Ful.Pl.Not.); fulcral plate with deep notch, head region with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘ ———.Ful.PI.Sp. —<— — ‘a abt — Ful PLNot. 2, ge _ _ —-0ps.in.PI. Fig. 1. Female genitalia of Bruchophagus gibbus (Boheman). pointed apex; outer ovipositor plate fused with eighth tergite (8th.Teg.), sclerotized, plate in area near attachment of fulcral plate, ventral area membranous; eighth tergite with single row of setae, number of setae ranging from 31 to 49 (mean of 38.6 for 30 speci- mens); setal region plated similar to dorsal fused area of outer ovipositor plate; eighth tergite separated from ventral region of out- er Ovipositor plate by two dark striae that extend from fused dorsal area of outer ovi- positor plate to cercus (Ce.); eighth tergite with a series of long setae beyond cercus, similar in structure to other eighth tergite setae located on epipygium (Ep.); cercus with five setae of different sizes and shapes; ovi- positor inner plate (Ops.In.Pl.) separated from second valvifers by darkened region that connects apodemes of outer and inner rami and apodemes of laminated bridge; this darkened area separates plated region on inner Ovipositor plate from membranous VOLUME 93, NUMBER 1 eS OSHS SSaS6 Aeg. S. Por. % — —.Par. Pl. Set. : 3 : = Se eRaniPl Sats rt ———-Fus. Par. Pl. 1) / } / eal ! | ; y ; | of_ = — = VS. Pl | Ceo | Ai | | | Ste || NS \ | / (TA \| / — — 4 — — Aeg. Apd. yi || ae 4 eg. Ap Hae \\ | / \ —/——— -Dor. Opn. Shp. { / / r . / \ / \ \ \ ; ae, 127 a —amiessien Vite OL. Ae. Nl a aren = Ogi meth: —.—Dgi. Apd. ee ee US.) ale els aes ae VOSnL Figs. 2,3. Male reproductive apparatus of Bruchophagus gibbus (Boheman) dorsal and ventral respectively. centers of second valvifers; second valvifer rami fused with fulcral plate extending past darkened region forming a groove in which fulcral plate apex fits; above this groove are four monitoring spines (Ful.P1.Sp.); this re- gion fused to rami apodemes forming shaft for ovipositor (Ops.); inner ovipositor plate with plated region extending to region of fused ovipositor sheath (Ops.Sh.); ovipos- itor sheaths not articulated, connected to each other by ovipositor ligament (Ops.Sh.Lg.); ovipositor sheath length (Fig. 4) 285 to 420 microns for 30 specimens; Ovipositor sheaths lightly striated; ovipos- itor sheath setae (Ops.Sh.Set.) grouped to- gether at apex with from 8 to 16 setae ex- tending from apex setae toward ovipositor sheath ligament area; ovipositor (Ops.) with typical Bruchophagus teeth on the shaft with a single median tooth followed by paired lateral teeth. Male (Figs. 2, 3): Reproductive apparatus with four dorsal aedeagal sensory pores (Aeg.S.Por.), these paired or staggered, two pairs located within aedeagal striae, aedea- gal dorsal striae (Aeg. Dos.Str.) reaching sec- ond aedeagal sensory pores; ventral apex of aedeagus with three sensory pores on each side, these smaller than dorsal sensory pores; aedeagus (Aeg.) elongated; aedeagal appa- ratus length measured (Fig. 4d) from digital apodemes (Dgi.Apd.) 360 to 440 microns (mean of 382 microns for 20 specimens); aedeagus capable of extending beyond digi- ti; attached to dorsal surface is aedeagal epi- pygium ligament (Aeg.Ep.Lg.) (Fig. 4a, b), that extends to epipygium (eighth tergite); parameres with three setae; apex setae nor- mally hidden between digiti and aedeagus; paramere plate setae (Par.P1.Set.) located on outer margin of curved paramere plate; par- amere plate setae next to apex seta longer and larger than other paramere plate setae; paramere plate setae located near fused par- amere and volsellar plate (Vos.PI.) similar in size to apex paramere setae; digiti (Dgi.) 128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON a Fig. 4. 2 Dgi. Apd. Pa Th I I a 0 Se a ee a— Male reproductive apparatus with aedeagus epipygium ligament attached to aedeagus associated with membranous sheath bearing two acorn-like setae. b—Male eighth tergite with attachment of aedeagus epipygium ligament and membranous sheath. c—Ovipositor sheath showing how length is measured. d— Male reproductive apparatus showing how length is measured. Abbreviations: (Aeg.Ep.Lg.) Aedeagus epipygium ligament; (Aeg.) Aedeagus; (Ce.) Cercus; (8th.Teg.) Eighth tergite; (Ops.Sh.) Ovipositor sheath; (Ops.Sh.Set.) Ovipositor sheath setae; (Dgi.Apd.) Digiti apodemes. with two finger-like teeth and a rounded projection on outer margin of molar, each molar with a single pore on base; digital apodemes (Dgi.Apd.) fused to aedeagal apo- demes; (Aeg.Apd.); aedeagal apodemes pro- trude from caulis (Ca.) dorsally; caulis, par- amere plate and volsellar plate (Vos.Pl.) fused, pigmentation darker on paramere plate section that contains paramere plate setae; center region of caulis transparent al- lowing observation of aedeagal apodemes; caulis forms opening on dorsal surface where aedeagal apodemes protrude. Bruchophagus roddi Female (Fig. 5): Second valvifers (Vf2) with two setae near apodemes of laminated bridge (Lam.Br.); rami spines (Ra.Sp.) range from 24 to 41 on right valvifer (mean of 28.6 for 30 specimens) and from 25 to 39 on left valvifer (mean of 28.5 for 30 spec- imens); spines widely spaced in laminated bridge region, close together near fulcral plate (Ful.Pl.) region; fulcral plate attach- ment of ovipositor outer plate (Ops.Ot.PI.) located near notch of fulcral plate; fulcral VOLUME 93, NUMBER | late Ss Gussakovskii. Female genitalia of Bruchophagus roddi plate notch (Ful.Pl.Not.) deep; fulcral plate apex pointed, fulcral plate margin bordering rami of lower second valvifer with pro- nounced rounded hump; apex of fulcral plate fits into notched ovipositor inner plate (Ops.In.Pl.); ovipositor outer plate fused with eighth tergite (8th.Teg.) lightly scler- otized with plating concentrated on and around eighth tergite region that contains eighth tergite setae (8th. Teg.Set.); dorsal area of ovipositor outer plate membranous; eighth tergite with double row of setae, number of setae range from 44 to 64 (mean of 54.0 for 30 specimens); setal region plat- ed similar to ventral area of outer ovipositor plate, plating absent on area near attach- ment of outer ovipositor plate and fulcral plate; eighth tergite separated from dorsal region of outer ovipositor plate by two dark striae that extend from fused ventral area of ovipositor outer plate to cercus (Ce.); eighth tergite with a series of long setae be- yond cercus, similar in structure to other eighth tergite setae, these associated with epipygium (Ep.); cercus with five setae of different lengths and sizes; ovipositor inner 129 plate (Ops.In.Pl.) separated from second valvifers by darkened region that connects apodemes of upper and lower rami of sec- ond valvifers and apodemes of laminated bridge; this darkened area separates plated region of ovipositor inner plate from mem- branous centers of second valvifers; lower second valvifer ramus which articulates with fulcral plate extends beyond darkened re- gion and forms groove in which fulcral plate apex fits; above groove are four monitoring spines (Ful.Pl.Sp.); region fused forming shaft for ovipositor (Ops.); ovipositor inner plate with plated region extending to fused Ovipositor sheath (Ops.Sh.); ovipositor sheaths not articulated, connected by ovi- positor sheath ligament (Ops.Sh.Lg.); ovi- positor sheath length (Fig. 4c) 250 to 330 microns (mean 276 microns for 30 speci- mens); Ovipositor sheath lightly striated; Ovipositor sheath setae (Ops.Sh.Set.) grouped together at apex with from 6 to 16 setae extending from grouped apex setae to- ward ovipositor sheath ligament area; ovi- positor (Ops.) with typical Bruchophagus teeth on shaft, with single median tooth fol- lowed by paired lateral teeth. Male (Figs. 6, 7): Reproductive apparatus with four aedeagal sensory pores (Aeg. S.Por.), these paired or staggered, one pair located within striae region of aedeagus, striae not reaching to second pair of sensory pores; paired sensory pores may be close to each other or spaced far apart (in B. roddi the staggered position is the normal pat- tern); aedeagal dorsal striae (Aeg.Dos.Str.) of different lengths; ventral apex of aedeagus with three sensory pores on each side, these smaller in size than dorsal sensory pores; aedeagus (Aeg.) elongated length measured (Fig. 4d) from digital apodemes (Dgi.Apd.) 240 to 315 microns (mean of 283 microns for 20 specimens); aedeagus capable of ex- tending beyond aedeagal digiti (Dgi.); at- tached to dorsal surface is the aedeagal epi- pygium ligament (Aeg.Ep.Lg.); parameres with three setae; apex setae hidden between digiti and aedeagus; paramere plate setae 130 Aeg. dos. str. »~ en eae AO Go Or aan Pl. Set. Pails % — ——-Par. PL Set. | er Par. Pl. ee od oe oe ee Al. Ard. Eigss6y 7. YQ — Ful. Pl. Sp. Nee nas Ful. Pl. Not. — Ss cD “Ta ! ‘ ! ‘ 1 fe) J no] a > eu 5 Px by ry y oe ey eo ee ERS By; s. Shae! = STE +5 soo ie Fag a il -a-Teg. Set. 1 4.———Ops. Sh. Lg. —-——-Ops. Sh. Fig. 8. Female genitalia of Bruchophagus platyp- terus Walker. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Aeg. Vt. Por. oe Aeg. SSolertis TEKIN SS SS SS SS Dgi. Por c 4 Cena ea Wee \ice eta Meet Dgi. Apd. ——=—-Fus. Par. Pl. -+——-Vos. Pi. '—-Dgi. Apd. Male reproductive apparatus of Bruchophagus roddi Gussakovskii, dorsal and ventral respectively. (Par.Pl.Set.) located on outer margin of curved paramere plate (Par.Pl.); paramere setae next to apex seta longer and larger than other paramere setae; paramere setae lo- cated near fused paramere and volsellar plate (Vos.Pl.) similar in size to apex paramere setae; digiti (Dgi.) with two finger-like teeth and a rounded projection on outer margin of molar, molar region with single pore; dig- ital apodemes (Dgi.Apd.) fused to volsellar plate; aedeagal apodemes (Aeg.Apd.) pro- trude from caulis (Ca.) dorsally; caulis, par- amere plate and volsellar plate fused; par- amere plate region that contains paramere setae pigmented; caulis center region trans- parent; caulis forms dorsal opening where aedeagal apodemes protrude. Bruchophagus playpterus Female (Fig. 8): Second valvifers (Vf2) with two setae near apodemes of laminated bridge (Lam.Br.); rami spines (Ra.Sp.) range VOLUME 93, NUMBER 1 from 18 to 29 on right valvifer (mean of 24.6 for 30 specimens), and from 19 to 28 on left valvifer (mean of 24.3 for 30 spec- imens); spines widely spaced in laminated bridge region, close together near fulcral plate (Ful.Pl.) region; fulcral plate attach- ment of ovipositor outer plate (Ops.Ot.PI.) located near apex of fulcral plate; fulcral plate notch (Ful.Pl.Not.) deep; fulcral plate apex pointed; fulcral plate margin bordering rami of lower second valvifer without pro- nounced rounded hump; apex of fulcral plate fits into notched ovipositor inner plate (Ops.In.Pl.); ovipositor outer plate fused with eighth tergite (8th. Teg.) sclerotized with plating extending to attachment of outer Ovipositor plate and second valvifer (plating may be absent or lightly marked); dorsal area of ovipositor outer plate membranous; eighth tergite with double row of setae; number of setae range from 37 to 67 (mean of 51.2 for 30 specimens); setal region of eighth tergite plated; eighth tergite with a series of long setae beyond cercus, similar in structure to other eighth tergite setae, these associated with epipygium (Ep.); cercus with five setae of different lengths and sizes; ovi- positor inner plate (Ops.In.Pl.) separated from second valvifers by darkened region that separates plated region of ovipositor inner plate from membranous center of sec- ond valvifers; lower second valvifer ramus which articulates with fulcral plate extends beyond darkened region forming a groove in which fulcral plate apex fits; above groove are four monitoring spines (Ful.P1.Sp.); this region fused forming shaft for ovipositor (Ops.); ovipositor inner plates plated region extends to fused ovipositor sheath (Ops.Sh.); Ovipositor sheath not articulated, connected by ovipositor sheath ligament (Ops.Sh.Lg.); Ovipositor sheath length (Fig. 4c) 185 to 220 microns (mean of 208 microns for 30 spec- imens); Ovipositor sheath lightly striated; Ovipositor sheath setae (Ops.Sh.Set.) grouped together at apex with from 4 to 12 setae extending from grouped apex setae to- ward ovipositor sheath ligament area; ovi- 131 positor (Ops.) with typical Bruchophagus teeth on shaft, with single median tooth fol- lowed by paired lateral teeth. Male (Figs. 9, 10): Reproductive appa- ratus with four aedeagal sensory pores (Aeg.S.Por.), these paired or staggered, paired pores may be close to each other or spaced far apart; normal spacing is far apart; aedeagal striae absent; aedeagal ventral apex with three pairs of sensory pores on each side, these smaller in size than aedeagal dor- sal sensory pores; aedeagus (Aeg.) elongat- ed, length measured (Fig. 4d) from digital apodemes (Dgi.Apd.) 260 to 300 microns (mean of 279 microns for 20 specimens); aedeagus capable of extending beyond ae- deagal digiti (Dgi.); aedeagal epipygium lig- ament present; parameres with three setae; apex setae hidden between digiti and ae- deagus; paramere plate setae (Par.Pl.Set.) located on outer margin of curved paramere plate; paramere setae next to apex setae longer and larger than other paramere setae; paramere setae located near fused paramere and volsellar plate (Vos.PI.); similar in size to apex paramere setae; digiti (Dgi.) with two finger-like teeth and a rounded projec- tion on outer margin of molar, molar with- out pores; digital apodemes (Dgi.Apd.) fused to volsellar plate; aedeagal apodemes (Aeg.Apd.) protrude from caulis (Ca.) dor- sally; caulis, paramere plate and volsellar plate fused; paramere plate region contain- ing paramere setae pigmented; caulis forms dorsal opening where aedeagal apodemes protrude. KEY TO FEMALES AND MALES OF BRUCHOPHAGUS GIBBUS, B. RODDI AND B. PLAYPTERUS USING REPRODUCTIVE STRUCTURES FEMALES 1. Ovipositor sheath length greater than 310 mi- crons (Fig. 4c); eighth tergite with a single row Ofisetaei(kic=sl)) be eres eae eis cane B. gibbus Ovipositor sheath length less than 310 mi- crons; eighth tergite with paired setae form- ingvaidoubleirow: (Figs) 5578) <45---.-. 2+ 2 132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ae ahs. Par. Pit a enin OS PIE my Or —Aeg. Apd. ——-Par. Pl. Set. EN LE IL SS eS Aeg. S. Por —\|.—-Vos. Pl. las Be ——-Dor. Opn. Shp. i Aeg. Apd. Figs.9,10. Male reproductive apparatus of Bruchophagus platypterus Walker, dorsal and ventral respectively. Abbreviations used for female genitalia of all three species; (Lam.Br.) Laminated bridge; (Ra.) Ramus; (Ra.Sp.) Ramus spines; (Vf2) 2nd Valvifer; (Ful.Pl.Sp.) Fulcral plate spines; (Ful.Pl.Not.) Fulcral plate notch; (Ful.PI.) Fulcral plate; (Ops.In.Pl.) Ovipositor inner plate; (Ops.Ot.PIl.) Ovipositor outer plate; (Ops.) Ovipositor; (8th.Teg.) 8th tergite; (8th.Teg.Set.) 8th tergite setae; (Ops.Sh.Lg.) Ovipositor sheath ligament; (Ops.Sh.) Ovipositor sheath; (Ops.Sh.Set.) Ovipositor sheath setae; (Ce.) Cercus; (Ep.) Epipygium. Abbreviations used for male reproductive apparatus of all three species, dorsal and ventral: (Aeg.) Aedeagus; (Aeg.Dos.Str.) Aedeagal dorsal striae; (Aeg.S.Por.) Aedeagal sensory pore; (Par.Pl.Set.) Paramere plate setae; (Fus.Par.Pl.) Fused paramere plate; (Vos.Pl.) Volsellar plate; (Ca.) Caulis; (Aeg.Apd.) Aedeagal apodemes; (Dor.Opn.Shp.) Dorsal opening shape; (Vt.Por.) Ventral pores; (Dgi.) Digiti; (Dgi.Teth) Digiti teeth; (Dgi.Apd.) Digiti apodemes. 2. Ovipositor sheath length between 260 and 290 MICKOnseinlength ey.e, se. ae ee B. roddi Ovipositor sheath length less than 225 mi- Cronshinilengthween ie. oe ee B. platypterus MALES 1. Aedeagus with striae associated with one or more pairs of dorsal sensory pores; dorsal sen- sory pore pairs close together (Figs. 2,6) .... 2 Aedeagus without striae associated with dorsal sensory pores (Figs. 9, 10); length of reproductive apparatus less than 325 mi- crons (see Fig. 4d); dorsal sensory pore pairs separated from each other by more than half the length of aedeagus ........ B. platypterus . Aedeagus striae encompassing both pairs of dorsal sensory pores (Fig. 2); length of repro- nN ductive apparatus greater than 350 microns (see Fig. 4d); dorsal opening shape as shown in Fig. DN bP adh kor ae ROP eae B. gibbus Aedeagus striae not encompassing both pairs of dorsal sensory pores (Fig. 6); length of reproductive apparatus less than 350 mi- crons (see Fig. 4d); dorsal opening shape as Shownbine bigs O mn ee eee B. roddi The aedeagus epipygium ligament (Aeg.Ep.Lg.) (Fig. 4b) is here for the first time given a name. This structure was re- ferred to by McDaniel and Boe (1990) as a covering attached to the eighth tergite and reference was made to the small setae that covered the structure. The aedeagus epi- VOLUME 93, NUMBER 1 Table 1. roddi, and B. platypterus. Species Ovipositor Sheath Length Rami Spines 133 Means and standard deviations of quantitative female genitalia characteristics for B. gibbus, B. Ovipositor Sheath Setae Eighth Tergite Setae microns No. No. No. B. gibbus 349 + 37a! 65.3 + 3.9a 12.6 + 2la 38.4 + 5.la B. roddi 276 + 16b 57.1 + 6.5b 8.9 + 2.7b 54.0 + 6.0b B. platypterus 208 + 12c 48.9 + 4.9c 6:8) == Ie7e¢ pase Halll, ' Means in same column followed by different letter are significantly different at the 0.01 level. pygium ligament is attached to a clear membrane that surrounds the reproductive apparatus. This membranous covering con- tains two acorn-like setae. Associated with a similar membrane attached to the epi- pygium of females are two acorn-like setae identical to those found on males. The ae- deagus epipygium ligament is beset with spine-like setae and covered with a fatty globular material. This material masks the structure of most of the setae on the liga- ment. However, the setae located on the section of the aedeagus epipygium ligament attachment of the epipygium can be seen clearly and their structure noted. If this lig- ament remains attached to the aedeagus when extracting the reproductive apparatus it will occlude the dorsal striae and dorsal sensory pores of the aedeagus. In these cases, length of the reproductive apparatus can be used in separating the males of the three species. The dorsal caulis opening shape can be used to separate the males of all three species. However, it was found that this area of the reproductive apparatus can be dam- aged in the process of removing the whole Table 2. Means and standard deviations of male reproductive apparatus lengths for B. gibbus, B. roddi, and B. platypterus. Male Reproductive Species Apparatus Length microns B. gibbus 382 = 18a! B. roddi 283 + 22b B. platypterus 279 + 8b ' Means followed by different letter are significantly different at 0.01 level. male reproductive apparatus from its con- nection to the eighth tergite. In dried pinned specimens this area is very brittle and the shape of this opening is many times de- stroyed during the removal of the repro- ductive apparatus. The dorsal sensory pores have been found to be a valuable species character in separating the three species. These pores are paired, and may be opposite to each other or staggered. The distance of the pairs of sensory pores from each other has been found to be important in separat- ing B. platypterus from B. roddi and B. gib- bus. The female genitalia of all three species tend to be similar for most structures. There is a size difference between the whole re- productive structure of B. gibbus and B. rod- di and B. platypterus. If the extracted ovi- positor of B. gibbus is placed next to the ovipositor of either B. roddi or B. platyp- terus the larger size of B. gibbus is very ev- ident. Another character that is of value in separating B. gibbus from B. roddi and B. platypterus is the single row of setae on the eighth tergite. B. roddi and B. platypterus have a double row of setae on the eighth tergite. An occasional single seta may occur but the setae form a definite double row in contrast to the single row found on B. gib- bus. There may appear to be a difference between the plating on both the outer and inner oOvipositor plates; however, it was found that some individuals may have very light plating causing difficulty in using the presence or absence of plating in different regions for separating species. To the trained 134 eye the shape of the darkened region be- tween the valvifers is diagnostic. ANALYSIS OF QUANTITATIVE CHARACTERS Highly significant (P less than 0.01) dif- ferences were found among the three species for ovipositor sheath length, number of rami spines, number of ovipositor sheath setae, number of eighth tergite setae, and male reproductive apparatus length (Tables 1, 2). Ovipositor sheath length was significantly longer for B. gibbus than B. roddi and sig- nificantly longer for B. roddi than B. pla- typterus. The same relationships between species were observed for numbers of rami spines and ovipositor sheath setae (Table 1). Number of eighth tergite setae was sig- nificantly higher for B. roddi and B. platyp- terus than B. gibbus. Male reproductive apparatus length was significantly greater for B. gibbus than B. roddi or B. platypterus (Table 2). The ap- paratus of B. gibbus was approximately 35% longer than those of the other two species. Although significant differences were found among species for the four female and single male genitalia characteristics, the use- fulness of any single characteristic for tax- onomic purposes is limited by relatively wide ranges of intraspecific variation. For example, the ovipositor sheath length of 23% of the B. roddi individuals fell within the range for B. gibbus, but no overlap was found for B. gibbus and B. platypterus. A similar relationship between species was also ob- served for number of rami spines. Ranges overlapped for B. roddiand B. gibbus as well as B. roddi and B. platypterus, but ranges of B. gibbus and B. platypterus did not. Length of the male reproductive apparatus was use- ful for separating B. gibbus from B. roddi and B. platypterus since the length of the shortest B. gibbus apparatus was approxi- mately 50 microns longer than the longest apparatus for B. roddi and B. platypterus. Overlaps in the ranges of all three species were observed for numbers of ovipositor PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sheath and female eighth tergite setae. How- ever, B. roddi and B. platypterus can be sep- arated from B. gibbus by arrangement of eighth tergite setae. B. roddi and B. platyp- terus have a double row of eighth tergite setae compared to a single row for B. gibbus. We observed the relative angle formed by the second rami and second valvifers used by Strong (1962) to separate the three spe- cies. We concur with his observations but found this angle to be influenced by the amount of pressure applied during slide preparation. Strong (1962) also pointed out that the male genitalia of B. gibbus were significantly larger than those of B. roddi and B. platypterus. However, he examined only 3 specimens of B. platypterus and thus was not able to accurately compare the size of the aedeagi of B. roddi and B. platypterus. These data indicated large interspecific quantitative differences in male and female genitalia characteristics of B. gibbus, B. rod- di, and B. platypterus. However, they also revealed substantial intraspecific variation that makes species separation based strictly on quantitative genitalia characteristics somewhat difficult. We have described mor- phological differences among species for ar- rangement of eighth tergite setae on the fe- males and aedeagal striae on the males that are useful for separating these three species. These traits, when used in combination with the easily measured quantitative traits, pro- vide additional guidelines for researchers who are uncomfortable with Strong’s (1962) characters for separation of the three spe- cies. LITERATURE CITED Fedoseeva, L.I. 1956. New species of the genus Bru- chophagus Ashm. (Hymenoptera: Chalcidoidea) developing upon leguminous plants. Doklady Akad. Nauk. SSSR III: 491-493. Graham, M. W. R. deV. 1970. Taxonomic notes on some Palearctic Eurytomidae (Hymenoptera: Chalcidoidea). Proc. R. Entomol. Soc. Lond. 39: 139-152. McDaniel, B. and A. Boe. 1990. Life history studies, VOLUME 93, NUMBER 1 host records, and morphological descriptions of genitalia of Eurytoma tylodermatis Ashm. (Eury- tomidae) from South Dakota. Proc. Entomol. Soc. Wash. 93: 96-100. Neunzig, H. H. and G. G. Gyrisco. 1958. Host re- lationships of seed chalcids reared from birdsfoot trefoil. Jour. Econ. Entomol. 51: 409-410. Sorenson, C. J. 1930. The alfalfa-seed chalcis-fly in 135 Utah. 1926-1929, inclusive. Utah Agric. Exper. Stat. Bull. 218. Strong, F.E. 1962. Studies on the systematic position of the Bruchophagus gibbus complex (Hymenop- tera: Eurytomidae). Ann. Entomol. Soc. Amer. 55: 1-4. Urbahns, T. D. 1920. The clover and alfalfa seed chalcid-fly. USDA Bull. 481. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 136-158 NORTH AMERICAN CHARIPIDAE: KEY TO GENERA, NOMENCLATURE, SPECIES CHECKLISTS, AND A NEW SPECIES OF DILYTA FORSTER (HYMENOPTERA: CYNIPOIDEA) ARNOLD S. MENKE AND H. H. EVENHUIS (ASM) Systematic Entomology Laboratory, Agricultural Research Service, USDA, %U.S. National Museum of Natural History, Washington, D.C. 20560; (HHE) Marijkelaan 2, 6721 GN Bennekom, Netherlands. Abstract.—The nomenclature of the 35 North American species of Charipidae is re- viewed. The following new synonymies are proposed: A/lotria ambrosiae Ashmead, 1898, Glyptoxysta necans Kieffer, 1909, Bothrioxysta numidica Kieffer, 1909, and Charips fla- vipes Ionescu, 1963 = Phaenoglyphis villosa (Hartig), 1841; Alloxysta vandenboschi An- drews, 1978 = Alloxysta obscurata (Hartig), 1840; Allotria tritici Fitch, 1861 and Charips areolata Kieffer, 1909 = Alloxysta victrix (Westwood), 1833. Synonymic checklists are presented for the North American species, and those in the primarily Old World genera Apocharips and Dilyta. Characters previously used to separate the Charipinae and Allox- ystinae are reviewed and illustrated (several are unreliable), and an illustrated key to the five genera in the family is provided. Adelixysta Kierych, 1988 is synonymized with Alloxysta Forster, 1869. A new species, Dilyta rathmanae, is described from the state of Washington; it is the first valid North American member of the genus. D. rathmanae is a hyperparasitoid of pear psylla, and its biology is summarized. Key Words: The Charipidae has five genera and about 200 valid species that are hyperparasitoids of Homoptera, mostly aphids. Dilyta and Apocharips, containing about ten species, have been reared only from psyllids. Ex- amination by Menke of material from the state of Washington, reared from psyllids by Robin Rathman, indicated that it be- longed to Dilyta, but that it was not the widespread European subclavata Forster. In the catalog of Hymenoptera of North America, Burks (1979) listed only one spe- cies of Dilyta, necans Kieffer (1909), de- scribed from Massachusetts, the syntypes of which had never been examined by any au- thor. Andrews (1978) suggested that the type material of species described in Kieffer’s (1909) paper might be at the Entomology Insecta, Hymenoptera, Charipidae, Di/yta, Alloxystinae, Charipinae Department of the University of Arkansas, Fayetteville. The syntypes of necans as well as Charips areolata Kieffer and Lytoxysta brevipalpis Kieffer were found there. Ex- amination of necans showed it is not a Dily- ta, but a synonym of the holarctic Phaeno- glyphis villosa (Hartig). Thus, Rathman’s wasp is the first valid Di/yta in the New World and is undescribed. The nearctic genera that are hyperparasit- oids of aphids (A//oxysta, Phaenoglyphis and Lytoxysta) were revised by Andrews (1978). However, he did not treat several North American species, including A//oxysta tritici (Fitch) (1861), the oldest name in our fauna. Andrews established considerable new syn- onymy, but did not appreciate the holarctic nature of some species. His paper includes VOLUME 93, NUMBER 1 a world catalog of the aphid hyperparasi- toids but it is incomplete; he excluded Dil- yta, believing it belonged to a “new subfam- ily.” Burks (1979) cataloged the nearctic Cha- ripidae (as Alloxystidae), but he did not in- clude the new taxa or synonymy of Andrews (1978), or the synonymy of Evenhuis (1973) and Evenhuis and Barbotin (1977). Subse- quently Evenhuis (1982), Evenhuis and Kiriak (1985), Evenhuis and Barbotin (1987) and Fergusson (1986) established new syn- onymies for European species, especially in Alloxysta. A few charipids are now known to be holarctic, and some of the synonymy by these authors directly affects the North American fauna. In this paper we describe the first valid North American species of Dilyta, and in- clude a biological summary of it written by Robin Rathman and G. Paulson. Their summary conclusively demonstrates that the new species is a hyperparasitoid. The Eu- ropean D. subclavata is often cited as a pri- mary parasitoid, although Herhard (1986) considers it a hyperparasitoid. Another as- pect of this paper is the presentation of notes on the status and generic placement of some charipid species, and the designation of lec- totypes for several. We also provide syno- nymic checklists for North American gen- era; the checklists for Di/yta and Apocharips include all currently recognized species. Complete citations for the 35 North Amer- ican charipid species are not given here be- cause they occur elsewhere (Andrews 1978, Burks 1979, Fergusson 1986, etc.). Our syn- onymy is largely derived from the previ- ously cited papers, but some is new. A key to the five valid genera of Charip- idae is presented because: 1) Hemicrisis Forster was synonymized by Evenhuis (1973), and 2) Apocharips Fergusson (1986) and Adelixysta Kierych (1988) were recently described. We synonymize Adelixysta with Alloxysta here. Bothriocynips (Diaz 1979) from South America, has been attributed to the Charipidae, but from her illustrations 137 and the fact that the female has 11 and the male 13 flagellomeres, the genus appears misassigned; it may be a eucoilid. Bothrio- cynips was taken from an ant nest, a habitat atypical of charipids. TAXONOMIC HISTORY AND CHARACTERS The family name has vacillated between Alloxystidae and Charipidae. The history of this, and the confusion that has existed over type species of certain genera, has been out- lined by Quinlan and Evenhuis (1980) and Fergusson (1986). The oldest family-group name, “‘Allotriina” Thomson (1862: 397, 406), based on the junior homonym 4/llotria Westwood, is unavailable under the pro- visions of Article 39 of the 1985 edition of the Code of the International Commission on Zoological Nomenclature. Incidently, Quinlan and Evenhuis (1980) and Fergus- son (1986) attribute the name Allotriinae to Forster (1869), but Thomson’s usage of the name is older. The next available family- group name is Charipinae Dalla Torre and Kieffer, 1910, based on Charips Haliday. Although Charips is a junior synonym of Dilyta Forster, the family-group names Charipidae and Charipinae remain avail- able under the provisions of Article 40 of the Code. The name Alloxystidae, which has been commonly used for the family un- til recently, is much younger, being pro- posed in 1931 by Hellén. The genera were segregated into two sub- families by Kierych (1979b): Alloxystinae and Dilytinae (= Charipinae). Biology sup- ports this grouping; the alloxystines are hy- perparasitoids of aphids and the charipines are hyperparasitoids of the Psylloidea (no other Cynipoidea are associated with Ho- moptera, and we regard the biology of cha- ripids as an apomorphy for the family). Kie- rych (1979b) described and illustrated the two subfamilies (see also Kierych 1979a). The important differences between Allox- ystinae and Charipinae noted by Kierych are summarized on Table 1. Some of Kierych’s characters are unreli- ey 138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table |. Character Antennal flagellomeres Mandibular teeth three at apex Frontoclypeal sulcus present Pronotum nae Scutellum Marginal cell closed or open Gastral terga I-II Spiracles of gastral tergum VI widely separated able for defining the two subfamilies. An- tennal differences are questionable when the New World genus Lytoxysta is examined; the flagellomeres are broadly joined throughout (Figs. 1, 7-9), but this condition may be a homoplasy. Kierych (1979b) in- dicated that the mandible was tridentate in Alloxystinae and bidentate in Charipinae, but this differs bilaterally in the few Allox- ysta and Lytoxysta that we examined. Typ- ically the left mandible has one less tooth. The right mandible has two large teeth, and one or two much smaller ones. The left mandible has two large teeth and sometimes a third small one. In Dilyta both mandibles have two large teeth, but the inner one on the left mandible is very broad. Similar mandible asymmetry was noted in /balia (Ibaliidae) by Ronquist and Nordlander (1989). Thus, the number of mandibular teeth, even if restricted to one mandible, is not useful in delineating subfamilies. The frontoclypeal sulcus varies in Al/oxysta from present to evanescent; the sulcus is absent in Lytoxysta. The forewing marginal cell (see especially A//oxysta) and pronotal charac- Figs. 1-9. Alloxystinae all separated by constrictions with or without two anterior cari- rounded apically, without carinae approximately equal Kierych’s (1979a, b) diagnostic characters for charipid subfamilies. Charipinae last two flagellomeres broadly joined two at apex absent with pair of carinae with carinae apically that form a projection open fused or I very small narrowly separated at midline ters vary in Alloxystinae and are therefore unreliable subfamily features. The remaining characters (presence or ab- sence of scutellar carinae apically, form of terga I and II, and location of the spiracles on tergum VI), and host differences, are the only reliable delimiters of the two subfam- ilies. The scutellar difference between Charip- inae and Alloxystinae has not been used since Kierych introduced it, but the pro- truding carinae found in Dilyta (inverted U-shaped, Figs. 13, 14) and Apocharips (M-shaped, Figs. 11, 12) support recogni- tion of the Charipinae. Presumably the presence of these carinae is an apomorphy. Fergusson (1986) indicated that the scutel- lum in Apocharips is also margined laterally by a carina, but none is visible in SEM pho- tographs of our material (Fig. 12). The re- duction of tergum I or complete fusion with II, and the narrowly separated spiracles on tergum VI in Charipinae are also synapo- morphies. Although the Charipinae is de- fined by synapomorphies, none are known a Antennal details. 1, female of Lytoxysta brevipalpis. 2, female of Alloxysta sp. 3, male of Dilyta rathmanae. 4, last four flagellomeres of female Lytoxysta brevipalpis. 5, last four flagellomeres of female A//oxysta sp. 6, last four flagellomeres of female Di/yta rathmanae. 7-9, female Lytoxysta brevipalpis; 7, scape, pedicel and flagellomere I; 8, flagellomeres I-III; 9, flagellomeres IV—VI (note that hexagonal reticulation diminishes on VI). 139 VOLUME 93, NUMBER 1 140 for the Alloxystinae, and the latter is prob- ably paraphyletic. Lytoxysta is the most distinctive genus in the family, differing from all others in: lack- ing a triangular depression on the meso- pleuron (Fig. 16); having a fine, more or less hexagonal, reticulate sculpture on the head, antenna, thorax and legs (Figs. 7-9, 15, 16, 21); having a tight cluster of special setae on the back of the head that is opposed by a similar group of setae on the anterolateral area of the pronotum (Fig. 15); having all flagellomeres connate; and lacking long fringing setae around the apical half of the forewing. All of these character states with the possible exception of the form of the antenna, are probably apomorphies. The marginal cell of Lytoxysta is open on three sides (only 2r and part of RS present, Fig. 24), but an incomplete cell is not unique to the genus. Although Lytoxysta is substantially dif- ferent from all other genera in the family, it can be assigned to the Alloxystinae on the basis of its simple scutellum, the approxi- mately equal size of gastral terga I-II, and broadly separated spiracles on tergum VI, all presumed pleisomorphies. But this placement only emphasizes the paraphylet- ic nature of Alloxystinae. Moreover, the thoracic makeup of Lytoxysta, including general shape, is obviously quite different from the other alloxystine genera (compare Fig. 16 with 17, 18). Although the autapo- morphies of Lytoxysta suggest that the ge- nus could be placed in its own subfamily, recognition of three subfamilies for the five charipid genera is taxonomically irrational. More likely no subfamilies should be rec- ognized in Charipidae. KEY TO GENERA OF CHARIPIDAE 1. Mesopleuron without triangular depression dorsally (Fig. 16); forewing margin without long fringing setae apically; head and thorax with fine reticulate sculpture (Figs. 7, 15); North AMIMETICA hee ec: Lytoxysta Kieffer — Mesopleuron with triangular depression dor- sally (Figs. 17-20); forewing margin with long PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fringing setae apically; head and thorax usually smooth, polished (Figs. 17—20) 2. Lower part of mesopleuron with horizontal sul- cus (Fig. 17); scutum sometimes with notauli; scutellum usually with pair of large foveae ba- sally (Fig. 22); holarctic ... Phaenoglyphis Forster — Mesopleuron without horizontal sulcus (Figs. 18-20); scutum without notauli; scutellum without basal foveae (Fig. 23) 3. All flagellomeres separated by constrictions, antenna moniliform (Figs. 2, 5); visible gaster represented by at least two large terga whose dorsomedian lengths are subequal (Fig. 27): apex of scutellum rounded, without carinae (Fig. lO) R ns a ks ee eee eee Alloxysta Forster — Last two flagellomeres broadly joined, preced- ing flagellomeres with constrictions between them (Figs. 3, 6, 34, 35, 38, 39); visible gaster represented single tergal plate (Figs. 29-32), or if two, then basal one much shorter than second along middorsal line (Fig. 28); apex of scutel- lum with M or inverted U-shaped carina (Figs. 12 U4). oo Aes eee eee eee 4 4. Gaster with small basal tergum (Fig. 28); inner (R,) and outer veins of forewing marginal cell essentially reaching wing margin (Fig. 25); Old World: . .....A8e6 eee Apocharips Fergusson — Gaster without small basal tergum (Figs. 29- 32); inner vein (R,) of forewing marginal cell often not reaching wing margin (Fig. 26); Hol- arctic and Ethiopian Regions ..... Dilyta Forster SPECIES NOTES Subfamily Alloxystinae Genus Phaenoglyphis Forster Phaenoglyphis ambrosiae (Ashmead) Allotria ambrosiae Ashmead, 1898: 156. Lectotype female, Cambridge, Mass., designated by Andrews (1978); in Na- tional Museum of Natural History, Washington, D.C. Andrews (1978) placed ambrosiae in Phaenoglyphis after studying the types. He did not realize, however, that it was a junior synonym of the European species villosa (Hartig) (NEW SYNONyMy), now known to be holarctic. The two Ashmead syntypes are mounted on a single point, and Andrews (1978) claims to have placed a lectotype label on the pin, with the indication that the specimen clos- est to the tip of the point is the type. How- VOLUME 93, NUMBER 1 141 . Li OA AUING /, > %s 7} Ny ‘ 4 ibe 4, ? = Ny ‘4 7%! weet I) | Figs. 10-14. Details of scutellum, metanotum and adjacent structures. 10, Al/oxysta megourae, left three- quarter rear view, arrow points to apical edge of scutellum. 11, 12, Apocharips trapezoidea, 11, left profile, arrows point to apical carinae of scutellum; 12, left three-quarter rear view, arrows point to M-shaped carina. 13, 14, Dilyta rathmanae; 13, left profile, arrow points to apical carina; 14, left three-quarter rear view, arrow points to inverted U-shaped carina. 142 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ey yey — : al Figs. 15-20. Left side of thorax and rear of head. 15, Dense setal patches on rear of head and front of pronotum, Lytoxysta brevipalpis. 16, Lytoxysta brevipalpis. 17, Phaenoglyphis sp. (upper arrow points to tri- angular depression, lower arrow to sulcus). 18, A//oxysta sp. 19, Apocharips trapezoidea (arrow points to pro- truding apical carinae of scutellum). 20, Dilyta rathmanae (arrow points to protruding apical carina of scutellum). VOLUME 93, NUMBER | Figs. 21-23. scutellar foveae. 23, Alloxysta sp. ever there is no such label. We have placed a lectotype label on the pin with Andrews’ name. Andrews (1978) noted that males are very rare in North American populations of this species (as ambrosiae). Of 158 specimens studied only 3 were males, and only females were obtained by him in rearing experi- ments and from Malaise traps. Andrews suggested that the species was parthenoge- netic, and his dissection of the female re- vealed no spermatheca, supporting his con- tention. In Europe villosa reproduces sexually, and the first male flagellomere is bent. According to Andrews (1978: fig. 46) flagellomere I in North American males is straight, suggesting that his specimens may have been misassociated. Rearing experi- ments are needed to ascertain the true sex ratios of villosa in North America. Phaenoglyphis americana Baker Phaenoglyphis americana Baker, 1896: 131. Lectotype male, Fort Collins, Colorado, present designation; in National Museum of Natural History, Washington, D.C. Andrews (1978: 35) misidentified the 143 Dorsum of head and thorax. 21, Lytoxysta brevipalpis. 22, Phaenoglyphis sp., arrow points to above specimen as a female. He also treated it as the holotype in spite of indicating that a second syntype is in the Cornell Univer- sity collection. We have placed a lectotype label on the USNM specimen. Phaenoglyphis flavipes (Ionescu), New COMBINATION Charips flavipes lonescu, 1963: 174. Holo- type male, Nehoiu, Romania; in Institute of Biology, Academy of the Rumanian People’s Republic, Bucharest. In a letter to Evenhuis in 1971, Ionescu indicated that the male type of flavipes has a transverse sulcus on the mesopleuron, and a pair of foveae basally on the scutellum. These features assign flavipes to the genus Phaenoglyphis. The illustrations of the an- tenna in Ionescu (1963, 1969) clearly in- dicate that flavipes is a synonym of villosa (Hartig) (New Synonymy). Although Io- nescu describes the marginal cell as closed, it is open basally in vi//osa but often difficult to ascertain. The scutum lacks notauli and the second abdominal tergum is much long- er than the first along the midline. 144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 24 LYTOXYSTA BREVIPALPIS 25 APOCHARIPS TRAPEZOIDEA 26 DILYTA RATHMANAE Figs. 24-26. Right forewing (setae not shown). VOLUME 93, NUMBER 1 Phaenoglyphis necans (Kieffer), NEw COMBINATION Glyptoxysta necans Kieffer, 1909: 481. “Forest Hills,’ Mass., col. by “Hay- hurst,” ex “Aphis rumicis L.?” on “Che- nopodium album.” Lectotype female with no locality data, just the number 385 and a label with the species name, present des- ignation; in University of Arkansas, Fay- etteville. Burks (1979) placed necans in Dilyta sim- ply because G/yptoxysta was a generic syn- onym. The syntypes indicate however, that necans is a member of the genus Phaeno- glyphis and also a junior synonym of the common European (and holarctic) villosa (Hartig) (NEw SYNONYMy). The University of Arkansas possesses two female syntypes. Neither has any labels that identify them as type material of necans, but they are glued to the same distinctive paper points used for Kieffer’s Lytoxysta brevipennis and Charips areolata. Both specimens have an identification label in- scribed with the name G/yptoxysta necans. These labels were probably placed on the material by someone other than Kieffer. Each specimen also has a label with the number 385. Similar numbers appear on the type material of other species described in the same paper, and we are confident that these two specimens are Kieffer’s syntypes of necans. One with most of the right an- tenna missing has been labeled as lectotype. Phaenoglyphis numidica (Kieffer), NEw CoMBINATION Bothrioxysta numidica Kieffer, 1909: 482. Holotype female, ““Algerien, Blidah, im Juli (Quedenfeld)”’; in Zoologisches Mu- seum der Humboldt-Universitat, Berlin. Kieffer described numidica from a single female, and in 1984 Evenhuis examined the type and found it to be a synonym of Phaenoglyphis villosa (Hartig) (NEW Synonymy). The type has two labels that read “Charips numidica K.” and “‘Blidah- 145 Médéah, Algerien, Juli-August 84, Queden- feldt.”” The last label is on blue paper. Ev- enhuis placed two additional labels on the type: “holotype 1984, H. H. Evenhuis” (or- ange paper) and “Phaenoglyphis villosa (Hartig) female, det. H. H. Evenhuis 1984” (white paper). Genus Alloxysta Forster Alloxysta alaskensis Ashmead Alloxysta alaskensis Ashmead, 1902: 142. Lectotype female, Muir Inlet, St. Paul Is- land, Alaska, designated by Andrews (1978) unknowingly; in National Muse- um of Natural History, Washington, D.C. Andrews (1978: 53) regarded the above lectotype as the holotype, but it was actually one of five syntypes (3 males, 2 females). According to Article 74(b) of the Code, An- drews’ recognition of a holotype constituted a lectotype designation. We have placed a lectotype label on the specimen. The mght antenna past the scape 1s missing, as 1s the abdomen and the right legs beyond the cox- ae. The other syntypes present in the USNM, two males, are in worse condition. Alloxysta areolata (Kieffer) Charips areolata Kieffer, 1909: 481. “‘Forest Hills,’ Massachusetts, col. ““Hayhurst.”’ Ex an aphid “‘Macrosiphum sp.?” Lec- totype female, labelled Forest Hills, Mass., present designation; in National Museum of Natural History, Washington, D.C. This species was not treated by Andrews (1978) although the name appears in his catalog as a species of Al/loxysta. Burks (1979) also listed the name in A//oxysta. Kieffer had eight syntypes of areolata; one is in the University of Arkansas collection and four are in the National Museum of Natural History collection. The latter were apparently obtained by Lewis Weld in 1924 via an exchange with the University of Ar- kansas. All of the specimens are identical with the widespread species Alloxysta vic- trix (NEW SYyNOoNYMy). We have selected 146 one of the USNM specimens as the lecto- type. The left antenna is gone. The lectotype has a machine-printed label “Forest Hills, Mass.” and the date May 29, 1909. This specimen and the others have a second label written in ink with the number 384, and a third greenish gray label. Only the lectotype has locality data. The University of Arkan- sas specimen has a handwritten label, ap- parently by Kieffer, that says ‘““Charips ar- eolata K Alle mit Griinem Zettel.” Alloxysta circumscripta (Hartig) Xystus circumscripta Hartig, 1841: 352. Lectotype female, no locality, designated by Evenhuis, 1982; in Zoologisches Staatssammlung, Munich. Alloxysta circumscripta (Hartig), a Eu- ropean wasp, was synonymized under vic- trix by Fergusson (1986) without any dis- cussion, but the species 1s quite distinct (NEW Stratus). Unlike victrix, circumscripta lacks pronotal carinae and the pronotum is cov- ered with vestiture except for a longitudinal glabrous strip at the middle. The pronotum of victrix 1s more broadly glabrous at the middle and has carinae. Alloxysta fuscicornis (Hartig), New Status Xystus fuscicornis Hartig, 1841: 350, 352. Lectotype female, Braunschweig, Ger- many, designated by Evenhuis, 1982; in Zoologische Staatssammlung, Munich. Allotria ancylocera Cameron, 1886: 85. Ho- lotype male, Carruber Glen, Scotland; in British Museum (Natural History). Allotria brassicae Ashmead, 1887: 14. Lec- totype female, Jacksonville, Florida, des- ignated by Evenhuis, 1974; in National Museum of Natural History, Washing- ton; Die: Allotria infuscata Kieffer, 1902a: 16. Lec- totype male, Amiens, France, designated by Evenhuis and Barbotin, 1987; in Mu- sée de Picardie, Amiens. Alloxysta fuscicornis and its synonyms PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON were synonymized under the common hol- arctic species victrix (Westwood) by Fer- gusson (1986). He stated that color, es- pecially of the head, is unreliable for separating species of A//oxysta, especially victrix and its relatives. He also said that the length of the flagellomeres varies with body size, and ““must be used with care.” Although victrix and fuscicornis are similar, Evenhuis has found that they are quite dis- tinct when reared from the mummies of Macrosiphum rosae (Linnaeus), and related species, as well as Brevicoryne brassicae (Linnaeus). A//oxysta victrix is larger, and the head is yellowish or reddish with the vertex darker. The head is quite dark in fuscicornis and the antennae and legs are darker than in victrix. The antennal flagel- lomeres are comparatively longer in victrix. On this evidence, Evenhuis hereby resur- rects fuscicornis from synonymy with victrix (NEw Stratus). It 1s possible that Fergusson simply confused a number of similar spe- cies, but clearly the status of fuscicornis and victrix needs further careful study. Alloxysta hayhursti (Kieffer) Charips hayhursti Kieffer, 1909: 481. ‘‘For- est Hills,’ Mass., ex “‘Aphis atriplicis,” col. ““Hayhurst.” Andrews (1978: 74) listed this name in the synonymy of A//loxysta xanthopsis (Ash- mead), but did not indicate new synonymy or give evidence for treating the species as a synonym. Apparently no Kieffer speci- mens exist at the University of Arkansas and none are to be found in the USNM. The identity of hayhursti thus remains in doubt. Alloxysta ignorata (Kieffer) Allotria testacea Hartig of Cameron, 1889: 56. Misidentification. Allotria testacea Hartig of Cameron, 1890: 250. Misidentification. Allotria castanea Hartig of Cameron, 1890: 256. Lapsus for testacea. Dilyta ignorata Kieffer, 1900: 114. Replace- VOLUME 93, NUMBER 1 ment name for Allotria testacea of Cam- eron, nec Hartig, 1841. Cameron (1889, 1890) misidentified a species that had an open marginal cell as testacea Hartig (Quinlan 1978a, transferred testacea Hartig to Synergus in Cynipidae). Hartig’s (1841) description leaves no doubt that testacea has a closed marginal cell. Aware of this, Kieffer (1900) proposed a replacement name for Cameron’s “‘testa- cea.” Quinlan (1978b) tried to locate type material of ignorata under the name festa- cea in the BMNH, but was unsuccessful. Cameron (1890) used the name festacea in his key to species, but in the same work the name castanea appears where testacea ought to be listed in the enumeration of species. Thus the type material of ignorata may be labeled “‘castanea,”’ not “‘testacea.” Kieffer (1902b) and Dalla Torre and Kief- fer (1910) placed ignorata in the genus A/- loxysta, and it is likely that the species is correctly assigned to that genus, rather than Dilyta. Alloxysta rufipleura Baker Alloxysta rufipleura Baker, 1896: 135. Lec- totype male, Fort Collins, Colorado, pres- ent designation; in National Museum of Natural History, Washington, D.C. Andrews (1978: 51-52) correctly synon- ymized rufipleura under Alloxysta affinis (Baker). He listed only one specimen which he regarded as the holotype, Baker had two males however, and we have selected and labelled one as lectotype. The lectotype has Baker’s red type label also. Alloxysta tritici (Fitch) Allotria tritici Fitch, 1861: 841. No locality mentioned. “‘Swept from wheat, aphis in- fested,” July 8, 1861. Lectotype female, labeled “15,341” and “‘Allotria tritici,” present designation; in National Museum of Natural History, Washington, D.C. This species is the earliest one described from North America. Weld (1951) listed tri- 147 tici from various States in the eastern United States and assigned the species to A//oxysta, but Andrews (1978) did not treat the name in his revision. Burks (1979) indicated that the proper name of the aphid from which Fitch’s material was reared is Schizaphis graminum (Rondani), and that the host is Aphidius nigripes (Ashmead). Barnes (1988) indicates that insects from the Fitch Collection are scattered but that some material was obtained by the National Museum of Natural History. In his catalog of Fitch insect material Barnes lists tritici, but does not mention the existence of type material, the inference being that no spec- imens are known to exist. There is a single specimen in the National Museum of Nat- ural History, however, that seems to qualify as Fitch material of tritici. It has a genuine Fitch catalog number label, #15,341, as de- scribed and illustrated by Barnes (1988). A second label is machine printed and reads ‘*Fitch’s Coll.” The specimen also has a very old handwritten ink label that reads ‘“Al- lotria tritici”’ followed by what appears to be an initial that is undecipherable. This label also says “‘a. nigriceph.”” The hand- writting may not be Fitch’s because it is unlike that depicted in Barnes (1988: 86). The National Museum has the type of A/- lotria avenae Fitch, a wasp now assigned to the Eucoilidae. Allotria avenae was de- scribed right after tritici and the type has a Fitch catalog number and “‘Fitch’s Coll.” label like those on the specimen of tritici. Thus it appears that the specimen of fritici can be considered as type material of Fitch’s species. The specimen of tritici is pinned sideways through the thorax with a very fine pin. That pin is stuck through a piece of cork affixed to a regular insect pin. The abdomen is missing, but the antennae are intact and in- dicate that the specimen is a female. All legs and wings are present. In Andrews’ (1978) key to species of Al/oxysta, this specimen runs to victrix (Westwood) and it compares favorably with Old and New World mate- 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rial of that species. The specimen agrees with Fitch’s original description. The lower part of the head is yellow, and the upper part is brownish. The scape, pedicel, first two fla- gellomeres and legs are yellow. The pro- portions of the first five flagellomeres agree fairly closely with the description, and are the same as those in victrix. We are treating this specimen as a Fitch syntype of tritici, designating it as the lectotype, and synon- ymizing it with victrix (NEW SYNONYMY). Alloxysta vagans Kieffer Alloxysta vagans Kieffer, 1909: 480. ‘‘For- est Hills, Massachusetts,” ex ““Hyalopte- rus dactylidis Hayhurst.” Andrews (1978) listed this species in his catalog but did not treat it in the main part of his revision. Apparently no type material exists at the University of Arkansas or in the USNM. Thus the identity of this species remains uncertain. Alloxysta vandenboschi Andrews Alloxysta vandenboschi Andrews, 1978: 69. Holotype male, 2 mi. s. Kelsey Bay, Brit- ish Columbia, Canada; in National Mu- seum of Natural History, Washington, D.C: Examination of Andrews’ type by Even- huis has established that vandenboschi is a synonym of the holarctic species obscurata (Hartig) (NEW SYNONYMy). Genus Lytoxysta Kieffer Lytoxysta brevipalpis Kieffer Lytoxysta brevipalpis Kieffer, 1909: 480. “Forest Hills, Massachusetts,” ex Aphis rumicis.” Lectotype female designated by Andrews, 1978: 23; in National Museum of Natural History, Washington, D.C. Kieffer described this species from 9 spec- imens. One is in the University of Arkansas collection, and three are in the National Museum of Natural History. The latter were obtained by Lewis Weld in 1924 in an ex- change with the Univ. of Arkansas. An- drews (1978) selected one of the USNM specimens as lectotype and supposedly placed a “‘red”’ lectotype label on it in 1970; however, we found no such label on the specimen. It is clear which specimen An- drews designated because it is the only spec- imen with a locality label that reads “Forest Hills, Mass.” with the date Sept. 22, 1908. A second handwritten label gives the num- ber 388. Below it is a plain white square. We have placed a lectotype label on the pin with Andrews’ name on it. The other spec- imens lack locality labels, but all have the 388 label and the plain white square. They do not have Andrews’ paralectotype labels either, although he says they were applied. The specimen belonging to the University of Arkansas museum has an additional handwritten label, apparently Kieffer’s, that reads ““Lytoxysta brevipalpis (Alle mit ro- tem Zettel).’’ Obviously the red square la- bels have faded. None of the specimens available to us show any evidence of being the material upon which Kieffer (1909: 480) described the “Var. nigra n. var.” The status of this name remains in doubt, although Andrews (1978) synonymized it under brevipalpis. Generic assignment unknown Glyptoxysta bifoveata Girault Glyptoxysta bifoveata Girault, 1931: 2. Syn- types: Victoria, Australia; in Queensland Museum, Brisbane. The description of this wasp is too brief to permit positive generic assignment. The notation that it was “‘on peach aphis” sug- gests that it belongs in Al/oxysta. Subfamily Charipinae Genus Apocharips Fergusson Apocharips eleaphila (Silvestri), NEw COMBINATION Alloxista eleaphila Silvestri, 1915: 274 (footnote). Syntypes, male and female, It- aly; in Museo Zoologico della Universita di Napoli? VOLUME 93, NUMBER | This species was reared from the psyllid Euphyllura olivina (Costa) which immedi- ately suggests that it belongs in the subfam- ily Charipinae. The species is briefly de- scribed in a footnote after the extensive description of another new species, Al/ox- ista peraptera Silvestri, which is well illus- trated. The latter is clearly an Apocharips because it has two abdominal terga. Silvestri did not describe the abdomen of eleaphila, but compared it with peraptera; we, there- fore, assume that it also has two terga and tentatively assign it to the genus Apocharips (NEW COMBINATION). Apocharips peraptera (Silvestri), NEw COMBINATION Alloxista peraptera Silvestri, 1915: 272. Syntypes, male and female, Nefasit, Eri- trea, Ethiopia; in Museo Zoologico della Unversita di Napoli? This species was reared from the psyllid Euphyllura aethiopica Silvestri, and the de- scription and figures clearly indicate that peraptera belongs in the genus Apocharips (NEw COMBINATION). Apocharips talitzkii (Belizin), NEw CoMBINATION Glyptoxysta talitzkii Belizin, 1966: 7. Ho- lotype male, Kishinev, Moldavian SSSR; in Zoological Museum, Academy of Sci- ences of the USSR, Leningrad. The host of talitzkii, Psylla pyri, suggests that it belongs in the Charipinae. One of the distinctive features of the subfamily is the connate nature of the last two flagellomeres of the antenna, but Belizin does not mention this in his description. Nevertheless, it seems probable that ta/itzkii belongs in the genus Apocharips based on Belizin’s description of the abdomen, and that is where we have assigned it (NEW COMBINATION). Genus Dilyta Forster Dilyta africana (Benoit), New ComMBINATION Alloxysta africana Benoit, 1956: 439. Ho- lotype male (“‘female’’), Zaire: Kivu: Ru- 149 tshuru; in Musée Royal de l’Afrique Cen- trale, Tervuren, Belgium. Benoit had 25 specimens when he de- scribed africana. Evenhuis examined nine, including the holotype, and the species be- longs in the genus Dilyta (NEW COMBINA- TION). The holotype is a male but Benoit mistakenly identified it as a female. He gave the relative lengths and widths of the fla- gellomeres which agree with the holotype, except that one of the flagellomeres between the sixth and the twelfth (all with the same relative lengths of 7.5 and widths of 3) was obviously omitted. Benoit described flagel- lomere I as being slightly curved, indicating that his holotype was a male, not a female. Although Benoit said the gaster had two visible segments and even gives their rela- tive lengths, the type, as well as the para- types, have only one tergum. The species thus clearly belongs in Dil/yta. The species exhibits other characteristics typical of the genus Dilyta: scutellum with U-shaped api- cal carina; marginal cell shape typical for the genus, except both R, and RS reach the wing margin. Generic assignment unknown Dilyta mascagnini Girault Dilyta mascagnini Girault, 1935: 2. Syn- types, Sandgate, Australia; in Queensland Museum, Brisbane. The description of this wasp is too brief and vague to determine if it belongs in Di- lyta. Girault stated that the “club” of the antenna was “*2-jointed”’; this suggests it be- longs in the subfamily Charipinae. But his mention of “abdomen glabrous, 2 exceeding 3 and over half the abdomen” makes that assignment doubtful. Girault compared mascagnini with Alloxysta thoreauini, an- other Australian species which he described as new in the same paper. The abdomen of thoreauini was described as “‘non-segment- ed,’ suggesting that this species really be- longs in Dilyta. Only a study of Girault’s 150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON type material will clarify the status of mas- cagnini. Checklists of Charipidae North American species of subfamily Alloxystinae Hellén, 1931 (= Allotriinae Thomson, 1862) Phaenoglyphis Forster 1869 Generic synonyms Hemicrisis Forster, 1869. Syn.: Evenhuis (1973), Fergusson (1986). Auloxysta Thomson, 1877. Bothrioxysta Kieffer, 1902. Charipsella Bréthes, 1913. americana Baker, 1896; Canada: Northwest Territories, British Colombia, Alberta, Manitoba, Ontario, Quebec, New Bruns- wick; U.S.: Alaska, California, Colorado, Utah, Maryland. brevicornis Kieffer, 1907. Syn.: Andrews (1978). nevadensis Kieffer, 1907. Syn.: Andrews (1978). calverti Andrews, 1978; U.S.: California. falcata Andrews, 1978; Canada: British Co- lumbia. gutierrezi Andrews, 1978; Canada: British Columbia, Saskatchewan; U.S.: Mon- tana. laevis Andrews, 1978; Canada: British Co- lumbia, Alberta. pecki Andrews, 1978; U.S.: Tennessee. pilosa Andrews, 1978; Canada: British Co- lumbia, Alberta; U.S.: Idaho, Colorado, California. ruficornis (Forster), 1869 (Hemicrisis); Hol- arctic: Europe, North America. Canada: British Colombia, Saskatchewan, Ontar- 10, Quebec; U.S.: California, North Car- olina. pubicollis (Thomson), 1877 (Allotria). Syn.: Evenhuis (1973). stenos Andrews, 1978; U.S.: Alaska villosa (Hartig), 1841 (Xystus); Holarctic: Europe, North America. Canada: North- west Territories, British Colombia, Al- berta, Ontario, New Brunswick, Nova Scotia; U.S.: California, Massachusetts. piciceps (Thomson), 1862 (Allotria). Syn.: Evenhuis and Barbotin (1977). collina (Cameron), 1889 (Allotria). Syn.: Evenhuis and Barbotin (1977). ambrosiae (Ashmead), 1898 (Allotria). NEw SYNONYMY carpentieri (Kieffer), 1902 (Allotria). Syn.: Evenhuis and Barbotin (1977). foveigera (Kieffer), 1902 (Allotria). Syn.: Evenhuis and Barbotin (1977). curvata (Kieffer), 1902 (Allotria). Syn.: Evenhuis and Barbotin (1977). recticornis (Kieffer), 1902 (Allotria). Syn.: Evenhuis and Barbotin (1977). subaptera (Kieffer), 1904 (Alloxysta). Syn.: Evenhuis and Barbotin (1977). campyla (Kieffer), 1904 (Alloxysta). Syn.: Evenhuis and Barbotin (1977). necans (Kieffer), 1909 (Glyptoxysta). NEw SYNONYMY numidica (Kieffer), 1909 (Bothrioxysta). NEw SYNONYMY flavipes (Ionescu), 1963 (Charips). NEw SYNONYMY Alloxysta Forster, 1869 Generic synonyms Allotria Westwood, 1833, nec Hiibner, 1823 Xystus Hartig, 1840, nec Schoenherr, 1826 Pezophycta Forster, 1869 Nephycta Forster, 1869 Thoreauana Girault, 1930 Adelixysta Kierych, 1988, NEw SYNONYMY Alloxista is incorrect subsequent spelling Our efforts to obtain type material of Ade- lixysta have been unsuccessful, but we are convinced that Kierych’s (1988) new genus isa synonym of Alloxysta (NEW SYNONYMY). Clypeal characters are the only features that seem to differentiate Adelixysta sawonie- wiczi, type species of the new genus, from most species of Al/oxysta. The sulcus be- tween the tentorial pits (frontoclypeal sul- cus) is faintly impressed, and the free mar- gin of the clypeus is truncate instead of VOLUME 93, NUMBER 1 arcuate. A/loxysta pallidicornis (Curtis) has a truncate clypeus also, but the species has a strongly impressed frontoclypeal sulcus. Both taxa have the same propodeal carinae but they differ in antennal and wing char- acters. Alloxysta pallidicornis attacks species of Pauesia that parasitize lachnine aphids on conifers, and sawoniewiczi will likely be found in similar hosts. A third and as yet undescribed species is known in this com- plex that was reared from Pausia cendrobii on Cedrus in Morocco. affinis (Baker), 1896 (Dylita); Colorado. abdominalis Baker, 1896. Syn.: Andrews (1978). apicalis Baker, 1896. Syn.: Andrews (1978). coloradensis (Baker), 1896 (Dylita). Syn.: Andrews (1978). gracilis Baker, 1896. Syn.: Andrews (1978). magna Baker, 1896. Syn.: Andrews (1978). ruficeps (Baker), 1896 (Dylita), nec Zet- terstedt, 1838. Syn.: Andrews (1978). rufipleura Baker, 1896. Syn.: Andrews (1978). similis (Baker), 1896 (Dylita). Syn.: An- drews (1978). alaskensis Ashmead, 1902; Canada: Que- bec; U.S.: Alaska. bakeri (Kieffer), 1907 (Allotria),; U.S.: Cal- ifornia. Possibly a synonym of megourae, see Andrews (1978: 47). bicolor (Baker), 1896 (Dylita); Canada: Brit- ish Colombia; U.S.: Colorado. robusta Baker, 1896. Syn.: Andrews (1978). commensurata Andrews, 1978; U.S.: Mary- land. coniferensis Andrews, 1978; U.S.: Tennes- see. dicksoni Andrews, 1978; U.S.: Maryland. filimentosa Andrews, 1978; U.S.: Idaho. fuscicornis (Hartig), 1841 (Xystus); Holarc- tic Region (NEw Status) 151 ancylocera (Cameron), 1886 (Allotria). Syn.: Evenhuis (1982). brassicae (Ashmead), 1887 (Allotria). Syn.: Evenhuis (1982). infuscata (Kieffer), 1902 (Allotria). Syn.: Evenhuis (1982). grioti (De Santis), 1937 (Charips). Syn.: Diaz (1980). aphidiinaecida (De Santis), 1937 (Cha- rips). Syn.: Diaz (1980). halli Andrews, 1978; Canada: British Co- lombia, Alberta; U.S.: Washington. lachni (Ashmead), 1885 (Allotria), Canada: British Colombia, Manitoba, Ontario; U.S.: California, North Carolina, Florida. lackni ‘“‘Dalla Torre and Kieffer,” An- drews (1978: 63), apparently a lapsus. leguminosa (Weld), 1920 (Charips); U.S.: Idaho, Oregon, Utah. Possibly asynonym of megourae according to Andrews (1978: 65). longiventris Baker, 1896; U.S.: Colorado. Not treated by Andrews (1978). megourae (Ashmead), 1887 (Allotria); U.S.: Florida. minuscula Andrews, 1978; Canada: Ontar- 10. miniscula Andrews, 1978: 48, lapsus. obscurata (Hartig), 1840 (Xystus); Holarc- tic: Europe, North America. Canada: British Colombia; U.S.: Alaska. homotoma Kieffer, 1904. Syn.: Evenhuis and Barbotin (1987). vandenboschi Andrews, SYNONYMY. pallidicornis (Curtis), 1838 (April 1) (Cyn- ips); Holarctic: Europe, North America. Canada: Alberta, Quebec; U.S.: Alaska, Colorado. Tax.: Quinlan and Fergusson (1981). minuta (Zetterstedt), 1838 (Dec. 31) (Cynips). Syn.: Evenhuis and Kiriak (1985). forticornis (Giraud), 1860 (Allotria). Syn.: Quinlan and Fergusson (1981). basalis (Thomson), 1862 (Allotria). Syn.: Evenhuis and Kiriak (1985). 1978. NEw HS anthracina Andrews, 1978. Syn.: Even- huis and Kiriak (1985). quebeci Andrews, 1978; Canada: Quebec. rauchi Andrews, 1978; Canada: British Co- lombia. schlingeri Andrews, 1978; U.S.: Arizona. vagans Kieffer, 1909; U.S.: Massachusetts. Not treated by Andrews (1978). victrix (Westwood), 1833 (Allotria); Nearc- tic, Palearctic and Neotropical Regions. ruficeps (Zetterstedt), 1838 (Cynips). erythrocephalus (Hartig), 1840 (Xystus). Syn.: Evenhuis (1972, 1982). tritici (Fitch), 1861 (Allotria). NEw SYNONYMY. macrocera (Thomson), 1877 (Allotria). Syn.: Evenhuis (1982). atriceps (Buckton), 1879 (Cynips). Syn.: Fergusson (1986) but types apparently not examined. curvicornis (Cameron), 1883 (Allotria). Syn.: Fergusson (1986). amygdali Buckton, Kieffer, 1902a (Cyn- ips), lapsus for atriceps (Buckton). Syn.: Fergusson (1986). lateralis (Kieffer), 1902b (Allotria). Syn.: Evenhuis and Barbotin (1987). luteiceps (Kieffer), 1902b (Allotria). Syn.: Evenhuis and Barbotin (1987). luteicornis (Kieffer), 1902b (Allotria). Syn.: Evenhuis and Barbotin (1987). grandicornis (Kieffer), 1904 (A/lotria). Syn.: Evenhuis and Barbotin (1987). aerolata (Kieffer), 1909 (Charips). New SYNONYMY. xanthopsis (Ashmead), 1896 (Allotria); Canada: British Colombia; U.S.: Alaska, California, Utah, Massachusetts, Florida. ? hayhursti (Kieffer), 1909 (Charips). Syn.: Andrews (1978) implied this synony- my but it remains unproven. Lytoxysta Kieffer, 1909 brevipalpis Kieffer, 1909; Canada: British Colombia, Alberta, Manitoba, Saskatch- ewan; U.S.: California, Colorado, Idaho, Massachusetts, Ohio, Utah, Washington. nigra Kieffer, 1909. Syn.: Andrews (1978). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON World species of subfamily Charipinae Dalla Torre and Kieffer, 1910 (= Dilytinae Kierych, 1979) The best general morphological descrip- tion of this subfamily is by Kierych (1979a, b), although his nomenclature is incorrect in part. Morphology and taxonomy of the group are also treated by Fergusson (1986). So far as known, all charipines are hyper- parasitoids in Psylloidea. Apocharips Fergusson, 1986 eleaphila (Silvestri), 1915 (Alloxista). Italy. Reared from Euphyllura olivina (Costa). NEw ComMBINATION. peraptera (Silvestri), 1915 (Alloxista). Eri- trea, Ethiopia. Reared from Euphyllura aethiopica Silvestri. NEw COMBINATION. talitzkii (Belizin), 1966 (Glyptoxysta). Mol- davian SSR. Reared from Psylla pyri (L.). NEw ComBINATION. trapezoidea (Hartig), 1841 (Xystus). Europe. Reared from Psylla sp. NEw CoMBINA- TION. xanthocephala (Thomson), 1862 (Allo- tria). Syn.: Evenhuis (1982). Dilyta Forster, 1869 Generic synonyms Dylita Forster, 1869, an incorrect subse- quent spelling. Charips Haliday in Marshall, 1870. Glyptoxysta Thomson, 1877. africana (Benoit), 1956 (Alloxysta); Zaire. NEw CoMBINATION. rathmanae Menke and Evenhuis, new spe- cies. Washington State. Reared from Cacopsylla alba (Crawford) and C. pyri- cola (Forster). subclavata Forster, 1869. Europe. Reared from Psylla pyri(L.) and Psyllopsis fraxini (E.): microcera (Haliday) in Marshall, 1870 (Charips). Syn.: Quinlan and Evenhuis (1980). heterocera (Thomson), 1877 (Glyptoxys- ta). Syn.: Hellén (1963). VOLUME 93, NUMBER 1 thoreauini (Girault), 1935 (Alloxysta); Aus- tralia. NEw COMBINATION. Dilyta rathmanae Menke and Evenhuis, NEW SPECIES (Figs. 3, 6, 13, 14, 20, 26, 31-34, 36, 38, 40) Black except antenna and legs yellow, api- cal half of flagellum and mid and hindlegs sometimes suffused with brown. Thorax with patches of dense, wool-like, white setae as follows (the white color may be due to some exudate because it is missing on a few specimens): on pronotum dorsolaterally and laterally, on scutellum laterally, on subalar fossa of mesopleuron, on metapleuron just beneath hindwing base, on propodeum pos- terolaterally just above hindcoxa, and also posteromedially. Setal girdle at base of ter- gum I similar. Female antenna as in Figs. 6, 34, 36; comparative lengths of pedicel and flagellomeres I—X1:9.5:5.0:2.75:3.2:3.5: 4.5:5.4:7.75:8.5:9.0:9.25:9.0; flagellomere I length less than twice width (10.5:5.5); fla- gellomeres I-VI without linear tyli, follow- ing articles with them. Male antenna as in Figs. 3, 33, 38, 40; comparative lengths of pedicel and flagellomeres I—XII: 6.5:7.25: B25=3-025:0 01 D311 5:7. 1528.2528-55025.9.0: 9.0; flagellomeres I-III or IV without linear tyli, following articles with them. First ter- gum usually without obvious pitting on api- cal half (Figs. 31, 32). Body length 0.9-1.2 mm. Types: Holotype male: WASHINGTON, Chelan Co.: Nahahum Canyon near We- natchee River (13 km n. Wenatchee), 180 m, ex Cacopsylla alba on Salix exigua, May 31, 1986, Robin Rathman, voucher #86-2- 11 (in collection of National Museum of Natural History, Washington, D.C.). Para- types, 23 females, 1 1 males, some with same data as holotype except different dates in 1985-1986, others from following localities in Chelan County, Washington: McManus Orchard, Yaxon Canyon, 6.6 km s. Cash- mere, elev. 242 m, June 18, 1987, June 23, 1988, G. S. Paulson; Pflugrath Orchard, 153 Derby Canyon, 8.3 km se. Leavenworth, July 8 and 15, 1986, G. S. Paulson; Dryden, July 8, 1987, G. S. Paulson. Paratypes in the Entomology Collection, Washington State University, Pullman; the National Museum of Natural History; the H. H. Ev- enhuis Collection; the British Museum (Natural History); and the American En- tomological Institute, Gainesville, Florida. Discussion: Dilyta rathmanae differs from the Old World species subclavata Forster in several ways. The easiest feature to appre- ciate is the apparent absence in most spec- imens of rathmanae of the rather dense field of pits found on tergum I of subclavata (compare Figs. 29, 30 and 31, 32). Actually a few scattered pits are present in most spec- imens of rathmanae but they are very shal- low and barely visible even in SEM pho- tomicrographs. One exception is a single male of rathmanae that has a clearly seen field of rather deep pits. Differences between the two species are also found in the distribution of antennal tyli (compare Figs. 33, 36, 40 and 37, 41), especially in the female. Specifically, tyli are absent on the first six flagellomeres of the female of rathmanae (Fig. 36) but only ab- sent on the first four of subclavata (Fig. 37). Tyli are absent on flagellomeres I-III or IV of the male in rathmanae (Fig. 33), and are absent on IJ-II or III of subclavata (Fig. 41). The overlap between the two species in the distribution of male antennal tyli makes them more difficult to distinguish in this sex. In both species the basalmost male fla- gellomere with tyli seems to have only one or two of these structures, and determining their presence can be very difficult, even at 250 magnification. Subsequent flagello- meres typically have several tyli that are rather conspicuous. Four of the seven males of rathmanae have tyli on flagellomere III which suggests that these structures are ab- sent on that flagellomere about 50% of the time. The proportions of the pedicel and fla- gellomeres I-II also differ between the two — ore 154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SSS SSS RON Figs. 27-32. Left view of gaster. 27, female Alloxysta megourae. 28, female Apocharips trapezoidea. 29-30, female and male, respectively, Dilyta subclavata. 31-32, female and male, respectively, Dilyta rathmanae. on Figs. 33-41. Antennal features in Dilyta. 33, right male flagellomeres I-VII of rathmanae (arrows point to tyli on V-VI). 34, 35, female, rathmanae and subclavata, respectively. 36, 37, apex of pedicel, and first six flagellomeres of female rathmanae and subclavata, respectively. 38, 39, male, rathmanae and subclavata, re- spectively. 40, 41, apex of pedicel, and first four or five flagellomeres of male rathmanae and subclavata, respectively. 155 VOLUME 93, NUMBER 1 156 species. In the female of rathmanae flagel- lomere I is less than twice as long as wide, and its length is about half that of the pedicel (Figs. 34, 36). In the female of subclavata flagellomere I is more than twice as long as wide (13:5), and its length is more than three- fourths as long as the pedicel (13:16) (Figs. 35, 37). In males of rathmanae flagellomere II is about one-third the length of I (Figs. 33, 40). Whereas, in males of subclavata it is about half the length of I (Figs. 39, 41). Dilyta rathmanae is a smaller insect (.9- 1.2 mm) than subclavata (1.2-1.5 mm). Distribution: Known with certainty only from Chelan Co., Washington. However Ferris and Hyatt (1923) reared a species of “Alloxysta” from the psyllid Neophyllura arbuti (Schwarz) in California (we are fol- lowing Hodkinson, 1988, for current nearc- tic psyllid names in this paper). It seems likely that what they really had was a species of Dilyta. Etymology: We take pleasure in naming this little wasp after its discoverer, Robin Rathman. BIOLOGICAL SUMMARY OF DILYTA RATHMANAE R. J. Rathman, Dept. of Entomology, Univ. of Hawaii, 3050 Maile Way, Honolulu, Ha- wall 96822, and G. S. Paulson, Dept. of Entomology, Washington State Univ., Pull- man, Washington 99164 Dilyta rathmanae was first reared from Cacopsylla alba (Crawford) on willow, Salix exigua L., in 1985 in Washington State (Rathman and Brunner 1988). Adults were also collected in unsprayed pear orchards and may be an important component of the pear psylla complex, Cacopsylla pyricola (Forster). Recent European authors disagree whether species of the Charipinae are pri- mary or hyperparasitoids. Fergusson (1986, 1988) reported that Dilyta subclavata For- ster, the single European species of the ge- nus, is a primary parasitoid of psyllids. In PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON southern France, however, this species was reported by Herhard (1986) to be a hyper- parasitoid of Psylla pyri (L.), through an en- cyrtid, Prionomitus mitratus (Dalman). Our field observations and rearing experiments have shown that Dilyta rathmanae is a hy- perparasitoid of psylla through an encyrtid, Trechnites sp. In laboratory rearing experiments con- ducted in July and August, 1987, parasit- oid-free psylla nymphs were placed in cages (about 4 mm x 20 mm diam.) designed to confine the psylla to the leaf surface of pear seedlings. Adult 7rechnites collected from pear trees and adults of Dilyta collected from Salix were added to one group of cages. Adults of Dilyta were only added to the sec- ond group. At the completion of the study adults of Dilyta and Trechnites had emerged from pear psylla mummies in group one, but in the second group that had been ex- posed only to Dilyta, the pear psylla were not parasitized. This indicates that Dilyta rathmanae is not a primary parasitoid of pear psylla. Observations of field collected mummies of C. pyricola and C. alba con- firmed the laboratory results. On 7 and 8 August 1988 parasitized psyl- la were collected from Salix and examined in the laboratory. One parasitized psylla contained a partially consumed adult of Trechnites and a cynipoid larva. Similarly in July 1988 a partially consumed pupa of Trechnites and a cynipoid larva were found within a parasitized C. pyricola. In this in- stance the cynipoid larva pupated and an adult female of Dilyta emerged. In August 1988 a mummy of C. pyricola was collected in which the forms of two distinct parasitoid pupae could be seen through the cuticle of the psylla. Two days later an adult male of Dilyta emerged. The psylla mummy was then dissected and a partially consumed Trechnites was recovered. These studies support the findings of Herhard (1986) and indicate that Dilyta rathmanae is a hyper- parasitoid of C. alba and C. pyricola. Thus members of both the Charipinae and Alloxystinae are hyperparasitoids. The Al- VOLUME 93, NUMBER 1 loxystinae are hyperparasitoids of aphids via Aphidiinae (Braconidae) and Aphelinidae (Chalcidoidea) (Fergusson 1988), while the Charipinae are hyperparasitoids via Encyr- tidae (Chalcidoidea). Both hyperparasitism and the use of Homoptera as hosts are prob- able behavioral synapomorphies of the Al- loxystinae and Charipinae that indicate the Charipidae is a monophyletic group. ACKNOWLEDGMENTS Chris Carlson (Dept. of Entomology, Univ. of Arkansas, Fayetteville) lent the Kieffer material for our study. Eliane de Coninck (Musée Royal de l’Afrique Cen- trale, Tervuren, Belgium) lent type material of Alloxysta africana Benoit to Evenhuis for study. Robin Rathman (formerly of the Tree Fruit Research Center, Wenatchee, Wash- ington) and G. S. Paulson (Dept. of Ento- mology, Washington State University, Pull- man) furnished the material of the new Dilyta. Goran Nordlander (Swedish University of Agricultural Sciences, Uppsala) and Nigel Fergusson (British Museum (Natural His- tory)) reviewed several drafts of our manu- script. Allan Norrbom, Sueo Nakahara, and Eric Grissell (Systematic Entomology Lab- oratory, USDA, Washington, D.C.), John Sorensen, California Dept. of Food and Ag- riculture, Sacramento. California, Byron Alexander, University of Kansas, Law- rence, Kansas, and David Wahl, American Entomological Institute, Gainesville, Flor- ida reviewed the final ms. LITERATURE CITED Andrews, F.G. 1978. Taxonomy and host specificity of nearctic Alloxystinae, with a catalog of the world species (Hymenoptera: Cynipidae). Occ. Pap. Ent., Calif. Dept. Food Agric., no. 25. 128 pp. Ashmead, W. H. 1887. Report on insects injurious to garden crops in Florida. U.S. Dept. Agric., Div. Ent., Bull. 14: 9-29. 1898. Descriptions of new parasitic Hyme- noptera. Proc. Entomol. Soc. Wash. 4: 155-171. . 1902. 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Les Cynipides (Suite), 3° Tri- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bu: Allotriines (Allotriinae), pp. 5-78, 592-602. In André, E., ed., Species des Hyménoptéres d’Eu- rope & d’Algérie, vol. 7, part 2. 748 pp. [pp. 5-78 = 1902, 592-602 = 1904] 1909. Beschreibung neuer in Blattlausen schmarotzender Cynipiden. Naturwis. Zeit. Forst- Landwirtschaft 7: 479-482. Kierych, E. 1979a. Notes on the genera Dilyta For- ster, 1869, and G/yptoxysta Thomson, 1877 (Hy- menoptera, Cynipoidea, Alloxystidae). Part I. Ann. Zool. 34: 453-460. 1979b. Notes on the genera Dilyta Forster, 1869 and Glyptoxysta Thomson, 1877 (Hyme- noptera, Cynipoidea, Alloxystidae). Part II. Di- lytinae subfam. n. Ann. Zool. 35: 59-64. 1988. A new genus and a new species of cynipoids (Hymenoptera, Cynipoidea, Charipi- dae) from Poland. Ann. Zool. 41: 351-354. Quinlan, John. 1978a. Xystus testaceus Hartig, 1841 and Synergus variabilis Mayr, 1873 redefined and redescribed (Hymenoptera: Cynipoidea). Zeitschr. Arb. Osterr. Ent. 30: 71-74. 1978b. On the identity of some British Al- loxystinae described by P. Cameron and by J. J. Kieffer (Hymenoptera, Cynipidae). Ent. Bericht. 38: 124-128. Quinlan, J. and H. H. Evenhuis. 1980. Status of the subfamily names Charipinae and Alloxystinae (Hymenoptera: Cynipidae). Syst. Ent. 5: 427-430. Quinlan, J.and N. D. M. Fergusson. 1981. The status and identity of the Cynipoidea (Hymenoptera) de- scribed by J. Curtis. Ent. Gazette 32: 251-256. Rathman, R. J. and J. F. Brunner. 1988. Observa- tions on the biology of a new species of Dilyta (Hymenoptera: Charipidae) from Washington State. Pan-Pac. Ent. 64: 93-97. Ronquist, F. and G. Nordlander. 1989. Skeletal mor- phology of an archaic cynipoid, /balia rufipes (Hy- menoptera: Ibaliidae). Ent. Scandinavica, suppl. 33. 60 pp. Silvestri, F. 1915. Contributo alla conoscenza degli insetti dell’olivo dell’Eritrea e dell’Africa meri- dionale. Boll. Lab. Zool. Gen. Agra. R. Scuola Super. Agric. Portici 9: 240-334. Thomson, C. G. 1862. Foérs6k till uppstallning och beskrifning af Sveriges Figiter. Ofvers. K. Vet.- Akad. Forh. 18: 395-420. (Although dated 1861 it was published in 1862 according to Horn & Schenkling, /ndex Litteraturae Entomologicae.) Weld, L. 1951. Superfamily Cynipoidea, pp. 594—- 654. In C. F. Muesebeck, K. V. Krombein, H. Townes, et al., eds., Hymenoptera of America North of Mexico. U.S. Dept. Agric., Agric. Mon- ogr. 2. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 159-162 SYNONYMY IN THE TREEHOPPER GENERA HOPLOPHORION, METCALFIELLA, AND OCHROPEPLA (HOMOPTERA: MEMBRACIDAE) STUART H. MCKAMEY'! AND LEwis L. DEITz Department of Entomology, North Carolina State University, Raleigh, North Carolina 27695-7613. Abstract. —The nomenclatorial status of the Neotropical treehopper genera Hoplopho- rion Kirkaldy, 1901, Metcalfiella Goding, 1929, and Ochropepla Stal, 1869, and of the tribe Hoplophorionini Goding, 1926, is reviewed. Hoplophorion is considered a junior synonym of Ochropepla; Metcalfilla is reinstated as a valid genus. The type species and three other species of Hoplophorion are placed in Ochropepla, producing one new com- bination, Ochropepla triangulum (Germar); most of the other species recently placed in Hoplophorion are referred to Metcalfiella. The nominal species (and subspecies) of Och- ropepla (8) and Metcalfiella (25) are listed. Key Words: phora, nomenclature Funkhouser (1951) noted that the tree- hopper genus Metcalfiella “‘has had a check- ered nomenclatorial career of misadven- ture.”’ Indeed, revisionary work at the species level disclosed the need to review the status of the genera Metcalfiella Goding, 1929 (type species: Hoplophora pertusa Germar, 1835, by original designation), Ochropepla Stal, 1869 (type species: Hoplophora corrosa Fairmaire, 1846, by subsequent designa- tion), and Hoplophorion Kirkaldy, 1901, a replacement name for Hoplophora Germar, 1833 (preoccupied by Hoplophora Perty, 1833) (type species: Membracis triangulum Germar, 1821, by subsequent designation), as well as the availability of the family-group name Hoplophorionini, of the subfamily Membracinae (Deitz 1975). ' Current address: Department of Ecology and Evo- lutionary Biology, Box U-43, University of Connect- icut, Storrs, Connecticut 06269-3043. Neotropical, Hexapoda, Insecta, Membracinae, Hoplophorionini, Hoplo- STATUS OF GENUS-GROUP NAMES The generic name Metcalfilla, first pub- lished (Goding 1929a) with no description or indication (Article 12, International Commission on Zoological Nomenclature 1985), became available when Goding (1929b) designated a type species. Goding (1929a, c) considered Hoplophorion trian- gulum to be congeneric with Ochropepla corrosa and clearly intended (1929a) to pro- pose Metcalfiella as a new genus for the oth- er species previously placed in Hoplopho- rion. Although initially (1929a) ambiguous, Goding later (1929c) gave nomenclatorial priority to Hoplophorion (1901) over Och- ropepla (1869) (perhaps because Hoplopho- rion replaced the older name Hoplophora [1833]). In view of the current rules of zoo- logical nomenclature, Ochropepla has pri- ority. Although Funkhouser (1951:68, foot- note) stated that he agreed with Goding, Funkhouser’s arrangement of the relevant 160 type species contradicted this statement— M. triangulum was not placed as congeneric with H. corrosa, but rather with H. pertusa. Furthermore, given his arrangement, Funk- houser erred in listing Hoplophorion (1901) as a junior synonym of Metcalfiella (1929). Metcalf and Wade (1965) corrected this pri- ority error, listing Hoplophorion as the se- nior synonym of Metcalfiella, but did not question the generic synonymy. Conse- quently, Ochropepla and Hoplophorion were considered valid in Metcalf and Wade’s (1965) catalog, and accepted by most later workers (see Wood and Morris 1974, Deitz 1975, Ceballos-Bendezu 1980, Wood 1984), bringing the nomenclature back to its pre- Goding condition. Our concept of H. corrosa Fairmaire, the type species of Ochropepla, is based on the original description and on a female spec- imen in the Signoret collection (Naturhis- torisches Museum Wien, Vienna, Austria) with labels “‘Bogota|Coll. Signoret.,”’ *““corrosa|det. Signoret.,”’ “‘corrosa| det. Fowler.,” “‘Coll. Nat.-Mus. Wien,” ‘““Hop- lophora,” and ‘‘POSSIBLE|LECTO- TYPE|Hoplophora corrosa|Fairmaire|see Fowler 1894c|det. McKamey & Deitz 1990” (the last a red-bordered label). This female (from Bogota, the type locality; in the Signoret collection, one of two reposi- tories listed by Fairmaire; and examined by Fowler) is likely to be the specimen that Fowler (1894) referred to as “‘the type-spec- imen of O. corrosa,” thus fixing it as the lectotype (Article 74a, International Com- mission on Zoological Nomenclature 1985). The type series of M. triangulum could not be located. Nevertheless, our analysis of the original description in conjunction with species-level revisions of Ochropepla and Metcalfiella confirmed that M. trian- gulum is, as Goding (1929a) believed, con- generic with H. corrosa. This decision will be justified in a forthcoming revision of Ochropepla. Thus, after correcting Goding’s priority decision, the genera here consid- ered valid are Ochropepla (with Hoplopho- rionas its junior synonym) and Metcalfiella. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The pronota of Ochropepla (Figs. 1, 2) and Metcalfella (Figs. 5, 6) are similar (short in length, conspicuously punctate, and with- out horn-like projections), but these genera are apparently distant relatives within the tribe Hoplophorionini (McKamey 1989). Whereas Ochropepla has an r-m crossvein in the hind wing (Fig. 3), Metcalfiella has the veins R,,; and M,,, confluent distally (Fig. 7). Furthermore, Metcalfilla is unique among membracids in that the pronotum has many more pale setae than pits (Fig. 4), rather than just one seta per pit. STATUS OF FAMILY-GROUP NAME HOPLOPHORIONINI GODING, 1926 The family-group name Hoplophorionini is retained in accordance with Article 40a (International Commission on Zoological Nomenclature 1985), although Hoplopho- rion, the nominotypical genus of the tribe, is here considered a junior synonym of Och- ropepla. Goding (1926) proposed the name Hoplophorionini (as Hoplophorioninae) to replace Hoplophorinae Amyot and Serville, 1843, which was based upon the junior homonym Hoplophora Germar. Lists OF NOMINAL SPECIES IN METCALFIELLA AND OCHROPEPLA The following placements, including one new combination, are based on examina- tion of type material (indicated by an as- terisk*) or of original descriptions of the rel- evant nominal species. Forthcoming revisions of Metcalfiella and Ochropepla will include lists of type material examined, spe- cies Synonymies, new species, and institu- tional locations for specimens of each spe- cies. Metcalfiella (all species names represent transfers from Hoplophorion, sensu Met- calf and Wade 1965): 1. Hoplophora apiformis Buckton, 1902 2. Hoplophorion carinulatum Schmidt, 1906 3. Hoplophora cinerea Fairmaire, 1846 VOLUME 93, NUMBER 1 161 * © » Chet a Red ek alt 1 ss oh Ate asa at bl * papa ow” en p Ve Bette Spt bee = OSS PIM IS. Rais+ Mise / / ahd ay a =e + FS aye Figs. 1-7. Comparison of Ochropepla and Metcalfiella. 1, 2, Ochropepla corrosa (Fairmaire), female, head and pronotum. 1, dorsal view. 2, lateral view (including forewing). 3, Ochropepla sp., right hind wing. 4, Metcalfiella sp., electron micrograph of pronotal setation (courtesy of C. H. Dietrich). 5, 6, Metcalfiella pertusa (Germar), female, head and pronotum. 5, dorsal view. 6, lateral view (including forewing). 7, Metcalfella sp., right hind wing. 4. *Hoplophora cinerea obfuscata Fowler, 14. Hoplophorion ohausianum Schmidt, 1894 1906 5. *Hoplophora cribrum Fairmaire, 1846 15. *Hoplophora pertusa Germar, 1835 6. *Hoplophora disparipes Fowler, 1894 16. *Hoplophora porosa Walker, 1851 7. Hoplophorion erectum Schmidt, 1906 17. *Hoplophora proxima Walker, 1851 8. Hoplophorion erectum nigromacula- 18. *Hoplophora pubescens Buckton, 1902 tum Schmidt, 1906 19. *Hoplophora rubripes Funkhouser, 9. *Hoplophora fimbriata Stal, 1862 1922 10. *Hoplophora gigantea Fairmaire, 1846 20. Hoplophora sanguinosa Fairmaire, 11. Hoplophora haenschi Schmidt, 1906 1846 12. Hoplophora monogramma Germar, 21. *Hoplophora semitecta Walker, 1858 1835 22. *Hoplophora signoreti Fowler, 1894 13. *Hoplophora obtusa Stal, 1862 23. *Hoplophora unicolor Fowler, 1894 162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 24. *Hoplophora variegata Fairmaire, 1846 25. *Hoplophora vicina Fairmaire, 1846 Ochropepla (species names 1, 4, 5, and 8 represent transfers from Hoplophorion, sensu Metcalf and Wade 1965): 1. *Hoplophora concinna Fowler, 1894 2. *Hoplophora concolor Walker, 1851 3. *Hoplophora corrosa Fairmaire, 1846 4. *Triquetra hebes Walker, 1851 5. *Ochropepla inaequalis Fowler, 1894 6. *Ochropepla pallens Stal, 1869 7. *Hoplophora punctum Fairmaire, 1846 8. Membracis triangulum Germar, 1821, NEw ComMBINATION: Ochropepla trian- gulum (Germar, 1821) Four species (Ochropepla carinata Funk- houser, 1822; Ochropepla dubia Fowler, 1897; Ochropepla fuscata Fowler, 1897; Hoplophora gloveri Goding, 1893) most re- cently placed in Hoplophorion (Metcalf and Wade 1965) belong to neither Metcalfiella nor Ochropepla and must be referred to oth- er genera (see McKamey 1989). ACKNOWLEDGMENTS We thank C. H. Dietrich, M. H. Farrier, J. W. Hardin, H. H. Neunzig (all of North Carolina State University), and T. J. Henry (USDA, % Smithsonian Institution) for nu- merous helpful suggestions relating to this work. We also thank U. Aspéck (Naturhis- torisches Museum Wien, Vienna, Austria) and the many other individuals and insti- tutions (see forthcoming revisions of Met- calfiella and Ochropepla) that loaned type material. This is paper No. 12390 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, N.C. 27695- 7643; an earlier version formed part of a thesis submitted by S. H. McKamey in par- tial fulfillment of the M.S. degree in ento- mology, North Carolina State University at Raleigh. Funding for McKamey was pro- vided by the NCSU Department of Ento- mology and Alumni Association during the year 1986-1987 and by a National Science Foundation Graduate Fellowship during the period 1987-1989. LITERATURE CITED Ceballos-Bendezt, I. 1980. Nueva sinopsis de los Membracidae (Homoptera: Auchenorrhyncha) del Peru. Revista Peruana de Entomologia 23: 39-58. Deitz, L. L. 1975. Classification of the Higher Cat- egories of the New World Treehoppers (Homop- tera: Membracidae). North Carolina Agricultural Experiment Station Technical Bulletin 225. [iv] + 177 pp. Fowler, W. W. 1894. Order Rhynchota. Suborder Hemiptera-Homoptera. (Continued.) Biologia Centrali-Americana 2: 24-56. Funkhouser, W. D. 1951. Homoptera family Mem- bracidae. Genera Insectorum 208: 1-383. Goding, F. W. 1926. Classification of the Membrac- idae of America. Journal of the New York Ento- mological Society 34: 295-317. 1929a. Notes on some South American Membracidae. Journal of the New York Ento- mological Society 37: 7-9. . 1929b. The Membracidae of South America and the Antilles. IV. Subfamilies Hoplophorion- inae, Darninae, Smiliinae, Tragopinae (Homop- tera). Transactions of the American Entomologi- cal Society 55: 197-330. International Commission on Zoological Nomencla- ture. 1985. International Code of Zoological No- menclature. 3rd ed. International Trust for Zoo- logical Nomenclature, London, University of California Press, Berkeley. xx + 338 pp. McKamey, S. H. 1989. Revision of the Genus Met- calfiella (Homoptera: Membracidae) with Review of the Nomenclature and Systematics of the Tribe Hoplophorionini and Description of a New Ge- nus. Unpublished M.S. thesis, Department of En- tomology, North Carolina State University, Ra- leigh. vil + 315 pp. Metcalf, Z. P.and V. Wade. 1965. General Catalogue of the Homoptera. A Supplement to Fascicle I— Membracidae of the General Catalogue of Hem- iptera. Membracoidea. In two sections. North Car- olina State University, Raleigh. 1552 pp. Wood, T. K. 1984. Life history patterns of tropical membracids (Homoptera: Membracidae). Socio- biology 8: 299-344. Wood, T. K. and G. K. Morris. 1974. Studies on the function of the membracid pronotum (Homop- tera). I. Occurrence and distribution of articulated hairs. Canadian Entomologist 106: 143-148. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 163-175 NEW AND LITTLE-KNOWN SPECIES OF FORCIPOMYIA (DIPTERA: CERATOPOGONIDAE) ASSOCIATED WITH COCOA POLLINATION IN BRAZIL WILLIS W. WIRTH Research Associate, Florida State Collection of Arthropods, and Cooperating Scientist, Systematic Entomology Laboratory, U.S. Department of Agriculture, 1304 NW 94th St., Gainesville, Florida 32606. Abstract. —Two new species of Forcipomyia Meigen that are common in cocoa plan- tations in Brazil, and are involved in cocoa pollination, are described and figured: For- cipomyia (Forcipomyia) soriai Wirth and Forcipomyia (Lepidohelea) winderi Wirth. A key is given to distinguish the subgenus Forcipomyia s. str. from the closely related subgenus Lepidohelea Kieffer. Keys are also presented to distinguish the Neotropical species of the squamitibia Group of Forcipomyia s. str. which includes F. soriai, and the Neotropical species of the bicolor Group of Lepidohelea, which includes F. winderi. Information on the immature stages and biology for these and related species is summarized and the immature stages of F. (F.) pinamarensis Spinelli, also associated with cocoa, are described and figured. Key Words: Biting midges of the genus Forcipomyia Meigen are important pollinators of cocoa (Theobroma cacao L.) wherever it is grown throughout the tropics. A voluminous lit- erature has accumulated in the past 30 years dealing with the taxonomy, biology, and re- lation to cocoa pollination of numerous spe- cies of Forcipomyia, especially of the sub- genus Euprojoannisia Bréthes. That which appeared up to 1977 was reviewed by Win- der (1978). Approximately three-fourths of the 70 species of Ceratopogonidae that have been collected in cocoa flowers belong to the genus Forcipomyia. Nearly a third of the Forcipomyia midges belong to the subgenus Forcipomyia s. str., while in decreasing or- der of importance the subgenera Eupro- jJoannisia Bréthes, Thyridomyia Saunders, Lepidohelea Kieffer, Microhelea Kieffer, Warmkea Saunders, and Lasiohelea Kief- fer, are also involved. There is considerable Ceratopogonidae, Forcipomyia, cocoa, pollination variation in the abundance and importance in pollination between these taxa, between individual species, from region to region, and also with the season. From 1967 to 1983 intensive studies were carried out by John A. Winder and Saulo de Jesus Soria at the Cacao Research Center (CEPLAC) at Itabuna, Bahia, Brazil. More recently Winder has resumed his cocoa pol- lination studies at the Fazenda Almirante, near Itajuipe in Bahia, and I have had the opportunity to cooperate in the determi- nation of the ceratopogonids from his col- lections. In Bahia, and especially at the Fa- zenda Almirante, by far the most abundant ceratopogonids collected in the cocoa plan- tations belong to the Forcipomyia bicolor Group in the subgenus Lepidohelea, and the F. squamitibia Group in the subgenus For- cipomyia. The purpose of this paper is to describe the two most important species in 164 Winder’s collections to make the names available for his biological studies, and to provide means for distinguishing them from closely related species. Explanation of the taxonomic characters used can be found in the general papers on Ceratopogonidae by Wirth et al. (1977) and Downes and Wirth (1981), and the revision of the North American Euprojoannisia by Bystrak and Wirth (1978). Holotypes and allotypes of new species are deposited in the National Museum of Natural History (USNM), Smithsonian Institution, Wash- ington, D.C.; paratypes as available in the Florida State Collection of Arthropods, Gainesville, Florida; California Academy of Sciences, San Francisco, California; Museo de La Plata, La Plata, Argentina; Instituto Oswaldo Cruz, Rio de Janeiro, Brazil; Brit- ish Museum (Natural History), London; and Museum National d’Histoire Naturelle, Paris. Taxonomic work on the cocoa-pollinat- ing Forcipomyia midges has as its main pur- pose the identification of the most impor- tant pollinating species, the location of their most important breeding places, and the ac- cumulation of enough information on their biology so that a practical regimen of cocoa culture and midge habitat manipulation can be devised to maximize cocoa pollination and crop production (Young 1982, 1983). Although the study of Forcipomyia breed- ing places and identification of immature stages has been pursued intensively in America since the early work of Saunders (1956, 1959), progress has been difficult and slow. Nevertheless, important contribu- tions have been made by Winder and Silva (1972), Winder (1977a), Soria et al. (1979), and Soria and Wirth (1979) in Brazil, and Young (1982, 1983, 1986) in Costa Rica. Larval habitats of Forcipomyia midges are somewhat correlated with their taxonomic group. The habitats of Euprojoannisia spe- cies are usually semiaquatic situations such as algae-covered rocks or mud, wet moss or leaves, mats of decaying aquatic vegetation, and leaf axils of water-holding plants. Spe- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cies of Thyridomyia usually are found in association with moss or algae in rather damp habitats. Immature stages of Warmkea species are frequently found in leaf axils of Pandanus, aroids, and epiphytic and terrestrial bromeliads, and less often in rotting cocoa leaves on the ground, in ba- nana stems, and in bracts of Heliconia. Spe- cies of the subgenera Forcipomyia, Lepi- dohelea, and Schizoforcipomyia are less aquatic and are more commonly associated with rotting plant material, which in cocoa plantations and their environs often in- volves heaps of cocoa pods, old banana stems, cocoa leaves, coconut debris, bracts of Heliconia and Calathea, and rotting fruits of coconut, calabash, palm nuts, etc. The taxonomic key to the subgenera of Forcipomyia published by Wirth and Ra- tanaworabhan is out-dated and has been re- placed by that of Debenham (1987a). Until the work of Debenham (1987a, b, c,) it was not possible with confidence to distinguish between the subgenera Forcipomyia s. str., Lepidohelea, and Schizoforcipomyia, but she pointed out characters in all stages to make possible a key to these taxa for the Western Hemisphere. KEY TO SUBGENERA OF FORCIPOMYIA RELATED TO LEPIDOHELEA 1. Hind tarsal ratio about 1.0; wing usually with pale costal spot or pale mottling; second radial cell short, costal ratio about 0.5 — Hind tarsal ratio less than 0.5 or greater than 1.5; wing usually without pale markings; costal ratio and second radial cell various ys oe miscellaneous subgenera of Forcipomyia 2. Body wings, and/or legs with flattened striated scales ranging from long and 1-striated setae to broad, short, multistriated scales; palpus with third segment more or less spindle-shaped, slightly more swollen on basal half with sensory pit located at about '4 length; legs with char- acteristic banding of alternating pale and dark bands, at least on hind tibia; male genitalia often with whitish dististyle and bicolored ninth sternum and basistyle; dististyle without long setae on outer margin; aedeagus various; larva with b hairs of body, and p and q head hairs swollen near base and becoming filamentous VOLUME 93, NUMBER 1 distally, usually some about as long as body of larva; anal pseudopod of larva cleft with each branch bearing 6-8 hooklets apically and 3-4 hooklets on inner lateral side — Body, wings, and legs rarely with flattened stri- ated scales; palpus with third segment swollen on basal '3—'/, slender distally, with sensory pit located at basal 4; legs, if banded, not with alternating pale and dark bands on hind tibia; male dististyle not contrasting whitish, but with well-developed setae on outer margin; aede- agus shield-shaped, apex pointed with median ridge; larval b hairs and p and q head hairs not as above; anal pseudopod of larva not cleft, bearing unbroken row of strong hooks ...... 5 pe SAR ee ee Cee Subgenus Forcipomyia 3. Male parameres with basal apodemes forming a U-shaped arch, parameres a pair of slender to stout, straight or bowed, well-sclerotized blades or rods reaching to no more than middle of basistyles; aedeagus various, with low to high basal arch, distal process various, often with recurved points; dististyle straight or slightly curved, tapering to slender tips; palpus 5-seg- mented; two spermathecae; scales of body and legs never short and broad Subgenus Schizoforcipomyia — Male parameres separate or indistinctly joined in an anteromesal bridge; parameres various, usually straight slender rods nearly as long as basistyle and tapering to filamentous tip; ae- deagus usually much longer than broad, basal arch low or absent, usually stout distally; dis- tistyle often sinuate with expanded tip; palpus 4- or 5-segmented; one or two spermathecae; scales of body and legs often short and broad ation cho he ee ee ere ee Subgenus Lepidohelea Subgenus Forcipomyia, Ss. str. Wirth (1982) discussed the taxonomy and biology of the Neotropical Forcipomyia s. str. associated with cocoa pollination, in- cluding four species which he placed in the “argenteola Group.” He also diagnosed and keyed out F. (F.) squamitibia Lutz, which with similar species could be confused with species of the argenteola Group. Species of these two groups can be separated by the following key: 1. Female tibiae with row of hastate spines on extensor surface; pale wing spot at end of costa larger, including most of second radial cell; hind tarsal ratio about 1.0; male parameres with slender common base; pupa without promi- 165 nent setose tubercles; respiratory horn large and globular with spiracular openings in a straight TO Wee hte err ke squamitibia Group — Female tibiae without hastate spines; pale wing spot smaller, not covering end of second radial cell; hind tarsal ratio 0.50-0.76; male para- meres with bases joined in a broad plate; pupa with setose tubercles that are often elongate and prominent; respiratory horn small to mod- erate in size, with longer neck and spiracular openings in a more or less irregular row .... pe RY ey Ray set peal EN etn cre ted argenteola Group Forcipomyia (Forcipomyia) pinamarensis Spinelli Figs. 1-11 Forcipomyia (Forcipomyia) pinamarensis Spinelli, 1983: 121 (male, female; Argen- tina; figures). Types. — Holotype female, allotype male, 12 female, 29 male paratypes: Argentina, Buenos Aires Prov., Pinamar, Partido de General Madariaga, 27.111.1981, G. R. Spi- nelli (in Museo de La Plata). Note.—Shortly before his death in 1968, Professor L. G. Saunders of the University of Saskatchewan very generously donated to the Smithsonian Institution in Washing- ton his extensive worldwide collection of Forcipomyia midges, mostly reared from the immature stages, along with his manuscript notes and drawings. Among his manuscript notes were drawings and the description of a species that he collected and reared in Bra- zil in 1923 and designated as ““B-70.”’ Dur- ing the present study it was determined that Saunders’ ““B-70” was the same species that Spinelli described as Forcipomyia pina- marensis from Argentina. This species is abundant and widespread in cocoa planta- tions; therefore this opportunity is taken to publish Saunders’ description and figures of the adults and immature stages, and to list new distribution records. The following de- scriptions are adapted from Saunders’ manuscript notes: “Larvae of this species were taken in fair numbers under the bark of a rotting log in the forests of Corcavado, Rio, 18.vi1i.1923. 166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. l-11. Forcipomyia pinamarensis: 1-4, female; 5, 6, pupa; 7-10, larva; 11, male: 1, antenna; 2, radial veins of wing; 3, hastate spines of tibia; 4, hind leg; 5, respiratory horn; 6, pupa; 7, last two abdominal segments; 8, head and body setae, lettering as in text; 9, 10, integumental armature of body; 11, genitalia, partial (drawn by L. G. Saunders; scale in microns). VOLUME 93, NUMBER 1 Pupae and adults were obtained from them in due time.: “Larva. Color white, the macrochaetae black, their papillae light brown. Head ochreous to black around mouth. Protho- racic pseudopod of moderate length, forked in its distal half, with the crown of hooks disposed equally over the end of each branch. Chaetotaxy (Fig. 8): p and q hairs of head lanceolate; a hairs of body lanceo- late; b and d separate, b short, curved, spi- nulate distally. Dorsal hairs of last abdom- inal segment arising from a cap of chitin (Fig. 7). Cuticular armature in form of short stout stem with a number of apical arms (Fig. 10); simple cones or hooks on pleura and last abdominal segment (Fig. 9). Cauda pointed. Anal blood gills bilobed, the outer lobe much smaller than the inner. ‘‘Pupa (Fig. 6). Color (exuviae) pale ochreous throughout. Head devoid of pa- pillae. Thorax with one dorsal pair of setig- erous papillae and three other pairs of very small papillae. Abdomen with no cuticular spines, few microchaetae. Terminal abdom- inal processes of moderate length. Protho- racic respiratory horn (Fig. 5) in form of subspherical knob on short, broad-based stem. About 16 spiracular papillae having small external membranes curve up the pos- terior side and over the top. Length 2.3 mm. ‘Adult Female. Face pale, ocellar triangle faintly brown; dark ring around bases of antennae. Palpus brown with light bands at distal articulations; segment three much swollen in proximal one-third, distal neck tapering gradually; proportional length of segments, 9:11:37:21:13. Antenna (Fig. 1): Scape light brown, flagellum white tinged faintly with brown distally; proximal seg- ments flask-shaped with broad elongated neck; proportion of first 8 to last 5 segments, 43:34; last 5, 6:6:6:6:10; terminal style a large knob with indented extremity. Thorax uniformly brown on dorsum, with coarse golden pubescence; humeral angles pale; scutellum brown to dark brown around margins; postscutellum dark brown. Pleura 167 light brown on all the usual sclerites; a very small sclerite in front of the base of the wing dark brown. Legs ochreous, fore and middle coxae brown, hind coxa paler; a fuscous band just before extremity of hind femur (Fig. 4); duskiness of tibiae due to presence of nar- row black squamous hairs; outer surface of all tibiae with a few ochreous lanceolate to squamous hairs (Fig. 3), not striated like the more usual decumbent hairs of legs and ab- domen. Proportional length of hind tarsal segments, 15:12:7:6:5. Wing densely clothed with dark hairs, narrowly squamous; area of pale yellowish hairs at end of costa. Branches of radius dividing to form distinct cell (Fig. 2); end of costa at middle of wing, fork of Cu directly below. Halteres white. Abdominal tergites uniformly brown except 8 + 9 which are pale ochreous with faint lateral brown spots; sternites 3—7 pale brown with stronger lateral coloring, 8 yellowish. Cerci pale. Tergites and pleural membrane clothed with narrow black-striated hairs; pale simple hairs on venter. Spermathecae two; 0.090 by 0.056 mm, subspherical, no chitinization on neck. Length 1.6 mm; wing 2.11 by 0.45 mm; antenna 0.6 mm. ‘““Male. Specimen very imperfect. Palps much as in female, but terminal segment longer; proportional lengths, 9:11:38:20:16. Hypopygium (Fig. 11): Aedeagus long, rounded distally, hairy, without lateral ca- rinae. Parameres joined in middle, a slender pair of bowed stylets extends backwards be- yond end of aedeagus. Side pieces of forceps long, tapered slightly, with distinct brush of fine hairs on inner side near anterior end; claspers lost. “This is a very typical Forcipomyia con- forming closely to type. The larvae are very close to F. bipunctata (Linnaeus), differing only in the presence of a chitinous cap on the last abdominal segment and in the ex- traordinary dorsal cuticular armature. The very bare, spineless pupa is distinctive, pos- sessing as it does a row of unusually small respiratory papillae on the prothoracic horn. The adult female belongs to the small group 168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON exhibiting a row of lanceolate scales on the tibiae; apart from this character they may readily be distinguished from most other species by the minute dark sclerite in front of the wing base.” Distribution.— Argentina, Brazil, Costa Rica, Panama. Specimens examined.—ARGENTINA: Pinamar, Bs. As., 27.111.1981, G. Spinelli, 1 female (paratype). BRAZIL: Bahia, Itabu- na, 1970, J. Winder, emergence traps, 5 males, 6 females; Bahia, Ilheus, CEPEC, 15.vi.1974, S. Soria, 1 female; Para, km 3, Rodovia Castanhal-Curuca, Faz. Agricul- tura Bandeirante, 31.v.1980, S. Soria, ex cacao cherelles, 4 males, 2 females; Rio de Janeiro, Corcavado, 18.viii.1923, L. G. Saunders, under bark, 4 larvae, | pupal ex- uviae, 2 females. COSTA RICA: Heredia Prov., vic. La Virgen, Finca La Tigra, 30.vi- 3.vii.1980, 25.vii.1981, 11,30,vii.1982, A. M. Young, 2 males, 3 females. PANAMA: Canal Zone, Pipeline Road, vii.1967, F. S. Blanton, light trap, 1 female; Canal Zone, Balboa, fogging Cassia tree, 19.vii.1979, E. Broadhead, 1 female; Canal Zone, Gatun, humid forest, fogging Spondias and Cordia tree canopy, 9-10.v1i.1979, E. Broadhead, 2 males, 3 females. Forcipomyia (Forcipomyia) soriai Wirth, NEw SPECIES Figs. 12-21 A hairy yellowish species with dark hal- teres; female with abundant long, dark, one- striated scalelike setae on radial field of wing and on abdomen; wing with one small yel- lowish costal spot; tibiae with row of blunt- pointed, pale, hastate setae; hind tarsal ratio 0.92. Allotype female. — Wing length 1.14 mm; breadth 0.52 mm; costal ratio 0.47. Head: Pale brownish; overall clothed with numerous long, brown, one-striated setae giving a bushy appearance. Antenna (Fig. 13) with lengths of flagellar segments in pro- portion of 20-15-15-15-15-15-15-16-15-15- 16-16-22, antennal ratio (1 1—15/3-10) 0.67; distal segments unusually short for the ge- nus; proximal verticils appearing bushy. Palpus (Fig. 14) with lengths of segments in proportion of 10-11-22-13-10; third seg- ment slightly swollen at base (palpal L/W ratio 2.2), gradually tapering to tip, a small, round, shallow, sensory pit near base; seg- ments 4 and 5 with faint separation. Man- dible without teeth. Thorax: Pale yellowish brown, darker on pleura, scutellum, postscutellum, and cox- ae. Mesonotum with suberect fine yellowish hairs. Legs uniformly yellowish except tib- iae distally and all tarsi proximally, slightly infuscated; with yellowish and brownish, fine hairs, no broad flat scales; tibiae, and tarsi to a lesser degree, with long, slender, brown setae on extensor side; tarsomeres 1-3 with row of strong, blackish, ventral spines, a pair at apices of tarsomeres much stronger; all tibiae with a row of erect, nearly hyaline, hastate setae (Fig. 12) with petiolate bases and bluntly pointed or rounded tips. Claws moderately long and slender, strongly curved, tapering to sharp tips; empodium small. Hind leg with lengths from femur to tarsomere 5 as 82-80-24-26-16-12-8:; tarsal ratio (T1/T2) 0.92; hind tibial spur yellow- ish, straight, slender, tapering to sharp point. Wing densely and uniformly covered with brownish macrotrichia, except two anterior patches of blackish, one-striated, scalelike macrotrichia over radial field and just past end of costa, the two patches separated by a small pale spot of yellowish scales at end of costa. Halter brownish. Abdomen: Brownish, darker posteriorly with numerous long, brown setae, especially on pleura and on posterior segments. Sper- mathecae (Fig. 15) two, dark brown, rela- tively large; unequal, measuring 0.118 by 0.087 mm and 0.087 by 0.075 mm; the larg- er Oval, the smaller subspherical; both with lightly sclerotized, slender, almost thread- like entrances to the ducts. Holotype male.— Wing length 1.21 mm; breadth 0.43 mm; costal ratio 0.44. Similar to the female with the usual sexual differ- VOLUME 93, NUMBER 1 Figs. 12-21. 169 Forcipomyia soriai: 12-15, female; 16, pupa; 17, 18, male; 19-21, larva: 12, hastate spines of tibia; 13, antenna; 14, palpus; 15, spermathecae; 16, respiratory horn; 17, aedeagus; 18, genitalia, aedeagus omitted; 19, head; 20, body segment, lateral view; 21, last two abdominal segments, dorsal view (lettering as in text; scale in microns). ences. Antenna with well-developed plume of yellowish-brown verticils; lengths of fla- gellar segments in proportion of 25-15-15- 15-15-15-15-15-16-55-28-20-32, antennal ratio (12-15/3--11) 1.03. Palpus with lengths of segments in proportion of 9-11-27-12- 10; third segment more slender at base than in female, palpal ratio 3.4. Legs with exten- sor setae longer than in female, hastate spines absent on tibiae; tarsi with numerous long, slender, black spines ventrally; claws long, slender and curved; empodium slender with short rays. Hind leg with lengths from femur to tarsomere 5 as 86-80-24-26-17-13-10; 170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tarsal ratio 0.92; tibial spur pale yellow, straight, slender, 3 as long as basitarsus, tapering to sharp point. Wing with pale yel- lowish macrotrichia overall, a small clump of strong blackish macrotrichia over first radial cell and another just past end of costa, the two separated by a small area of yellow- ish macrotrichia. Halter brownish. Abdo- men yellow, with brownish pigmented areas laterally on each segment; with abundant, long, brownish, bristlelike setae, without broad striated scales. Genitalia (Fig. 18).— Ninth sternum about twice as broad as long, caudal margin slight- ly convex with shallow mesal concavity; basistyle about twice as long as broad, with- out stout setae on mesal face but with nu- merous long, bristlelike setae laterally and distally that are about twice as long as dis- tistyle, the latter straight, slender, and ta- pered to slender tip. Aedeagus (Fig. 17) shield-shaped, broad basally, length equal to basal breadth, with short basal arms, the basal arch only a slight concavity; sides con- vex, tapering to a blunt caudomedian point. Parameres with slender, straight, basal apo- demes; joined mesally on basal fourth; dis- tal portions relatively stout, nearly straight and parallel, with the blunt tips slightly curved laterad. Larva.—Length 4-5 mm when mature. Color yellowish white; head capsule pale brownish; dorsal body hairs borne on brownish conical prominences (Figs. 20, 21). Hairs p and q of head (Fig. 19) long and slender, with spear-shaped tip, that of p hair much larger than g. Antenna long and ta- pering, borne on a large subconical pedestal. Body hairs as in Fig. 20; a hairs with long slender stem and spear-shaped tip, even on last segment (Fig. 21). Cuticular armature absent. Prothoracic pseudopod cleft, each lobe with 6-7 small brown hooks; posterior pseudopod with a row of 14-16 brown hooks. Cauda short but pointed. Pupa.—Exuviae only, length not mea- sured. Color pale yellowish, integument without spinules or shagreening. Respira- tory horn (Fig. 16) irregularly oval in out- line, with very short, stout peduncle; in cross-section appearing slightly flattened with the spiracular openings forming a slight rim; 16-18 spiracular openings in a straight row up posterior side and around tip. Body tubercles not developed, setae scarcely vis- ible under high magnification. Distribution. — Brazil, Puerto Rico, Trin- idad. Types. — Holotype male, allotype female, Brazil, Itajuipe, Fazenda Almirante, 9.v.1988, J. A. Winder, in emergence trap (deposited in USNM). Paratypes, 37 males, 44 females, | larva, 2 pupal exuviae: Brazil, same data as types, but dates iv.1987- vul.1989, 28 males, 35 females; Bahia, Ita- buna, 1970, J. A. Winder, emergence trap, 6 males, 4 females; Bahia, IIheus, CEPEC, 4—9.viii.1972, S. Soria, at flowers of cacao, 1 male, 3 females. Puerto Rico, Maricao, 21.1.1953, L. G. Saunders, reared from ba- nana leaves, | larva, 1 pupal exuviae, | male. Trinidad, no locality, 7.v.1953, L. G. Saun- ders, reared from wet leaves in stream, 1 larva, 1 pupal exuviae, | male. Discussion.— This species is dedicated to Saulo de Jesus Soria of Bento Goncalves, R.S., Brazil, in recognition of his long and continued interest and his many important contributions pertaining to the role of cer- atopogonids in cocoa pollination through- out Central and South America. Lane’s description and figure of the male genitalia of F. argenteola Macfie from Brazil probably refers to F. squamitibia Lutz, as all species of this group have the hind tarsal ratio of about 1.0 and the parameres with narrow basal separation and quite slender proximally. Forcipomyia argenteola, to the contrary, belongs to a group of species re- viewed by Wirth (1982) which has the hind tarsal ratio about 0.5 and the male para- meres with bases fused in a broad plate with posterior concavity and the caudal process- es relatively stout and tapering to a sharp- pointed tip. Wirth (1982) described the male of F. argenteola from males with associated VOLUME 93, NUMBER 1 females from Nova Teutonia, Brazil, the type locality. Species of the Forcipomyia squamitibia Group can be distinguished by the following key: KEY TO SPECIES OF THE FORCIPOMYIA SQUAMITIBIA GROUP 1. Hastate spines of female tibiae in two dense irregular rows; large dark brown species, wing length 1.7-2.0 mm; hind tarsal ratio 1.0 .... 5 cid 6 b-6 CRSA ee ane ae chilensis (Philippi) — Hastate spines of female tibiae in one sparse row; large or small, pale or dark species ..... 2, 2. Mesonotum with dark vittae; small dark brown species, wing length 0.93 mm; hastate tibial spines brown, short, and broad with sharp tip; hind tarsal ratio 0.9 ........... sexvittata Wirth — Méesonotum not vittate; color, size and tibial SDINESAVATIOUSE tes cavsiie oo ec eters: « 3 3. Hastate tibial spines longer than width of tibia 4 — Hastate tibial spines as long as width of tibia OsSHOTLe Tarp re cere secur. sec Se cea emer ney 5 4. Yellowish brown species with unmarked yel- lowish legs; tibial spines slightly longer than width of tibia, swollen in midportion, tapering to sharp tip; antenna pale; third palpal segment swollen on proximal 3; wing length 1.3 mm; hind tarsal ratio 1.0; male parameres with fi- liform tips straight and parallel squamitibia Lutz - Dark brown species with brown legs; tibial spines much longer than width of tibia, slender and only slightly swollen in midportion; an- tenna dark brown; third palpal segment swol- len on proximal 7; wing length 1.5 mm; hind tarsal ratio 1.3; male unknown ............ é Sfadae cihtoene ene tenuisquamipes Wirth 5. Legs pale yellow, hind femur with narrow apex brownish; hastate tibial spines petiolate with broad apices; wing length 1.1 mm; hind tarsal ratio 0.9; spermathecae pale; male parameres with filiform tips crossing (Fig. 11) ......... 5 Oo Let eR ae IEE TO Oe eae pinamarensis Spinelli — Legs dusky yellowish with tibiae brownish dis- tally, hind femur without distal dark band; has- tate tibial spines slightly petiolate with tips moderately broad and slightly tapered to point; wing length 1.14 mm; hind tarsal ratio 0.92; spermathecae dark brown; male parameres stout distally with tips abruptly bent a short wayaigivergent (Fig. 8)! 222 SA eae Subgenus Lepidohelea Kieffer Debenham (1987b) has presented an ad- mirable discussion of the difficult taxonomy al of the subgenus Lepidohelea, including characters that may be used to separate the species groups occurring in the Australasian Region. The American species appear to fall in two groups differing from the Old World species, most of which would fall within what Debenham has named the chrysolopha Group. The American species may be dis- tinguished from the chrysolopha Group by the following key: 1. Palpus with four segments; one spermatheca; male dististyle more or less expanded distally (Western Hemisphere species) ............. ee Ber eT A sei ne annulatipes Group — Palpus with five segments; two spermathecae; male dististyle various ..................... 2 2. Male dististyle straight or slightly curved, ta- pering to slender tip (Western Hemisphere spe- CIES) ay ess ean TORU teed tn rtae tree: bicolor Group — Male dististyle straight to sinuate, tip more or less expanded (Eastern Hemisphere species) Be at 1 LD. ate Ba a Pe eae chrysolopha Group Forcipomyia (Lepidohelea) bicolor Species Group Forcipomyia (Lepidohelea) winderi Wirth, NEw SPECIES Figs. 22-30 A hairy, predominantly yellowish species with brownish markings; female wing dark with one large pale anterior spot, male wing pale except for dark costal field; palpus stub- by, third segment not swollen, with small shallow pit; antenna moderately long; legs banded with bicolor Group type markings, femora pale except hind femur with apical dark brown band, all tibiae and tarsi band- ed; male genitalia with brownish markings laterally on ninth sternum, distal halves of basistyles, and all of dististyles. Female holotype.— Wing length 1.11 mm, breadth 0.44 mm: costal ratio 0.47. Head: Pale brownish, overall clothed with numerous long, brown, one-striated setae giving head a bushy appearance, especially strong and numerous on clypeus. Antenna (Fig. 22) with lengths of flagellar segments in proportion of 20-16-16-16-16-16-15-15- 20-20-20-20-23, antennal ratio 0.79; anten- 172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON N ~ ‘fi! RES SESS Laie = ip SS SSSSAS SRR UMips SSS SERS SS SIE ~ Na / = RNA Dis SS AN TAN rn RAW wary TAN 3= = Soa SS mae ani 2 Rocco isa a) Ge oe See es Se Peers Ones Sea Se wn DMA 96 iat ee yey, TID —G eae LR Peels Per es Arid ray igi ts, Figs. 22-30. Forcipomyia winderi: 22-25, 29, 30, female; 26-28, male: 22, antenna; 23, palpus; 24, 26, wing; 25, striated scales of body and legs; 27, aedeagus; 28, genitalia, aedeagus omitted; 29, fore, mid, and hind legs (top to bottom); 30, spermathecae (scale in microns). nal verticils short and strong. Palpus (Fig. 23) with lengths of segments in proportion of 8-12-17-9-8; third segment slightly swol- len in midportion, spindle-shaped, with un- usually small, shallow, round, sensory pit. Mandible without teeth. Thorax: Reddish brown; mesonotum somewhat paler in midportion, clothed abundantly with long, pale brown, bristly, erect setae and fine, yellowish, appressed hairs. Scutellum paler in midportion. Legs (Fig. 29) yellowish, with brownish bands as figured: fore and mid femora pale, tibiae with narrow basal and broad apical bands; hind femur with broad apical, and hind tibia with broad basal and narrow subapical bands; bands more intense on hind leg; fem- ora and tibiae with some long, brownish, bristly setae, and abundantly clothed with broad, sharp-pointed, 4- or 5-striated scales (Fig. 25). Hind tibial spur short and pointed, nearly straight, about as long as breadth of tibia. Tarsomeres dark banded in midpor- tion; hind tarsal ratio 1.07; claws slender VOLUME 93, NUMBER 1 and curved. Wing (Fig. 24) dark brown, abundantly clothed with long, one-striated macrotrichia, these especially stout and dense over radial veins, this area appearing blackish; a moderately large pale anterior spot just past end of costa in cell R5, the macrotrichia pale yellowish in this area. Halter pale. Abdomen: Pale yellowish brown proxi- mally, becoming dark brown distally; clothed with sparse setae, and abundantly with appressed, flattened, striated, sharp- pointed scales (Fig. 25), latter varying in width, and with 2 to 5 striations. Two rather small ovoid spermathecae (Fig. 30) without distinct necks; slightly unequal, 0.072 by 0.046 mm and 0.061 by 0.043 mm. Male allotype.— Wing length 1.30 mm, breadth 0.43 mm; costal ratio 0.42. Similar to the female with the usual sexual differ- ences. Wing (Fig. 26) much narrower, uni- formly pale yellowish, with fine macrotrich- ia except costal field provided with abundant, broad, scale-like macrotrichia forming a prominent dark area, another small, dark-scaled area on anterior margin at midlength of cell RS. Antenna with prominent plume of strong verticils, these brown on basal half of plume, yellowish on distal half; lengths of flagellar segments in proportion of 26-15-14-14-15-15-15-16-16- 17-33-30-35, antennal ratio 1.00. Palpus slender; lengths of segments in proportion of 2-13-17-10-10; palpal ratio 2.4. Hind leg with lengths from femur to tarsomere 5 as 94-96-28-30-20-13-10; tarsal ratio 0.93; tibial spur longer than in female, 1.3 times as long as apical breadth of tibia. Abdomen yellowish; terga with posterolateral corners brownish forming broken segmental bands; distal segments provided with broad stri- ated scales, increasing in breadth and num- ber posteriorly, ninth tergum with dense broad scales. Genitalia (Fig. 28): Ninth sternum slight- ly concaved caudad, brownish on postero- lateral corners; basistyle moderately broad, brownish on distal half; dististyle slender 173 and nearly straight, entirely brownish. Ae- deagus (Fig. 27) unusually long, more than twice as long as basal breadth; basal arms unusually short, sides slightly convex, ta- pering gradually to pointed tip which ap- pears to be slightly cleft. Parameres forming nearly straight, slender rods reaching level of tip of aedeagus, their bases broadly sep- arated on the strongly sclerotized, mbbon- like connective between the basistylar apo- demes. Immature stages.— Unknown. Distribution. — Brazil, Eucador. Types.— Holotype female, Brazil, Bahia, Itajuipe, Fazenda Almirante, 4.v.1988, J. A. Winder; allotype male, same data but 23.v.1988 (in USNM). Paratypes, 76 males, 37 females, as follows: Brazil, Bahia, same data as types, but dates v.1987-x.1989, 50 males, 19 females; Bahia, Itabuna, 1970, J. A. Winder, reared from rotting cacao pods, 1 male, 1 female; same, but taken 1970- 1971 in emergence traps, 20 males, 12 fe- males. Ecuador, Quemado, Pichilingue, N.I.A.P., iv—v.1978, J. Mendoza, reared from rotting vegetation, 5 males, 5 females. Discussion.—This species is named in honor of John A. Winder of Curitiba, Brazil in recognition of his outstanding research on the ceratopogonid pollinators of cocoa in Brazil. Winder (1977a: 58) reported that this spe- cies (listed as a banded-legged species of Forcipomyia (Forcipomyia) comprised 3.8% of the total catch of ceratopogonid midges collected in cocoa flowers at CEPLAC in Bahia, Brazil. This species was also reared from rotting cocoa pods. KEY TO NEOTROPICAL SPECIES OF THE FORCIPOMYIA BICOLOR GROUP 1. Femora dark from base to tip (narrow knee spots may be pale, narrow bases may be pale) 2 — Femora extensively pale proximally, may be entirelygpaleyess fers eee ea Peis + 5 2. All tibiae entirely dark; antenna long, seg- ments with narrow base swollen, abruptly narrowed to long, slender, distal portion; 174 S(t): PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mesonotum with four pairs of lines of white Scales; Sayin eee squamithorax Clastrier At least hind tibia with broad sub-basal pale band; mesonotum and hind tibia various .. 3 Antenna short, segments short tapering; all tibiae with broad sub-basal to median pale bands (faint on fore leg); female wing with- out pale spot or pale areas; small species, wing length 1.1-1.5 mm .. abercrombyi Macfie Antenna long, segments with narrow bases swollen, abruptly narrowed to long slender distal portion; larger species, wing length | .3— IE Semin yarn ais mrata neh were 4 Basal dark brown band in hind tibia narrow, not as broad as second band; hind femur with narrow distal pale band at knee distinct NEE hss San bee aerated dubia Macfie The two dark brown bands on hind tibia about equal in width; hind femur with nar- row distal pale band at knee indistinct ... DOr er ee ON ey Pee Creer lacrimotorii Macfie Fore and mid femora entirely pale; female wing mottled with pale and dark areas; an- tenna short and stubby; third palpal segment Withtdeepepitw 7 se eee te seminole Wirth Fore and/or mid femora with brown bands; antenna long; female wing dark or mottled; palpal pit various Fore and mid femora pale except mid femur with basal dark band ...... flavifemoris Macfie Fore and mid femora dark on distal half or more Legs pale with narrow dark bands; female wing mottled; third palpal segment of female slender with shallow pit; spur of hind tibia slender, straight, pale; female genital scler- otization without spines or long setae; male basistyle and dististyle with pale bases ... a et et Bee GB is winderi new species Legs with extensive, broad dark bands; fe- male wing dark; third palpal segment of fe- male swollen to past midportion with deep pit; spur of hind tibia long, curved, dark with pale base; female genital sclerotization with tuft of long setae on ends, with several short spines along midportion; male basistyle and dististvlewanious?.: a ser SS \ < oe a a > 4 oe 6. “ue vg peta, 4>8° a8 a, «Os ww PRL Be POAT SS SS \ Sy, SS ac ASS ( 31 oO RAK . \ C4 \ os on -@ axe “ge z \ v7 poe \ > 7 PAR sa, 817” fe SS \ .oia R~ ‘e@ \ @S ofQ Sige: fo) - \ Ro \ i J af, g J a3 TAYE SEE ERTS al 6, 218 ral , \ era pee \ Vo ag ¢ a1 8" Or 8B, g42 BY Ye 8 2 Aptos ~ 7 dat @ By fre 164» gee, rAd, oe \ | | \ \ \ \ Mel : \ \ \| \l Se eS Se — == — 189 190 guishable from, and continuous with, rows of dorsal setae; about 8 on head; abdominal setae 13 (10-14) long; longest near coxae (Fig. 2E) 15 (14-15) long. Antennae (Fig. 2F) well developed, apparently 9-segment- ed, no specimens with antennae complete. Segments VI, VII and VIII with fleshy setae. Although segment 9 was missing on all available specimens, other species of Mat- sucoccus in North America with 9-seg- mented antennae have 4—S setae and 2 fleshy setae on segment 9. Segment II with 3-6 campaniform sensilla. Segments III though VI with 0-2 intersegmental sensilla and 0- 3 coeloconic sensilla. Scape with a dorsal cluster of 11-14 setae, 8 (7-9) long; longest seta on segment II 41 (38-43) long; fleshy setae 24 (21-26) long. Legs (Fig. 2G) well developed, 1084 (985-1204) long; 2 claw digitules and 2 setiform tarsal digitules. Trochanters with 10-13 sensoria and | long seta. First tarsal segment with 2 campani- form sensilla. Thoracic spiracles (Fig. 2H) typical of genus, anterior pair 80 (65-88) long from orifice to tracheae, width of atri- um 32 (32-33), width of sieve 12 (12-12); posterior pair 75 (67-85) long from orifice to tracheae, width of atrium 37 (33-40), width of sieve 14 (12-15). Seven pairs of abdominal spiracles (Fig. 21), sclerotized ring surrounding orifices; diameter 16 (14-21). Cluster of 71-92 multilocular disc pores (Fig. 2J) at apex of abdomen, total diameter 1 1 (10-12), diameter of locule ring 6 (6-6), width of tubular duct orifice 4 (4-4), 9 (7- 12) high. Locule ring with 10-13 loculi. Bi- locular tubular ducts in continuous series with, and similar to, dorsal bilocular tubular ducts. Four small discoidal pores (Fig. 2K), 4 (4-4) in diameter, midventrally between meso- and metathoracic legs. Three to 4 similar pores between pro- and mesotho- racic legs. Other stages. — First instars and cysts are Fig. 2. Matsucoccus oocarpae, n. sp., adult female. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON included in the dry and slide-mounted ma- terial. Both stages are typical of their re- spective stages of Matsucoccus. Type material. —Holotype adult female collected on Pinus oocarpa, Ostuncalco, Guatemala, July 10, 1944, by E. J. Ham- bleton. Three paratype adult females on 2 slides collected on P. oocarpa, Quezalte- nango, Guatemala, July 10, 1944, by E. J. Hambleton. The holotype and 1 paratype deposited in U.S. National Museum of Nat- ural History, Beltsville, Md.; one paratype had been retained in the Coccoidea collec- tion of Auburn University. KEY Ray and Williams (1984) provided a key to the North American species of Matsu- coccus which included 17 species. That key may be modified as follows to accommo- date the new species here described: 11. Marginal band of bilocular tubular ducts between last thoracic row and first abdom- inal row extending across abdominal dor- sum 11’. Marginal band of bilocular tubular ducts between last thoracic row and first abdom- inal row not extending across abdominal dorsum 12. Cicatrices in 4 bands, some specimens with slightly developed Sth band; feeding on P. QOUMIIHE! 5 asadenooa dows M. oocarpae, n. sp. 12’. Cicatrices in 5 or more bands; on P. pon- GerOSA 5.3 ee M. vexillorum Morrison Tubules of bilocular tubular ducts often ad- herent; feeding on pines of subsection Pon- LAR ONY? | deen Oey M. californicus Morrison Tubules of bilocular tubular ducts often di- vergent; not feeding on pines of subsection Ponderosae Cicatrices usually in 4 distinct bands, the anterior about '2 width of other bands; feeding on pines of subsection Australes . as SE Sie nts See M. alabamae Morrison Cicatrices in widely varying bands; feeding on P. banksiana of the subsection Contor- LAC <2... te ee M. banksianae, n. sp. 12a 12at: 12b 12b. 12b’. VOLUME 93, NUMBER 1 191 / «(Sam Pe / 1] Hai Be < = 8,e 8, 28 ,o @ .8 w/o@ sO Gs & 4 iA PLP 28, 29°86 epee 78 2 “ %G B.28, a" Bee a i Se ,47 rp A Teve “0 192 DISCUSSION Matsucoccus alabamae, M. banksianae, M. californicus, M. oocarpae and M. vex- illorum form a group of species which share the following characteristics. All 5 species lack fleshy sensory setae on antennal seg- ment 5, have legs and antennae well devel- oped, possess multilocular disc pores at the apex of the abdomen and possess only | size of body setae. Matsucoccus alabamae and M. californicus feed on the trunks of their hosts (personal observations) while M. vex- illorum feeds on twigs of its hosts (Mc- Kenzie 1943). All feed on pines of the sec- tion Pinus, each species feeding on pines of a single subsection of Pinus. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Cockerell, T. D. A. 1909. The Japanese Coccidae. Canadian Entomologist 41: 55-56. McKenzie, H. L. 1943. The seasonal history of Mat- sucoccus vexillorum Morrison (Homoptera: Coc- coidea: Margarodidae). Microentomology 8: 42- Spx Morrison, H. 1939. Descriptions of new species of Matsucoccus (Hemiptera: Coccidae). Proceedings of the Entomological Society of Washington 41: 1-20. Ray, C. H. and M. L. Williams. 1984. Two new spe- cies of Matsucoccus (Homoptera: Margarodidae) from Arizona and Mexico with a key to species in North America. Annals of the Entomological So- ciety of America 77: 765-769. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 193-196 NOMENCLATURAL CHANGES IN THE TREEHOPPER TRIBES HOPLOPHORIONINI, SMILIINI, AND TALIPEDINI (HOMOPTERA: MEMBRACIDAE) STUART H. MCKAMEY'! AND LEwis L. DEITz Department of Entomology, North Carolina State University, Raleigh, North Carolina 27695-7613. Abstract. —The following nomenclatural changes are made in the family Membracidae (Homoptera): Aconophoroides Fowler, 1894, is reinstated as a junior synonym of Potnia Stal, 1866; Microschema Stal, 1869, is considered to be a junior synonym of Potnia Stal, 1866, NEw Synonymy; Talipes Deitz, 1975, is considered to be a junior synonym of Trinarea Goding, 1926 (elevated to generic status), NEw SyNonymy. Also nine NEw COMBINATIONS are given: Metcalfiella spreta (Goding, 1893), Platycotis fuscata (Fowler, 1897), Potnia dubia (Fowler, 1894), Potnia orthosoma (Fonseca and Diringshofen, 1974), and Potnia straminicolor (Stal, 1862) (Membracinae: Hoplophorionin1); 7rinarea appen- diculata (Fonseca, 1936), Trinarea carinata (Funkhouser, 1922), and Trinarea fenestrata (Striimpel, 1974) (Membracinae: Talipedini); and Ophiderma gloveri (Goding, 1893) (Smi- liinae: Smiliini). A lectotype is designated for Platycotis spreta Goding, 1893. Key Words: While revising selected genera in the membracid tribe Hoplophorionini, we found several species misplaced at the ge- neric level, including three assigned to the wrong tribe. Some of the consequent trans- fers of species affect generic synonymy. Based on examination of type material (ex- cept as noted), nine new combinations are presented here under the higher categories to which the species are newly assigned. Subfamily Membracinae Tribe Hoplophorionini Metcalfiella Goding, 1929 Metcalfiella spreta (Goding, 1893), New COMBINATION ' Current address: Department of Ecology and Evo- lutionary Biology, Box U-43, University of Connect- icut, Storrs, Connecticut 06269-3043. Hexapoda, Insecta, generic synonymy, new combination A female specimen in the NMNH (Na- tional Museum of Natural History, Wash- ington, D.C.) is here designated lectotype of Platycotis spreta Goding. It bears the fol- lowing labels in which vertical lines (| ) are used to separate individual lines: ““FWGoding | Collection”’ and “*Platycotis | (Microschema) | spreta Godg. | Mex. (Ash) type”’ [underside with ‘“‘Hoplophora|Gdg. | Mex. (Ash) Type.”’] and “Type No. | 27388 |U.S.N.M.” [on red card] and “‘Platycotis| spreta | Godg. | Det.W.D.| Funkhouser” [on red-bordered card] and “LECTOTYPE | Platycotis | spreta | Goding, 1893 |L.L. Deitz 1989” [on red-bordered card supplied by Deitz]. A second specimen (without an ab- domen) from Goding’s collection (NMNH) is now labeled as a paralectotype. These specimens belong to the genus Metcalfiella Goding, 1926, based on the narrow head and the presence of many more pale setae 194 than pits on the pronotum (McKamey and Deitz 1991). Platycotis Stal, 1869 Platycotis fuscata (Fowler, 1897), New COMBINATION The lectotype of Ochropepla fuscata Fowler (BMNH: British Museum (Natural History), London, England), most recently placed in the genus Hoplophorion Kirkaldy, 1901 (Metcalf and Wade 1965), has a wide head and contrasting pale and dark bands across veins Sc and R on the forewing base, but lacks an r-m crossvein on the hind wing. This combination of features is diagnostic for the genus Platycotis Stal, 1869 (Mc- Kamey 1989). Potnia Stal, 1866 Potnia Stal, 1866 [Stal 1866b, not Stal 1866a as listed by Deitz (1975)] Aconophoroides Fowler, 1894, REINSTATED SYNONYMY Microschema Stal, 1869, NEw SYNONYMY Potnia dubia (Fowler, 1894), NEw Comst- NATION Potnia orthosoma (Fonseca and Diringsho- fen, 1974), NEw COMBINATION Potnia straminicolor (Stal, 1862), NEw COMBINATION The type species of Potnia Stal is Mem- bracis venosa Germar, 1821. Based on ex- amination of two probable syntypes of M. venosa, one from MLUH (Martin-Luther- Universitat Halle, Halle, German Demo- cratic Republic) and one from ZMUH (Zoologisches Museum, Universitat von Hamburg, Hamburg, Federal Republic of Germany), in conjunction with a review of generic limits in the tribe Hoplophorionini (McKamey 1989), the following combina- tion of features appears to be diagnostic for Potnia: head narrow; pronotum extending posteriorly nearly to apex of forewing in re- pose; hind wing with r-m crossvein and with cell R; (the cell posterior to vein R, of Deitz PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1975) longer than cell R;; metathoracic tibia with cucullate setae in row I (sensu Deitz 1975). Prior to the work by Fonseca and Di- ringshofen (1974), the genus Aconopho- roides (type species: Thelia gladiator Walk- er, 1857) was considered to be a junior synonym of Potnia (Goding 1929, Metcalf and Wade 1965). Based on examination of the holotype of 7. gladiator Walker (BMNH), we confirm this synonymy. Although the type material of Aconopho- roides orthosoma Fonseca and Diringshofen was not examined, the original description and illustrations (1974) indicate member- ship in the genus Potnia. Likewise, exami- nations of the lectotype of Ochropepla dubia Fowler (BMNH), a species formerly placed in Hoplophorion (Metcalf and Wade 1965), and of the holotype of Hoplophora stra- minicolor Stal (NHRS: Naturhistoriska Riksmuseet, Stockholm, Sweden), the type species of Microschema Stal, led to the con- clusion that these two species are also con- geners of Potnia venosa. Thus, we list above three new combinations and the new generic synonymy of Microschema, which was pre- viously considered a subgenus of Platycotis. The only other species formerly included in Microschema, Platycotis spreta, is trans- ferred to Metcalfiella, above. Hoplophora straminicolor was described from Brazil. Such a distribution for Platy- cotis would involve a geographic disjunc- tion and a host shift, because the families Fagaceae (including Quercus spp.) and Bet- ulaceae, which encompass all known hosts of Platycotis, do not occur in Brazil (Hey- wood 1985). This information alone sug- gested that the species was placed incor- rectly, and also suggests that a similar taxonomic error may be involved with the Brazilian species Platycotis sordida (Ger- mar, 1821), type material of which was not examined. VOLUME 93, NUMBER 1 Subfamily Membracinae Tribe Talipedini Trinarea Goding, 1926 Trinarea Goding, 1926, elevated to generic status, NEw STATUS Talipes Deitz, 1975, to Trinarea Goding, 1926, NEw SYNONYMY Trinarea appendiculata (Fonseca, 1936), New ComMBINATION Trinarea carinata (Funkhouser, 1922), NEw COMBINATION Trinarea fenestrata (Striimpel, 1974), NEw COMBINATION Trinarea was formerly placed as a sub- genus of Hoplophorion (Metcalf and Wade 1965). Although Metcalf and Wade (1965: 578) attributed the designation of a type species to Goding (1929), Goding (1926:305) wrote “‘type” before the name carinata in his key to subgenera. Thus, Ochropepla car- inata Funkhouser is the type of Trinarea by original designation (International Com- mission on Zoological Nomenclature 1985: Articles 68b, 67c). The holotype of O. car- inata (NMNH) and the holotype and para- types of Tylopelta fenestrata Strimpel (ZMUH) were examined and found to be congeneric with TJalipes appendiculata (Fonseca, 1936), the type species of Talipes Deitz, 1975. Members of the genus Jalipes differ from other membracids in having fo- liaceous pro- and mesothoracic tibiae but metathoracic legs with clavate tibiae and reduced tarsi (Deitz 1975). Trinarea is therefore given generic status, with 7alipes as its junior synonym, but the tribal name Talipedini Deitz, 1975, is retained (Inter- national Commission on Zoological No- menclature 1985: Article 40a). Subfamily Smiliinae Tribe Smiliini Ophiderma Fairmaire, 1846 Ophiderma gloveri (Goding, 1893), NEw COMBINATION 195 Goding (1893) described Hoplophora gloveri by reference to an illustration by Glover (1878), and no specimen could be located at either AMNH (American Mu- seum of Natural History, New York) or NMNH, known depositories of Goding’s type specimens. H. gloveri was placed most recently (Metcalf and Wade 1965) as a jun- ior synonym of Hoplophorion monogram- ma (Germar, 1835). Although the illustrat- ed specimen was identified in Glover’s figure legend as “Hoplophora n.s.,” its forewing has a preapical confluence of the radial and medial veins, which is characteristic of the Smiliinae and absent from Hoplophorioni- ni. Forewing venation and pronotal color- ation and contour, in lateral view, indicate the specimen illustrated is best referred to the genus Ophiderma Fairmaire (Smiliinae: Smiliini). The precise identity of H. gloveri is likely to remain uncertain in the absence of the original material on which Glover’s figure was based. ACKNOWLEDGMENTS We thank the following people and insti- tutions for loaning specimens and providing other assistance: M. Schwartz (AMNH); P. S. Broomfield, W. J. Knight, and M. D. Webb (BMNH); M. Dorn and J. Schuh (MLUH); P. Lindskog (NHRS); T. J. Henry, J. P. Kramer, and M. B. Stoetzel (U.S. Dept. Agric., % NMNH); R. C. Froeschner and G. H. Hevel (NMNH); and H. Striimpel (ZMUH). We also thank C. H. Dietrich, M. H. Farrier, J. W. Hardin, and H. H. Neunzig (all of North Carolina State University) who provided many helpful suggestions relating to this work. This is paper No. 12521 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, N.C. 27695- 7643; an earlier version formed part of a thesis submitted by S. H. McKamey in par- tial fulfillment of the requirements for the M.S. degree in entomology, North Carolina State University at Raleigh. Funding for 196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON McKamey was provided by the NCSU De- partment of Entomology and Alumni As- sociation during the year 1986-1987 and by a National Science Foundation Graduate Fellowship during the period 1987-1989. LITERATURE CITED Deitz, L. L. 1975. Classification of the higher cate- gories of the New World treehoppers (Homoptera: Membracidae). North Carolina Agricultural Ex- periment Station Technical Bulletin 225. [iv] + 177 pp. Fonseca, J. P. da. 1936. Contribuigao para o con- hecimento dos membracideos neotropicos. Ar- chivos [sic] do Instituto Biologico (Sao Paulo) 7: 157-166. Fonseca, J. P. da and R. von Diringshofen. 1974. Contribuicao ao conhecimento dos membracideos neotropicos (Homoptera: Membracidae, VII). Ar- quivos do Instituto Bioldgico (Sao Paulo) 41(4): 151-160. Fowler, W. W. 1894. Order Rhynchota. Suborder Hemiptera-Homoptera [continued]. Biologia Cen- trali-Americana 2: 25-56, plates 3-4. 1897. Order Rhynchota. Suborder Hemip- tera-Homoptera [continued]. Biologia Centrali- Americana 2: 169-184, plate 11. Funkhouser, W. D. 1922. New records and species of South American Membracidae. Journal of the New York Entomological Society 30(1): 1-35, plates 1-3. Glover, T. 1878. [Manuscript notes from my journal and illustrations of insects native and foreign. Or- der Hemiptera: suborder Homoptera. Washing- ton, D.C.]. Plates 1-2 + 5 pp. [apparently distrib- uted in very limited numbers; title above as cited by Metcalf and Wade’s (1963) “‘A Bibliography of the Membracoidea and Fossil Homoptera (Ho- moptera: Auchenorhyncha),” North Carolina State [University], Raleigh, iv + 200 pp.] Goding, F. W. 1893. Bibliographical and synonym- ical catalogue of the described Membracidae of North America. Illinois State Laboratory of Nat- ural History Bulletin 3: 391-482. 1926. Classification of the Membracidae of America. Journal of the New York Entomological Society 34(4): 295-317. 1929. The Membracidae of South America and the Antilles. 1V. Subfamilies Hoplophorioni- nae, Darninae, Smiliinae, Tragopinae (Homop- tera). Transactions of the American Entomologi- cal Society 55: 197-330, plates 10-11. Heywood, V. H. (ed.). 1985. Flowering Plants of the World. 1985. Prentice-Hall Inc., New Jersey. 335 pp. International Commission on Zoological Nomencla- ture. 1985. International Code of Zoological No- menclature. 3rd ed. International Trust for Zoo- logical Nomenclature, London, University of California Press, Berkeley. xx + 338 pp. McKamey, S. H. 1989. Revision of the Genus Met- calfiella (Homoptera: Membracidae) with Review of the Nomenclature and Systematics of the Tribe Hoplophorionini and Description of a New Ge- nus. Unpublished M.S. thesis, Department of En- tomology, North Carolina State University, Ra- leigh. vili + 315 pp. McKamey, S. H. and L. L. Deitz. 1991. Synonymy in the treehopper genera Hoplophorion, Metcal- fiella, and Ochropepla (Homoptera: Membraci- dae). Proceedings of the Entomological Society of Washington 93: 159-162. Metcalf, Z. P.and V. Wade. 1965. General Catalogue of the Homoptera. A Supplement to Fascicle I— Membracidae of the General Catalogue of Hemip- tera. Membracoidea. In two sections. North Car- olina State University, Raleigh. vi + 1552 pp. Stal, C. 1862. Bidrag till Rio Janeiro-traktens Hemip- ter-fauna. II. Kongliga [Svenska] Vetenskaps-Aka- demiens Handlingar 3(6): 1-75. 1866a. Hemiptera Homoptera Latr. Hemip- tera Africana 4. Norstedt, Stockholm. 276 pp. + 1 plate. 1866b. Analecta hemipterologica [contin- ued]. Berliner Entomologische Zeitschrift 10: 38 1- 394. 1869. Hemiptera Fabriciana. Fabricianska Hemipterarter, efter de i K6pehnhamn och Kiel forvarade typexemplaren granskade och beskrifne. 2. Kongliga Svenska Vetenskaps-Akademiens Handlingar 8(1): 1-130. Strimpel, H. 1974. Beitrag zur Kenntnis der neotro- pischen Membraciden Gattung 7y/lope/ta Fowler, 1894. Entomologische Mitteilungen Zoologisches Museum, Hamburg 4(88): 531-540. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 197-200 A DISTRIBUTION AND ESTABLISHMENT PROGRAM FOR CHILOCORUS KUWANAE (SILVESTRI) (COLEOPTERA: COCCINELLIDAE) IN THE UNITED STATES R. M. HENDRICKSON, JR., J. J. DREA, AND MIKE ROSE (RMH) Beneficial Insects Research Laboratory, Agricultural Research Service, U.S. Department of Agriculture, 501 South Chapel Street, Newark, Delaware 19713; (JJD) Beneficial Insects Laboratory, BBII, Agricultural Research Service, U.S. Department of Agriculture, BARC-East, Beltsville, Maryland 20705; (MR) Entomology/Biological Con- trol, Texas A&M University, College Station, Texas 77843. Abstract. —Chilocorus kuwanae, a coccinellid predator of euonymus scale, Unaspis eu- onymi (Comstock) (Homoptera: Diaspididae), was introduced from Korea (1984) and Japan (1985) into the United States. The beetle was released in 25 states in the eastern half of the country and became established in nine states (CN, DE, MA, MD, MI, NC, NJ, OH, PA) and the District of Columbia. Key Words: Insecta, Coccinellidae, Chilocorus kuwanae, euonymus scale, Unaspis eu- onymi, biological control, predator, Aprostocetus neglectus, Eaonymus plants Euonymus scale, Unaspis euonymi (Com- stock) (Homoptera: Diaspididae), is a seri- ous pest of many species of ornamental trees and shrubs in the United States (Gill et al. 1982). In the absence of adequate controls, this insect can attain populations severe enough to cause complete defoliation and death of the host plant (Johnson and Lyon 1988). Even moderate infestations impair photosynthesis and reduce growth (Cock- field and Potter 1987). The pest is a nui- sance to homeowners because insect-dam- aged shrubs are unsightly and require pesticide treatments or removal. Some at- tractive but scale-susceptible species or va- rieties of Euonymus, an ornamental shrub or tree, are no longer produced by nursery- men since the plants require frequent chem- ical pest control (Drea and Hendrickson 1988). A biological control method for eu- onymus scale could reopen markets for these Euonymus. The Agricultural Research Service (ARS), USDA, initiated a Small Farms Research Project in the early 1980’s for control of scale pests, one of which was euonymus scale. Since U. euonymi is believed to be Oriental in origin, the assistance of the ARS Asian Parasite Laboratory, Seoul, Republic of Korea (ROK) was requested to obtain natural enemies attacking the scale in Ko- rea. Several species of parasites and pred- ators of U. euonymi were collected by the laboratory personnel, including the cocci- nellid Chilocorus kuwanae (Silvestri). Spec- imens of this beetle were shipped to the ARS quarantine facilities at the Beneficial Insects Research Laboratory, Newark, Delaware. After clearance from quarantine, the beetle was sent to the ARS Beneficial Insects Lab- oratory, Beltsville, Maryland, for additional studies, culture, and release (Drea and Carl- son 1987). METHODS AND MATERIALS From 1984 to 1986 411 adult C. kuwanae were received from the laboratory in Korea. 198 Initially these specimens were used to es- tablish laboratory cultures. F, adults, lar- vae, and a small number of eggs from these cultures were released at the U.S. National Arboretum, Washington, D.C., on Euony- mus europaeus L. in 1984. Color photo- graphs of the predators can be found in Hen- drickson and Drea (1988). The predator became established at the Arboretum. By 1985 this primary release site had become a natural insectary and served as the main source of living material for laboratory cultures and for subsequent distribution of the predator to secondary release sites in MD, DE, PA, NJ and DC. Populations of beetles at these locations increased rapidly. For example, in Swarth- more, PA, a single shrub, ca. 2.5 m diam.., provided 7135 beetles in the year following the initial release of 84 C. kuwanae adults on the shrub. During the period 1985 to 1989 a total of 16,157 beetles were collected and redistributed from the primary and sec- ondary sites. Ultimately, beetles were re- leased directly or sent to cooperators in 24 states (AL, AR, CN, DE, FL, GA, KS, KY, MA, MD, MI, MO, NC, NH, NY, OH, OK, PA, VA, RI, TN, TX, VT, WV) and DC. In 1986, nine specimens of C. kuwanae, col- lected by one of us (MR) in Japan, were cultured at Beltsville and 53 specimens were supplied to the New Jersey Department of Agriculture, Trenton. To determine the success of the project by 1989, questionnaires requesting infor- mation relating to the outcome of releases made at various locations throughout much of the eastern United States, were mailed to cooperators in June and reported in this study. RESULTS AND DISCUSSION By 1987, the scale populations had been reduced to a very low level at the Arbore- tum, the primary release site (Drea and Carlson 1987). By 1989 the scale insects were almost absent from the trees and the coccinellid was difficult to find. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON All cooperators replied to the question- naire. From their replies and personal field observations we determined that C. kuwa- nae was established in CN, DC, DE, MA, MD, MI, NC, NJ, OH, and PA for one or more years (Fig. 1). In some areas, insufh- cient time had elapsed since release to de- termine winter survival. Releases made in Texas failed to estab- lish. One of us (MR) obtained 2311 adult, larval and pupal stages of C. kuwanae from Maryland and Delaware sites in 21 con- signments from August 1987 to September 1989. These were released on euonymus scale at numerous sites in Austin, Bryan, College Station, and Dallas/Ft. Worth. Only a few cast larval skins were found and by July 1990 only one adult was recovered. Release sites in Texas were ant-free and C. kuwanae adults readily fed on the abundant scale insects. Chilocorus kuwanae adults were observed to feed and remain alive for 10 days in branch sleeve cages following colonization. Failure to establish may be related to climatic conditions, especially heat. Surveys are now being conducted to determine overwinter survivalship and re- production of adult C. kuwanae released in October, 1989. The release date was select- ed to initially circumvent high summer temperatures and to study the potential col- onizations of the beetle at lower tempera- ture. An example suggesting the importance of heat in preventing establishment of this predator occurred in Delaware. Five releas- es of C. kuwanae totalling 60 individuals were made in 1986 and 1987 on 22 small specimens (less than 30 cm high) of a pros- trate, variegated variety of Euonymus for- tunei, infested with euonymus scale, grow- ing on the grounds of the Beneficial Insects Research Laboratory, Newark, DE. The beetles never remained on the plants more than 2 or 3 days and failed to establish. These plants were individually isolated and surrounded by bare soil. Heat rising from the exposed soil on sunny days may have VOLUME 93, NUMBER 1 199 Fig. 1. an open circle represents one or more releases without confirmed establishment in that state. repelled the predators. In contrast, beetles became established on an extensive ground- cover planting of the same species at Swarthmore, PA, that had no bare soil to reflect heat. The presence of ants may be another con- sideration when selecting release sites for the coccinellid. At some locations in DE, MD, and PA, the beetles failed to establish, or if established, failed to develop popula- tions high enough to control the scale over a several year period. These shrubs or trees were always ant infested. There were nu- merous ants tending honeydew producers such as aphids and planthoppers. The ants repeatedly stung larvae and pupae of C. ku- wanae and removed stung individuals from the shrubs. Ant species on shrubs were iden- tified as Camponotus nearcticus Emery, C. ferrugineus (F.), Formica pallidefulva La- treille, Lasius alienus (Foerster) and Tapi- Distribution of Chilocorus kuwanae in eastern United States. A solid circle indicates establishment, noma Sessile (Say) (Hymenoptera: Formic- idae). Larvae of C. kuwanae, collected from study sites in Delaware and Maryland dur- ing 1988 and 1989, were parasitized by Aprostocetus neglectus (Domenichin1) (Hy- menoptera: Eulophidae: Tetrastichinae). The identifier, John LaSalle, noted that this species is known from Europe, the Middle East and North Africa where it is a parasite of coccinellid larvae and pupae. The para- site was previously unknown from North America. No parasites have been reared from adults of C. kuwanae originating from field sites in the U.S. To eliminate the possibility of further distributing A. neglectus, only adult C. kuwanae are utilized for new field colo- nizations. Our best estimate of the rate of move- ment by C. kuwanae was determined from 200 adults reared from pupae collected by R. D. Gordon in Bowie, MD in August, 1989. Specimens probably dispersed from our closest colonization sites at either Beltsville Agricultural Research Center, Maryland, a distance of ca. 11 km in 5 years, or from the U.S. National Arboretum, Washington, DC, a distance of ca. 19 km in 6 years. It appears that the rate of movement is about 2-3 km per year. The use of cooperators for disseminating the beetle has greatly enhanced the slow rate of natural dispersal. As a result of this effort, C. kuwanae has become established in nine states and the District of Columbia during the six year period from 1984 to 1989. We are hopeful that additional cooperator as- sistance and future involvement of the An- imal and Plant Health Inspection Service, USDA, will result in the collection and wide dissemination of the predator throughout thesW:s: ACKNOWLEDGMENTS We thank Ho-Yeon Han and Jang-Hoon Lee, Asian Parasite Laboratory, USDA, ARS, Seoul, Republic of Korea who sup- plied the original material; and S. E. Barth, Beneficial Insects Research Laboratory, Ag- ricultural Research Service, USDA, New- ark, DE and E. McClunin, Beneficial Insects Laboratory, ARS, USDA, Beltsville, MD, who assisted in collecting, culturing, and re- leasing Chilocorus. Identifications were pro- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON vided by R. D. Gordon (Coccinellidae), D. R. Smith (Formicidae), both Systematic En- tomology Laboratory, Agricultural Re- search Service, USDA, Washington, DC, and J. LaSalle (Eulophidae), C.A.B. Insti- tute of Entomology, London, United King- dom. The manuscript was reviewed by R. D. Gordon, and D. E. Meyerdirk, Animal and Plant Health Inspection Service, USDA, Hyattsville, MD. LITERATURE CITED Cockfield, S. D. and D. A. Potter. 1987. Distribution development, and feeding impact of euonymus scales (Homoptera: Diaspididae) on Euonymus fortunei under greenhouse conditions. Environ- mental Entomology 16: 917-921. Drea, J. J. and R. W. Carlson. 1987. The establish- ment of Chilocorus kuwanae (Coleoptera: Cocci- nellidae) in eastern United States. Proceedings of the Entomological Society of Washington 89: 821- 824. Drea, J. J. and R. M. Hendrickson, Jr. 1988. Exotic predators. American Nurseryman 168(8): 66-71. Gill, S. A., D. R. Miller, and J. A. Davidson. 1982. Bionomics and taxonomy of the euonymus scale, Unaspis euonymi (Comstock), and detailed bio- logical information on the scale in Maryland. Uni- versity of Maryland Agricultural Experiment Sta- tion Miscellaneous Publications Number 969. 36 pp. Hendrickson, R. M., Jr. and J. J. Drea. 1988. Our insect allies: Beetles battle scale. Explorer 30(4): 4-8. Johnson, W. T. and H. H. Lyon. 1988. Euonymus scales, pp. 388-389. Jn Insects that Feed on Trees and Shrubs. 2nd ed. Cornell University Press, Ith- aca, NY. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 201-203 NOTE On Intraspecific Variation in the Alysiinae (Hymenoptera: Braconidae) There are few studies published that ex- amine intraspecific variation of the aly- siines. Thus, Wharton (1980), in his review of the neartic alysiines, states: ““A general disregard for intraspecific variation has characterized much of the recent taxonomic work on the Alysiini. This has not only cre- ated a misleading impression of diversity and biogeographic relationships, but also resulted in problems in identification.’ He then continues with a detailed analysis of this problem and concludes with: “The studies on intraspecific variation suggest that new species should not be described on the basis of minor morphological differences unless these are supported by biological in- formation.” It is therefore interesting to make an in-depth study of the intraspecific variability of the taxonomic characters of morphological type, such as the absolute or relative measurements of certain parts of the body, in order to elucidate to what ex- tent these data are of diagnostic value. The present article addresses this question. For this study, the Dacnusini species Dac- nusa sibirica Telenga, 1934, was selected Table I. because it is common in the study area and because it is easy to breed in the laboratory. Materials and Methods. — Measurements were made of 195 specimens (75 6 and 120 2) of Dacnusa sibirica, obtained from three species of Agromyzids [Chromatomyia hor- ticola (Goureau, 1851); Liriomyza sonchi Rydén, 1951 and Liriomyza strigata (Mei- gen, 1830)], on four species of plants [Car- daria draba L. (Desv.), Diplotaxis erucoides (L.) DC.; Eryngium maritimum L. and Pa- paver rhoeas (L.)], collected at four localities [(Lérida: Viella (31TCH1930); Teruel: Al- cala de la Selva (30TXK9472); Valencia: Albalat de la Ribera (30SYJ2876), Cullera (30SYJ3738)]. The characters treated (= measurement magnitudes; measurement characters) cor- respond with those most commonly used— see Griffiths (1964)—such as: width and length of the head (AC, LC), length and width of thorax (LT, AT), and length and width of the petiole (LP, AP), which are used for calculating the indices: AC/LC, LT/AT and LP/AP. The simple allometric equation (y = a.x>) was used used for the study of the Allometric equation: AC = a.LC? (see graph 1) Sex a b r Males 1.312 05332 0.671 Females 0.269 0.761 0.652 Allometric equation: LT = a.AT® (see graph 2) Sex a b 1; Males 0.456 0.854* 0.819 Females —1.143 1.220 0.892 Allometric equation: LP = a.AP (see graph 3) Sex a b it Males 0.377 0.819* 0.908 Females 0.058 0.907* 0.935 F g.l. Fe g.1 39.243 173 2 D 2.805 1,118 ore oty FE g.l. leh g.l 6.596 7g) ? 2 9) 2.849 1,118 fe: aa Je all E’ g.1 He ni 81.251 2,191 13.290 202 joint variation of the magnitudes consid- ered. The measurements were taken using a mi- crometric eyepiece with a margin of error of 0.0072 mm, fitted to an Olympus szh binocular microscope. Results.—Table | gives the results of the bivariate analysis of the magnitudes AC, LC, LT, AT, LP and AP, involved in the cal- culation of the indices AC/LC, LT/AT and LP/AP. Allometric equations for the three pairs of variables involved in calculating the in- dices: AL/LC, AT/LT and LP/AP (r being the correlation coefficient; F being the sta- tistic (= statiscal parameter) distributed as a Snedecor F variable, which allows one to distinguish between allometeric (b = 1) and isometric growth (b = 1); F’ being the sta- tistic (statiscal parameter) distributed as a Snedecor F variable, which allows the sig- nificant acceptance or rejection of the su- perposition of two given regression lines (De Renzi and Martinelli 1979). In four of the equations obtained (marked with an asterisk) the allometric coefficient (b) is significantly higher or lower than | (F > FO0.025). The hypothesis of the superposition of the regression lines, for each sex, can be rejected in all cases, with an optimum level of sig- nificance, x = 0.01, F’ > FO.001 (2, n + n’ — A) Discussion.—In taxonomic studies of members of the Braconidae, as in other groups from the series Parasitica, ratios be- tween measurements are frequently used as diagnostic characters. For some time now, the authors have ob- served certain individual, interpopulational and even intrapopulational variation in cer- tain measurements and we thus think that the joint variation of these measurements does not allow a specific value to be given to the respective ratios. The results obtained confirm secondary sexual characteristics 1n the relative increase of corporal measurements, in the three cor- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON lin! RIG = 02/6394. (OG Uns A\G Females Win UGS Se Byte ee 5 .3/5)9) = 0.67074 graph 1 1 A= 10/.74:5i635: ee 0 Bie Tiny OT females Lin, “A= soy P1429 > erate? iin, Tey r= 0.89239 graph 2 In AP= 0.0578 + 0.9067 In LP r= 93503 A males In AP= 0.3367 + 0.8187 In LP C= 0.90754 graph 3 poral indices examined (AC/LC, AT/LT and LP/AP). The equations obtained give allo- metric coefficients (b) higher or lower than 1, thus producing an increase or decrease in the value of the indices in accordance with the corporal measurements. These results are unfavourable for the use of indices of measurements in either keys or as basic diagnostic characters. VOLUME 93, NUMBER 1 Acknowledgments.—We wish to thank Dr. Griffiths (University of Alberta. De- partment of Entomology), Dr. Godfray (Im- perial College, London) and Dr. Wharton (Texas A&M University, Department of Entomology) for their observations and critical reading of the manuscript. LITERATURE CITED De Renzi, M. and J. Martinelli 1979. Algunos as- pectos de la problematica de la especie paleonto- logica: Aplicacion al caso de la diferenciacion biométrica de Nassarius semistriatus (Brocchi, 1814) y N. elatus (Gould, 1845). Studia Geologica, XV: 7-36. 203 Griffiths, G.C. D. 1964. The Alysiinae (Hym., Bra- conidae) parasites of the Agromyzidae (Diptera). I. General questions of taxonomy, biology and evolution. Beitr. Entomol., 14: 823-914. Wharton, R. 1980. Review of the Neartic Alysiini (Hym., Braconidae), with discussion of generic re- lationships within the tribe. Univ. Calif. Publ. Ent., 88: 112 pp. J. Tormos and S. F. Gayubo, Departa- mento de Biologia Animal, Ecologia, Eda- fologia, Parasitologia y Quimica Agricola, Facultad de Biologia, Universidad de Sal- amanca, 3707 1-Salamanca, Espana; J. D. Asis and A. Sendra, Fundaci6n Entomol6- gica “Juan de Torres Sala,” Passeig de la Petxina, n° 15, 46008-Valéncia, Espania. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 204-205 NOTE A Simple Device to Clean Insect Specimens for Museums and Scanning Electron Microscopy Insect specimens are often contaminated with debris that obscures structures of in- terest. Scales and hairs adhere to specimens taken from malaise traps or by other col- lecting methods that capture Lepidoptera with other insects. Dust accumulates on in- sects both in life and as mounted specimens in collections. When mounting insects for scanning electron microscopy, fragments of dissected structures often adhere to the specimen. Below I described a simple, in- expensive device that uses a fine jet of air to efficiently remove loose debris without damaging specimens. The device (Fig. 1) consists of a one liter filtering flask, a one-hole rubber stopper (no. 8), latex rubber tubing (6.4 mm I.D., ca. 80 cm long), plastic tubing (6.4 mm I.D., ca. 70 cm long), glass tubing (8 mm O.D., ca. 28 cm long), a small quantity of polyester aquarium filter fiber, and a modified dis- posable glass Pasteur pipet (14.6 cm long). Latex tubing facilitates manipulation of the pipet; plastic tubing is less likely to collapse and block air flow to the device. The poly- ester aquarium filter fiber removes moisture and other contaminants from the air supply. The pipet (Fig. 2) is modified as follows: Reduce the diameter of the hole in the tip of the pipet to about 0.25-0.5 mm by pass- ing it through the flame ofa Bunsen or Fish- er burner. For safety, use protective eye wear, and before making the vent hole attach a length of tubing to the pipet to keep your face away from the flame. Make the vent hole by holding one side of the shoulder of the pipet in the flame. When the glass begins to glow, blow through the tubing attached to the pipet until a bubble begins to swell from the area being heated. Continue blow- ing until the bubble bursts, thus forming the vent hole. Polish the edge of the vent hole 1 plastic ate in a glass tubing latex ‘ z Filtering eae flask filter oe ey i Cr; Figs. 1-3. Device for cleaning insect specimens. |, Complete device. 2, Modified pipet. 3, Holding the pipet. VOLUME 93, NUMBER i by passing it through the flame. The final diameter of the vent hole should be about 3-4 mm. Finally, bend the pipet near the tip by passing it through the flame. A low flame works well for modifying the tip of the pipet; a hotter flame is best for making the vent hole. Assemble the device as shown in Fig. 1 and attach the plastic hose to a source of compressed air. In many laboratories com- pressed air is available from stopcocks built into the benches. Air passing through the tip of the pipet is used to clean specimens. Use very low vol- umes of air from the compressed air source. The optimal volume depends on the di- ameter of the hole in the tip of the pipet. The air passing out of the vent hole should be barely detectable for pipets with tips re- duced to 0.25-0.5 mm. Air is forced through the tip of the pipet by covering the vent hole with your index finger (Fig. 3). Because the hole in the tip 1s much smaller than the vent hole, air pres- sure increases in the flask when the vent hole is covered. Simultaneously, the velocity of the air leaving the tip of the pipet increases. When cleaning specimens, use short bursts of air by opening and closing the vent hole at regular intervals, or by only partially cov- ering the vent hole. This will prevent the stopper from being blown out of the flask and will also keep the velocity of the jet from 205 becoming great enough to damage speci- mens. Loose debris is blown off of a spec- imen by directing the jet toward the area to be cleaned. This can be done under a dis- secting microscope. Debris that is glued to a specimen by oil or other such substances usually cannot be removed. Because of the low volume of air used, it is unlikely that pressure within the flask will become great enough to break the flask. Moreover, the stopper, if not placed in the flask with excessive force, acts as a safety valve. However, as an added precaution the flask should be wrapped with nylon-rein- forced tape. Additionally, by reversing the direction of air flow through the device, it can be used to pick up and manipulate small delicate objects. For this purpose, install the device onto a vacuum line. John W. Heckman, Center for Electron Optics, Michigan State University, East Lansing, suggested the method for making a vent hole in the pipet. An earlier draft of the manuscript was reviewed by Guy L. Bush, Judith M. Sirota and Frederick W. Stehr, Department of Entomology, Michi- gan State University, East Lansing, and an anonymous reviewer. Their assistance is gratefully acknowledged. John Jenkins, Department of Zoology, Natural Science Building, Michigan State University, East Lansing, Michigan 488 24- JVB PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 206-207 NOTE Lectotype Designation for Opostega heringella Mariani, a synonym of Opostega spatulella Herrich-Schaffer (Lepidoptera: Opostegidae) Because an examination of the type series of Opostega heringella Mariani was not pos- sible until after my synoptic catalogue of the family Opostegidae (Davis, 1989) had been accepted for publication, I wish at this time to report on the results of that study. Mariani (1937) described Opostega her- ingella (Fig. 1) from an unspecified number of specimens collected during July in Zap- pulla, Sicily. He considered the species to be intermediate in color and maculation be- tween O. constantiniella Costantini and O. spatulella Herrich-Schaffer. Through the cooperation of Marcello Arnone of the Mu- seo Civico di Terrasini, I have been able to examine the entire series of eight male and six female syntypes of O. heringella and have selected as lectotype a male bearing the fol- lowing labels: Sicilia, Zappulla, 5.VI.34, M. Mariani; 358; Opogona spatulella H. S., Rbl. & Htg. det.; photograph on file USNM; Lec- totype 6, Opostega heringella Mar., by D. Davis. The remaining 13 paralectotypes were collected at the same type locality and bear the following label data: 1 2, 20.VI.31; 1 4, 20.VI.33, M. Mariani; 1 9, 2.VI.34, M. Mariani; 2 6 (6 slide USNM 30498), 3 9, 5.VI.34, M. Mariani; 4 4, 1 2, 12.VI.34. Ex- amination of the male genitalia of one of the paralectotypes has verified the earlier, unpublished determination by Rebel and Hartig as well as the recently published de- cision by van Nieukerken (1989) that her- ingella is a junior synonym of spatulella. The ground color and forewing markings of Opostega spatulella are highly variable, as shown in two extremes previously figured (Davis, 1989, Figs. 262 and 263). Opostega costantiniella was also synonymized under spatulella by van Nieukerken but, as noted Fig. 1. Opostega heringella Mariani, lectotype. VOLUME 93, NUMBER 1 by him, the unique female holotype of the former is believed lost (formerly in the Tu- rati collection). I am indebted to Marcello Arnone and the Museo Civico di Terrasini, Terrasini, Italy for the loan of the type specimens. I also wish to thank Erik J. van Nieukerken of the Rijksmuseum van Natuurliyke His- torie, Leiden, Netherlands and Giorgio Baldizzone of Asti, Italy for assisting me in locating the type depository of O. heringel- la. LITERATURE CITED Davis, D. R. 1989. Generic revision of the Oposteg- idae, with a synoptic catalog of the world’s species 207 (Lepidoptera: Nepticuloidea). Smithsonian Con- tributions to Zoology, No. 478. 97 pp., 320 figs., 1 map. Mariani, M. 1937. Nuove species e forme di Lepi- dotten di Sicilia ed un nuovo parassita degli agru- mi. Giornale di Scienze Naturali ed Economiche 39(3):1-13, 1 pl. Nieukerken, E. J. van. 1989. Opostegidae, pp. 357- 375. In R. Johansson, E. S. Nielsen, E. J. van Nieukerken, and B. Gustafsson, eds., The Nepti- culidae and Opostegidae (Lepidoptera) of North- west Europe. Fauna entomologica Scandinavica, vol. 23, part 1. Donald R. Davis, Department of Ento- mology, NHB 127, Smithsonian Institution, Washington, D.C. 20560. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 208-209 NOTE Pseudomethoca simillima (Smith) (Hymenoptera: Mutillidae), a Newly Discovered Parasitoid of Andrena macra Mitchell (Hymenoptera: Andrenidae) Pseudomethoca simillima (Smith) are known from Massachusetts southward to Florida, westward to Nebraska and Texas (Krombein. 1979. Jn Catalog of Hymenop- tera in America North of Mexico. Vol. II. pp. 1276-1314). Investigations on the bi- ology of this species have dealt with sea- sonal abundance of adults (Fattig. 1943. Bull. Emory Univ. Mus. no. 1. 24 pp.; Schmidt and Hook. 1979. Florida Entomol. 62(2): 152) and mating behavior (Shappirio. 1947. Bull. Brooklyn Entomol. Soc. 42: 162- 163, Sci. Monthly. 44(4): 348-350). This note reports the first host record for P. simil- lima. The study areas were two Andrena macra Mitchell nesting sites located on the United States Marine Corps Reservation in Quan- tico, Virginia. Andrena macra built their nests in the soil and each female constructed and provisioned its own brood cells. Ob- servations were made during Spring and Summer 1984 and 1985 at A. macra nest- ing sites I and II. On 26 April 1984 P. simillima females were visible and extremely numerous on the surface of A. macra nesting site I. They were crawling over and under leaf litter. Twenty- seven females were trapped within 15 min- utes. However, A. macra males or females were not seen and emergence holes or nest entrances were not present on the surface of site I. Schmidt and Hook (1979) observed an unusually large population of P. simil- lima in Highlands Co., Florida on 20 March 1978. Males were not seen and the nests of the potential hosts were not found (Schmidt and Hook 1979). Andrena macra males and females were first observed on the surface of the nesting sites by 10 May 1984. Pseudomethoca si- millima females were also present on this day, but much less numerous as compared to their numbers seen on 26 April 1984. Pseudomethoca simillima females were seen at sites I and II during Spring and Summer 1985. For example, on 22 June 1985, al- most 3 weeks after the A. macra nesting season was over, two females were found 35 cm beneath the surface of nesting site II, some 2—4 cm away from brood cells that contained healthy A. macra prepupae. Oth- er neighboring brood cells were excavated from A. macra nests on this day and also on 29 June 1985. Forty-seven cells were ex- cavated and each contained a prepupa. Three A. macra and one Nomada sp. prepupae were paralyzed and a parasitoid egg was at- tached to their cuticle. All prepupae were placed, individually, inside 4-dram shell vi- als and taken to the laboratory. Prepupae were reared at room temperature (24—27 de- grees Celsius). The successfully reared parasitoids were P. simillima, a male and a female. Both con- sumed their 4. macra host and required ap- proximately three months to complete de- velopment, which included two weeks to develop to the prepupal stage, six weeks to metamorphose from prepupal to pupal stages, and another two weeks to metamor- phose into mature adults. The P. simillima male emerged from its cocoon by 17 Sep- tember 1985. The female constructed an ill- formed cocoon and therefore was visible throughout its development. It was a ma- ture adult by 22 September 1985. Observing adult P. simillima at the nest- ing sites, and rearing of a male and female from A. macra nests clearly establishes a host-parasitoid association between A. macra and P. simillima. This association is especially significant because it broadens the VOLUME 93, NUMBER 1 host taxa by including the bee family An- drenidae. Dr. A. Menke (USDA-ARS Systematic Entomology Laboratory) and Dr. K. V. Krombein (Smithsonian Institution, Natl. Mus. Nat. Hist. Department of Entomolo- gy) identified the P. simillima. This research was a portion of an M.S. thesis submitted to the graduate faculty (Department of Zo- ology) at Howard University. Partial sup- 209 port was provided by a Minority Biomed- ical Research grant (RR-08016) awarded to Drs. R. M. Dufheld and J. Wheeler from the Division of Research Resources, Na- tional Institutes of Health. Eric W. Riddick, Department of Zoology, Howard University, Washington, D.C. 20059. Present address: Department of En- tomological Sciences, University of Califor- nia—Berkeley, California 94720. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 210-215 OBITUARY William Edwin Hoffmann 1896-1989 On September 12, 1989, William E. Hoff- mann passed away at Lawrence, Kansas fol- lowing heart failure. He will be remembered for his outstanding contributions during a long, useful, and exceedingly adventurous career. He was professor of entomology, re- searcher, and curator of the Natural History Survey and Museum at Lingnan University, Canton, China. He was an enthusiastic col- lector of plants and animals. His speciality was research on the biology of aquatic and semi-aquatic Hemiptera. Because so many species of the Scutelleridae, Pentatomidae, and Reduviidae in South China were of eco- nomic importance, he conducted research on their biology and control, and developed extensive catalogues for each family. He also undertook research on the biology and con- trol of other insect species injurious to ag- ricultural crops. He was a naturalist at heart and collected and made observations on in- sects, fishes, birds, reptiles, and plants. My brother, known by his friends as Bill Hoffmann, grew up on a homestead in Grant County, Oklahoma. He graduated from Caldwell, Kansas High School in 1914 and soon after moved to Lawrence, Kansas. He obtained an A.B. degree from the Univer- sity of Kansas and a Master’s degree from the University of Minnesota. Bill was on the entomology staffs of both universities and also worked for the State Entomologists of Kansas and Minnesota. In June, 1924, Bill was a member of the Minnesota Pacific Expedition. The expe- dition, composed primarily of faculty mem- bers of the University of Minnesota, con- ducted research on marine life. Participants were also delegates to the International Pan- Pacific Food Conservation Congress and Bill was secretary of the Fisheries Section. Following the Congress, he joined Ling- nan University, a private Chinese-Ameri- can University, in Canton, Kwangtung Province, in southern China. His first ap- pointment was Associate Professor and Acting Head of the Department of Biology and, the following year, he was advanced to Full Professor and Head. He founded the Lingnan University Natural History Survey and Museum in 1932 and remained in the Department of Biology as Professor of En- tomology. He was managing editor of the Lingnan Science Journal for thirteen years and founded and published the Lingnan Sci- ence Bulletin. Professor Hoffmann made many biolog- ical collecting expeditions in the hinterland of Kwangtung Province of China but he was considered an authority on the island of Hainan where he collected many new spe- cies of animals and plants. He made a spe- cial study of the aborigines there and lived for a time in the house of a chief. VOLUME 93, NUMBER 1 Bill also made biological investigations in Taiwan and the Philippine Islands. He spent several months in Luzon Province in the Philippines in 1927 and again in 1940 and 1941. During his 27 year affiliation with Ling- nan University, Bill was granted leave twice to undertake entomological research in Eu- rope and England. He traveled extensively, went around the world three times by ship, visited about 75 countries, and represented China at six international congresses. Bill was involved in the Chinese-Japanese war for five years and the experience of hun- dreds of bombings left an indelible mark on him. His greatest disappointment in life was the loss of his manuscript and artists’ orig- inal illustrations for a book on the insects of China. Bill lived under the Japanese Army of Occupation and helped operate the Ling- nan University refugee camp which cared for 8000 refugees. He was in Canton on Pearl Harbor day and was interned by the Japa- nese. Bill was repatriated in December, 1943 in an exchange of American and Japanese internees. While on leave from Lingnan University, Bill was an Associate Curator of Insects at the Smithsonian Institution, from 1944 to 1947. He returned to China in 1947 out of sheer dedication to the university and with the keen desire to preserve and to keep it out of the hands of the communists. In 1951, he returned to the United States and com- pleted a research project for Lingnan Uni- versity. He retired from Lingnan University two years later and then became affliated with the University Press at the University of Kansas For many years Bill served as a consultant on China. He retired from the University Press in 1962 but continued to enjoy keeping up with world events, fishing, and gardening until overcome by health problems. At the request of personnel at the University Archives, Kenneth Spencer Re- search Library, University of Kansas, Bill spent much time during the last two years of his life assembling letters and documents 211 of historical value relating to the changing times he and others experienced in South China. Bill’s amazing memory and ability to tell jokes and stories both humorous and tragic kept his family and friends spell- bound. Bill Hoffmann was a veteran of World War I. He was a member of the Entomo- logical Society of America, the Washington Entomological Society, the Kansas Ento- mological Society, and the First Presbyte- rian Church of Lawrence. He was a life member of the Masonic bodies of Lawrence Lodge #6 and Scottish Rite. His wife, Winifred, preceded him in death on November 26, 1986, but he is survived by myself and a sister, Goldie L. Case of Rossville, Illinois. Clarence H. Hoffmann, 6906 40th Ave- nue, Hyattsville, Maryland 20782. PUBLICATIONS OF WILLIAM E. HOFFMANN 1. Hoffmann, W. E. and J. E. Tilden. 1924. The Minnesota Pacific Expedition. Bull. Pan-Pacific Union, New Series, 52: 5-9. 2. Hoffmann, W. E. 1924. Are you tired of your goldfish or your canary bird? Aquatic Life 8: 1-3. 1924. One way in which the isolated worker may contribute effectively toward the so- lution of fishery problems in the Pacific. Bull. Pan-Pacific Union, New Series 56: 9-10. . 1924. Handy collecting apparatus. En- tomol. News 35: 253-255. 1924. Biological notes on Lethocerus americanus (Leidy). Psyche 31: 175-183. 1924. Winter food for waterbugs in aquaria. Bull. Brooklyn Entomol. Soc. 19: 149- 150. . 1924. The relation of the crow to pecan culture. The Wilson Bull.: 175-184. . 1924. The life histories of three species of gerrids (Heteroptera, Gerridae). Ann. Ento- mol. Soc. Amer. 17: 419-430. 1925. The interrelations of conserva- tion. Public Affairs 4: 6, 8. 1925. Repressing our aversion for snakes. Aquatic Life 8: 139-140. . 1925. The relation of life history studies to food conservation. Mid-Pacific Mag. 29: 394— 395. 12. ——. 10. Nols 1925. Some aquatic Hemiptera having nN —_ No nN N 23; 24. Die 33. 34. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON only four nymphal stages. Bull. Brooklyn Ento- mol. Soc. 20: 93-94. 1925. The life history of Velia watsoni Drake (Heteroptera, Veliidae). Can. Entomol. 57: 107-112. 1925. Collecting Microvelia in Hawaii. Bull. Brooklyn Entomol. Soc. 20: 132-133. 1925. The food of lizards in captivity. Aquatic Life 9: 33-35. 1925. Carp fishing in America. China Jour. Sci. Arts 3: 464-466. | plate. . 1925. The common snapping turtle not entirely carnivorous. Aquatic Life 9: 49-50. 1925. Some observations on the snap- ping turtle and their biological significance. Mid- Pacific Mag. 30: 259-262. . 1925. The three-toed box turtle. Aquat- ic Life 9. . 1925. Anew species of Nepa from South China (Heteroptera, Nepidae). Lingnan Agric. Rev. 3: 39. 1925. Notes on snakes in captivity. Aquatic Life 9. 1925. The biology of a common Chi- nese waterstrider (Heteroptera, Gerridae). Ling- nan Agric. Rev. 3: 48-63. 1925. Notes on collecting Microvelia (Heteroptera, Veliidae). China Jour. Sci. Arts 3: 606-608. . 1926. The life history of Sphaerodema rusticus Fabr. (Hemiptera, Belostomatidae). Lingnan Agric. Rev. 3: 167-176. 2 tables, | plate. 1927. Coccidae from China with a list of host plants. Lingnan Agric. Rev. 4: 73-76. . 1927. Biological notes on Laccotrephes (Hemiptera, Nepidae). Lingnan Agric. Rev. 4: 77-93. . 1927. A further note on a common Chi- nese waterstrider (Hemiptera, Gerridae). Ling- nan Agric. Rev. 4: 191-194. 1927. Charles Fuller Baker (obituary). Lingnan Agric. Rev. 4: 197-202. 1 plate. 1929. Notes on a squash-bug of eco- nomic importance. Lingnan Sci. J. 5: 281-292. 1929. Charles Walter Howard (obitu- ary). Lingnan Sci. J. 5: 293-297. . 1929. A stink-bug injurious to citrus in South China. Proc. Third Pan-Pacific Sci. Con- gress, Tokyo 1926: 2030-2038. 1929. Carl Oscar Levine (obituary). Lingnan Sci. J. 5: 357-362. 1930. Note on the larva of Cyrtotrach- elus longimanus F. Lingnan Sci. J. 6: 364-366. J. E. Tilden and W. E. Hoffmann. 1930. Basi- cladia, a new genus of Cladophoraceae. Bot. Gaz. 89: 374-384. . Hoffmann, W.E. 1930. The wood of water pine 36. aT: 38. 39) 40. 41. 42. 43. 44, 45. 46. 47. 48. 49. 50. a1 52. 33: roots as a substitute for balsa wood for ento- mological uses. Lingnan Sci. J. 9: 138-139. . 1930. The food habits of Erthesina fullo (Thunb.) Lingnan Sci. J. 9: 139-142. . 1930. Notes on the life history of Ran- atra chinensis Mayr. Proc. Nat. Hist. Soc. Fukien Christian Univ. 3: 31-37. 1 table, 1 plate. 1930. A peculiar habit in the water- strider Rhadadotarsus kraepelini Breddin. Ling- nani Scia Ja 92 321¢ 1931. The life history of Rhynchocoris humeralis Thunb. (Hemiptera, Pentatomidae). Lingnan Sci. J. 7: 817-~820. 2 plates. 1931. Notes on the biology of Ranatra filiformis (Hemiptera, Nepidae). Lingnan Sci. J. 9: 385-395. 1931. Life history notes on Enithares sinica Stal (Hemiptera, Notonectidae). Lingnan Sci. J. 9: 432-433. 1931. A pentatomid pest of growing beans in South China. Peking Nat. Hist. Bull. 5: 25-26. 1931. Eradication of nightshade (So- lanum nigrum L.) as an aid in the control of insects of economic importance in South China. Lingnan Sci. J. 10: 113-115. 1931. Notes on the Coreid, Anoploc- nemis phasiana Fabr. Lingnan Sci. J. 10: 131- 133% . 1931. The food plants of the adult Cer- copid, Cosmoscarta bimaculata Walk. Lingnan Sci. J; 10133: 1931. Notes on Hemiptera and Ho- moptera at Canton, Kwangtung Province, Southern China, 1924-1929. Bur. of Entomol., U.S.D.A. Ins. Pest Survey Bull. 11: 138-151. . 1931. Revised bibliography of the works of the late professor C. W. Howard. Lingnan Sci. J. 10: 335-337. 1931. Coprophagus beetles from Hai- nan Island. Lingnan Sci. J. 10: 501-502. 1931. Critheus lineatifrons Stal injuri- ous to bamboo. Lingnan Sci. J. 10: 502-503. 1931. Studies on the bionomics of the water-bug Lethocerus indicus (Hemiptera, Belo- stomatidae) in China. Verh. Int. Verein. theoret. angewandete Limnologie 5: 661--667. 1932. A list of the Pentatomidae, Plat- aspidae, and Coreidae (order Hemiptera) of Chi- na, Korea, and Indo-China. J. Pan-Pacific Res. Inst. 7: 6-11. . 1932. Notes on the bionomics of some oriental Pentatomidae (Hemiptera). Archivio Zool. Italiano (atti dell’XI Congresso Interna- zionale di Zoolocia, Padova, 1930) 16: 1010- 1027. 1932. The economic status of the ly- VOLUME 93, NUMBER 1 54. 5: 56. Sli. 58. oe) 60. 61. 62. 63. 64. 65. 66. 67. 68. 69: 70. qh gaeids and notes on the life history of Lygaeus hospes Fabr. and Aphanus sordidus Fabr. (He- miptera, Lygaeidae). Lingnan Sci. J. 11: 119-135. . 1932. On pentatomids collected by the Lingnan University Fifth Hainan Island Expe- dition, 1929. Lingnan Sci. J. 11: 139-141. 1932. Interesting flies from the island of Hainan. Lingnan Sci. J. 11: 289-290. Hoffmann, W. E., P. Marshall, G. B. Cressey, W. Repetti, T. Wayland Vaughan, and C. M. Fraser. The Fourth Pacific Science Congress, Java, 1929. Lingnan Sci. J. 11: 292-304. Hoffmann, W. E. 1932. Brief note on Vitruvius Insignis Distant. Lingnan Sci. J. 11: 470. 1 plate. . 1932. The economic status of the genus Eurydema with biological notes on E. pulchrum Westw. (Hemiptera, Pentatomidae). Lingnan Sci. J. 11: 553-564. 1 plate. 1932. Oides decempunctata (Billberg), a chrysomelid pest of cultivated grape (Vitis lam- brusca Linn.). Lingnan Sci. J. 11: 565-566. 2 plates. 1932. The life history of Poecilocoris drurnaei L. (Hemiptera, Pentatomidae) in Can- ton, China. Lingnan Sci. J. 11: 567-569. 2 plates. 1933. Life history notes on some Kwangtung, China, coreids (Hemiptera, Corei- dae). Lingnan Sci. J. 12: 97-127. 10 plates. . 1933. Note on Membracidae (Homop- tera) from Hainan Island. Lingnan Sci. J. 12: 137- 138. 1933. The life history of a second spe- cies of Laccotrephes (Hemiptera, Nepidae). Ling- nan Sci. J. 12: 245-256. 1 plate. 1933. The biology and control of Lac- coptera chinensis F. (Coleoptera, Chrysomeli- dae). Lingnan Sci. J. 12: 259-260. 1 plate. . 1933. Some basic needs of Entomology in China with special reference to South China. Lingnan Sci. J. 12 (Suppl.): 233-240. 1933. A preliminary list of the aquatic and semi-aquatic Hemiptera of China, Chosen (Korea), and Indo-China. Lingnan Sci. J. 12 (Suppl.): 243-258. 1933. Observations on a weevil inju- rious to banana. Hong Kong Naturalist 4: 48-54. 1933. Remarks on Poecilocoris with special reference to P. Latus Dallas (Hemiptera, Pentatomidae). Lingnan Sci. J. 12; 435-439. 1 plate. 1933. Biological note on Cosmoscarta bispecularis (White) (Homoptera, Cercopidae). Lingnan Sci. J. 12: 447-451. 1933. Additional data on the life his- tory of Lethocerus indicus (Hemiptera, Belostom- atidae). Lingnan Sci. J. 12: 595-601. 3 plates. 1933. Notes on Cosmoscarta heros F. 72) 13% 74. 1. 76. His 78. 72 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 213 with a description of the last nymphal stage (Ho- moptera, Cercopidae). Lingnan Sci. J. 12: 605- 607. 1 plate. 1934. Preliminary notes on the fresh- water fish industry of South China, especially Kwangtung Province. Lingnan Univ. Sci. Bull. 5. 70 pp.; colored frontispiece, 11 plates. 1934. The life history of a species of Graptostethus (Hemiptera, Lygaeidae). Lingnan Scit F131 71176: 1934. The egg of the Lygaeid, Lygaeus hospes Fabr. (Hemiptera). Lingnan Sci. J. 13: 181. 1934. Notes on the Coreid, Hygia opa- ca (Uhler). Lingnan Sci. J. 13: 313-321. 1934. Abnormal tibia in nymph of Aquarius paludum (Fabr.). Lingnan Sci. J. 13: 331-332. . 1934. Robert Emerson Wall (obituary). Lingnan Sci. J. 13: 333-334. 1 plate. 1934. The status of entomology in Kwangtung Province with suggestions for pro- cedure. Agricultural Information Nos. 173-174: 145-171. (College of Agric., Sun Yatsen Univ., Canton, China). 1934. The life history and economic status of Sycanus croceovittatus Dohrn (Hemip- tera, Reduviidae). Lingnan Sci. J. 13: 505-516. 2 plates. . 1934. Cratonotus coloratus Distant 1879 in China. Lingnan Sci. J. 13: 523-525. 1934. Morphological notes on Megar- rhamphus truncatus (Westw.): Hemiptera, Pen- tatomidae. Lingnan Sci. J. 13: 693-696. 1934. The overuse of certain abbrevi- ations in scientific publications. Lingnan Sci. J. 13: 699. 1934. Tree borers and their control in Kwangtung [in Chinese, English summary]. Ling- nan Agric. J. 1: 37-59. . 1934. Biological notes on the stink-bug Metonymia glandulosa (Hemiptera, Pentatomi- dae). Hong Kong Naturalist 5: 272-278. 1935. The bionomics and morphology of Isyndus reticulatus Stal (Hemiptera, Reduvi- idae). Lingnan Sci. J. 14: 145-153. 4 plates. 1935. Description of the last nymphal stage and other notes on Brachycerocoris camelus Costa (Hemiptera, Pentatomidae). Lingnan Sci. J. 14: 165-168. 1935. Notes on the life history of Neo- Jurtina typica Dist. (Hemiptera, Pentatomidae). Lingnan Sci. J. 14: 261-265. 2 plates. . 1935. Further notes on Brachycerocoris camelus Costa (Hemiptera, Pentatomidae). Ling- nan Sci. J. 14: 327-328. Hoffmann, W. E., I. J. Condit and H. C. Wang. 1935. Observations on the culture of oranges 214 90. OF 92: 93. 94. 95: 96. 97. 98. 9S: 100. 101. 102. 103. 104. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON near Swatow, China. [in Chinese, English sum- mary]. Lingnan Agric. J. 1: 175-248. Hoffmann, W. E. 1935. Notes on eight Penta- tomids not hitherto recorded from Formosa. Trans. Nat. Hist. Soc. Formosa 25: 57-62. . 1935. The life history, economic status, and control of three injurious leaf beetles (Co- leoptera, Chrysomelidae). Lingnan Sci. J. 14: 505—- 517. 7 plates. . 1935. Life history notes on a species of Lawana (Homoptera, Flatidae). Lingnan Sci. J. 14: 521-524. 3 plates. Hoffmann, W. E. and Y. W. Djou. 1935. En- tomology for agricultural middle schools in Kwangtung Province, No. 1, Hemiptera or True Bugs [in Chinese, common names also in En- glish]. Lingnan Agricultural Middle School, Can- ton. 37 pp. (Abstr. in Lingnan Sci. J. 14: 729.) Hoffmann, W. E. 1935. An abridged catalogue of certain Scutelleroidea (Plataspidae, Scuteller- idae, and Pentatomidae) of China, Chosen, Indo- China, and Taiwan. Lingnan Univ. Sci. Bull. 7. iv + 294 pp. + 2 maps, 2 tables. . 1935. Observations on a Hesperid leaf- roller and a lace-bug, two pests of banana in Kwangtung. Lingnan Sci. J. 14: 639-649. 5 plates. 1935. A novel native method of con- trolling June-beetles. Lingnan Sci. J. 14: 689- 690. 1 plate. 1935. Injurious insects in Kwangtung. The Rocan 4: 8-17. Canton, China. 1935. Notes on Kwai Fa Shim (Le- thocerus indicus Lep. & Serv.), an insect of eco- nomic importance [in Chinese and English]. Lingnan Agric. J. 1: 167-174. . 1936, The bionomics of the rose sawfly, Arge victorina Kirby (Hymenoptera: Argidae), with notes on other injurious Tenthredinoid lar- vae in Kwangtung. Lingnan Sci. J. 15: 101-112. 1 plate. 1936. Diaphorina citri Kuw. (Homop- tera: Chermidae), a citrus pest in Kwangtung. Lingnan Sci. J. 15: 127-132. 1936. Notes on citrus insects [in Chi- nese, English summary]. Lingnan Agric. J. 2: 165— 218. Hoffmann, W. E. and B. W. Fuson. 1936. Notes on three widely distributed insect pests of vege- tables and their occurrence at Canton. [in Chi- nese, English summary]. Lingnan Agric. J. 2: 219- 236. Hoffmann, W. E. 1936. The life history of Lim- nogonus fossarum (Fabr.) in Canton (Hemiptera: Gerridae). Lingnan Sci. J. 15: 289-299. 2 plates. 1936. Alucita niveodactyla Pag. (Mi- crolepidoptera: Pterophoridae), a pest of sweet potatoes in Kwangtung. Lingnan Sci. J. 15: 311- 312: 105. 106. 107. 108. 109. 110. Pi: 12: 113. 114. 115. 116. 117. 118. 119: 120. WA 122 1936. Life history notes on Rhagado- tarsus Kraepelini Breddin (Hemiptera: Gerridae) in Canton. Lingnan Sci. J. 15: 477-482. 2 plates. 1936. The occurrence of Limnometra gigas China (Hemiptera: Gerridae) in Hainan Is- land, with description of the apterous form. Ling- nan Sci. J. 15: 489-492. 2 plates. 1937. Distributional notes on some Kwangtung and Kwangsi fishes. Lingnan Sci. J. 16: 95-115. 1937. Lawana increasing in impor- tance as a noxious insect. (Homoptera, Flatidae). Lingnan Sci. J. 16: 119-120. . 1937. Kwangtung Aphididae including host plants and distribution. Lingnan Sci. J. 16: 267-302. . 1937. Further notes on citrus insects in Kwangtung [in Chinese and English]. Lingnan Agric. J. 2: 378--396. . 1937. Coleoptera collected by the Ling- nan University Fifth Hainan Island Expedition. Lingnan Sci. J. 16: 631-637. 1938. The nidification of the bamboo bee, Xylocopa dissimilis Lepel. (Hymenoptera: Xylocopidae). Lingnan Sci. J. 17: 87-91. 1938. Records of Coleoptera, Hyme- noptera, and Isoptera from the island of Hainan. Lingnan Sci. J. 17: 91-94. 1938. Formosan tree sparrows (Frin- gillidae) damage brick wall. Lingnan Sci. J. 17: 94-96. Hoffmann, W. E., Y. C. Ng, and H. W. Tsang. 1938. Life history studies in nine families of Kwangtung butterflies (Lepidoptera: Rhopaloc- era). Lingnan Sci. J. 17: 227-246; 4 plates; 407- 424: 2 plates; 515-532; 6 plates. Hoffmann, W. E. 1938. Distributional and bi- ological notes on some Kwangtung Diptera. Ling- nan Sci. J. 17: 251-260. 1938. Coleoptera and Hymenoptera from Kwangtung including Hainan Island. Ling- nan Sci. J. 17: 439-460. 1940. The foodplants of Nezara viri- dula Linn. (Hemiptera: Pentatomidae), pp. 81 1- 816. Jn Proc. Sixth Int. Congr. Entomol., 1935, Madrid. 1941. Catalogue of aquatic Hemiptera of China, Indo-China, Formosa, and Korea. Lingnan Sci. J. 20: 1-78; 78A—78E; 1 map. 1942. Notes on beetles of the genus Sagra (Coleoptera, Sagridae). Lingnan Nat. Hist. Surv. Mus. Spec. Publ. 6: 1-8; 2 plates. . 1942. Bugs or Homoptera: the cicadas, plant hoppers, plant lice, scale insects, and others. Lingnan Nat. Hist. Surv. Mus. Spec. Publ. 9. 20 pp.; 30 plates. . 1942. Lingnan trees, a list of trees grow- VOLUME 93, NUMBER 1 123% 124. 125: ing on Lingnan University campus. Lingnan Nat. Hist. Surv. Mus. Spec. Publ. 11. 15 pp. 1944. Catalogue of the Reduviidae of China. Lingnan Univ. Sci. Bull. 10. 80 pp. 1947. A smarting anal fluid ejected by some exotic Pentatomids. Proc. Entomol. Soc. Wash. 49: 142-143. . 1948. First supplement to Catalogue of Scutelleroidea. Lingnan Sci. J. 22: 1-41. 126. WAT. 128. 129. 215 . 1949. Insects as human food. Proc. En- tomol. Soc. Wash. 49: 233-237. . 1950. Second supplement to Catalogue of Scutelleroidea. Lingnan Sci. J. 23: 21-42. 1953. Third supplement to Catalogue of Scutelleroidea. Philippine J. Sci. 82: 309-318. . 1954. The life history of Glaucias cras- sa (Westwood) (Hemiptera: Pentatomidae). Phil- ippine J. Sci. 83: 83-92. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 216-217 Book REVIEW Special Biotic Relationships in the Arid Southwest. Edited by Justin O. Schmidt. University of New Mexico Press, Albu- querque. 1989, 152 pp. Cloth. $22.50. To the uninitiated, deserts are barren and sterile. Hot, unquestionably (except cold de- serts of polar fringes of the Northern Hem1- sphere); bleak, perhaps; depauperate, cer- tainly not. Any sense of emptiness is illusory, for dry deserts usually abound in plant and animal life. Anyone taking time to explore them—rather than passing through on an interstate highway or viewing them from the window of an airplane—will encounter biological richness. Biodiversity is especial- ly characteristic of the Sonoran Desert. Bot- anists, zoologists, and ecologists continue to discover fascinating adaptations and spe- cializations related to coping with the in- tense heat and scarcity of water. Research- ers also must contend with complex interactions between organisms. Special Biotic Relationships in the Arid Southwest resulted from a symposium on desert animals and plants held in Tucson, Arizona, in March 1985. As stated in the preface, its purpose “‘is to present detailed analyses of ways in which various biotic forms have evolved and survived in the arid Southwest.” That goal has been met—and probably exceeded. An appropriate lead-off chapter is the in- triguing story of desert drosophila, cacti, and microorganisms. The system involves four Drosophila species endemic to the Sonoran Desert; columnar cacti, including the famil- lar organ pipe and saguaro, that differ in stem chemistry (alkaloids, lipids, triterpene gly- cosides, etc.); yeasts living on necrotic cactus tissue; and bacteria that apparently initiate the rotting process on cactus stems. Yeast abundance and distribution are largely de- termined by chemical differences among cac- ti, with distribution influenced by Drosoph- ila spp. and their host specificity. The flies, which are thought responsible for inoculat- ing young areas of decay with yeasts, breed either in necrotic stems of cactus or in soil soaked by rot juice. Of the 24 possible sub- strate-fly combinations—given five cactus species, soaked soils, and four Drosophila spp.—only six are normally used. Host plant selection and utilization involve attraction of the flies to volatiles, assimilation of ne- crotic cactus tissues, and ability to tolerate or detoxify antiherbivore compounds in the hosts. Much of the chapter emphasizes cac- tus stem chemistry as the main determinant of host use by the drosophilids and as the probable cause of the volatile patterns pro- duced in rots characteristic of each cactus species. The interactions described in this chapter, too complex to be presented here, form the chemical and biological basis of a model system in evolutionary biology. Chapter 2 details the evolutionary inter- actions between harvester ants, Pogono- myrmex spp., and horned lizards of the ge- nus Phrynosoma. Despite characteristics rendering them vulnerable to predation, seed-harvesting ants have few predators; they possess a powerful sting and a toxic venom that is injected through the sting shaft. Horned lizards, a highly derived group of iguanids native to the western states and Mexico, are ant predators that often spe- clalize on Pogonomyrmex. They have evolved a broad, flattened shape, with large stomach and digestive tract, necessary for a diet of nutrient-poor, chitinous prey. Their resistance to harvester ant venom involves a blood factor(s) that neutralizes or detox- ifies the mouse-lethal activity of the venom. Other lizards are not resistant to Pogono- myrmex venom, and horned lizards cannot neutralize snake or bee venoms. As the au- thors of this chapter point out, this is the first example of a vertebrate known to show VOLUME 93, NUMBER 1 specific resistance to the venom of an ar- thropod. Termite ecology is discussed in the next two chapters. Chapter 3 elucidates the role of a subterranean termite, Gnathamitermes tubiformans (Buckley), in breaking down or- ganic matter on the soil surface and in ni- trogen cycling in the northern Chihuahuan Desert. Interactions among termites, nitro- gen availability, and soil-water status are shown to affect the composition and phe- nology of the desert plant community. The presence of termites also affects populations of mites, collembola, and other soil microar- thropods. Foraging ecology of subterranean termites (of several genera) in the Sonoran Desert (Santa Rita Experimental Range near Tucson) is presented in chapter 4. Chapter 5 calls attention to the abun- dance of spiny and thorny plants (woody and suffrutescent perennials) and venomous animals in the Arizona Upland Sonoran Desert. Spines and venoms are important in protecting desert organisms against her- bivory and predation. The relative painful- ness of stings of solitary and social Hyme- noptera are compared, and the physiological and behavioral basis of the protective value of venoms against large animals is analyzed. The final chapter, in focusing on the bi- ological effects of agricultural development, departs from the previous chapters. Dis- cussed is pecan monoculture along the flood plain of the Santa Cruz River in southern Arizona and the biological and social re- percussions of this irrigated desert agroeco- system. Some of the topics considered are pesticide and water use, development of floodwater mosquito populations in or- chards, weed management, and burning of tree prunings. The biotic community asso- ciated with pecan farming is shown to be similar to, but distinct from, that of declin- ing native riparian habitats. No significant problems with this edited book were found. I noticed some minor in- consistencies in abbreviations used in the references, e.g., Am. and Amer., Ent. and 27, Entomol., and Nat. and Natl. (for National), and in capitalization (new world, Old World). There are the usual typographical errors, such as soitary (for solitary), salt- brushes (for saltbushes), Ekol. Polksa (for Polska), and cresotebush; in one instance there was failure to italicize (occidentalis, p. 35) or to close parentheses (Viperidae, p. 40). There is an occasional grammatical problem (for instance, p. 117: “The exact causes of this pain has not been investigated ...’) ora potentially confusing sentence (p. 35: “Unlike most lizards that are general predators or herbivores, horned lizards spe- cialize in ants as prey.” In the last sentence, it would have been better to say: “Unlike most lizards, which are general predators or herbivores, .. .”” On p. 112, readers are told twice that the number of spider species hav- ing vertebrate-active venoms is not known. Botanists disagree on the correct name for creosotebush in southwestern deserts, 1.e. whether Larrea tridendata is distinct from the South American L. divaricata. In the book, both names are used to refer to creo- sotebush in the desert Southwest. These, however, are nearly all trivial problems, ones many readers will not find objectionable. Overall this is a well-execut- ed book. Ecologists and evolutionary biol- ogists studying the biota of the arid South- west, as well as nonspecialists and naturalists, should find it useful. Editor (and chapter author and coauthor) Justin Schmidt and nine other contributors capture the uniqueness of the Chihuahuan and Sonoran deserts. I was sorry to see the book end— don’t stop at six case histories, give us more. But Special Biotic Relationships will stim- ulate research on plants and animals of the arid Southwest. Because equally fascinating relationships remain to be elaborated, we can look forward to a similar symposium on Southwestern desert life and to reading additional case histories. A. G. Wheeler, Jr., Bureau of Plant In- dustry, Pennsylvania Department of Agri- culture, Harrisburg, Pennsylvania 17110. PROC. ENTOMOL. SOC. WASH. 93(1), 1991, pp. 218-221 MANUSCRIPT PREPARATION GUIDELINES FOR THE PROCEEDINGS OF THE ENTOMOLOGY SOCIETY OF WASHINGTON ROBERT D. GORDON AND F. CHRISTIAN THOMPSON Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, % U.S. National Museum of Natural History Washington, D.C. 20560 GENERAL POLICY Articles for publication are ordinarily ac- cepted only from members of the Society and must be in English. A summary in French, German, Spanish or Russian is ac- ceptable. Such a summary should be placed immediately after the English abstract and may be a translation of that abstract. Man- uscripts should not exceed 15 printed pages (two typewritten pages are approximately equivalent to a printed page). Manuscripts are peer-reviewed before they are accepted and final acceptance is the responsibility of the Editor. Articles are published in the or- der received rather than in order of date of acceptance. Immediate publication can be had for payment of full page charges. Notes and book reviews are published as space is available. NAMES OF ORGANISMS The first mention of a plant or animal should include the full scientific name in- cluding the authority. For taxonomic papers a complete citation is required for all sci- entific names. Use only common names ap- proved in Common Names of Insects and Related Organisms published by the Ento- mological Society of America. Spell out the entire scientific name the first time it is men- tioned; thereafter abbreviate genus names except at the beginning of a paragraph or sentence. Do not abbreviate a genus name when it begins a sentence; e.g., use ““Xus albusis:...<” rather than ““Xialbus is’..25” Within sentences throughout the text do not use a specific name that is not preceded by the generic initial, e.g. “This species is sim- ilar to X. albus because... .” ILLUSTRATIONS No extra charge is made for line drawings or half tones. Authors should plan illustra- tions for reduction to the dimensions of the printed page and allow room for legends at the top and bottom. Do not make plates larger than 14/18”. Individual figures must be mounted on suitable board, and photo- graphs (glossy only) should be trimmed, grouped together, and abutted when mount- ed; the printer will insert the hairlines. In- clude title of paper, author’s name and ad- dress, and illustration numbers on the back of each plate. Figures should be numbered consecutively and in the sequence in which they are referred to in the text. Reference in the text should be in the style: (Fig. 1) and (Table 1). See Figure 3 for proper for- mat of figure legends. Each table or figure with its title or legend should be under- standable without reference to text. MANUSCRIPT PREPARATION Type manuscripts on letter size paper. All parts of the manuscripts should be double spaced including tables, table headings, lit- erature cited, etc. Allow 12 inch margins on all sides. Use a title page with name, address, and telephone number of the au- thor to whom all correspondence and proof is to be sent at the upper left of the first page. The abstract is to be typed on a separate page, complete with key words as in Fig. 2. Key words are a must. VOLUME 93, NUMBER 1 219 Return Proof to: John R. Doe 315 State St. Meriden, CT 06420 (203) 555-1212 A new species of Xus (Order: Family) injurious to hollies, Ilex spp. (Aquifoliaceae) John R. Doe and John Smith (JRD) Resident Biologist, 315 State St., Meriden, Connecticut 06420; (JS) Entomologist, City Parks, Hartford, Connecticut 06540. Abstract.- Xus albus, a new species of ... is described, illustrated, and compared with ... Key Words. Distribution, ornamental shrub, damage, leaf roller Figure Legends Figs. 1-4. Xus albus. 1, Habitus. 2, Male genitalia (lateral view). 3, Larva. 4, Pupa. Fig. 5. Damage to holly leaves. Literature Cited Doe, J. and J. Smith. 1970. Holly Insects. Jones and Case. New York, New York. 38 pp. Smith, J. and J. R. Doe. 1967. A list of insects injurious to hollies (Ilex spp.). Proceedings of the Entomological Society of Virginia 38:54-68. Figs. 1-4. Sample pages for submitted manuscripts. 1, Title page. The mailing address and phone number of the person(s) to whom the page proofs will be sent should be typed in the upper left-hand corner. A proof can be sent to each author of a two- or multi-authored paper if they desire. Affiliation and full address are typed as a paragraph below the author name(s). Spell out the state name. 2, Abstract page. Note that “Abstract” begins the paragraph. On the same page, follow abstract with 3-5 key words. 3, Figure legend. Note punctuation. Start a new paragraph for legends describing each plate. 4, Literature Cited page. Note punctuation, spacing, and complete journal titles. 220 Do not use all capitals for any reason. Underscore only where italics are intended in the body of the text. The sequence of material in submitted manuscripts should be: Title, Author(s), Abstract, Text, Acknowledgements, Liter- ature Cited, Author’s(s) Address(es), Ap- pendix, Figure Legends, Figure copies (each numbered and identified), Tables (each ta- ble numbered with an Arabic numeral and with heading provided). Titles of taxonomic papers must be qual- ified to indicated the systematic position of all the taxa treated. The order and family must be included except where only family level taxa are treated, in which case only the order is required. All taxa below family level mentioned in the title must be accompanied by the authority name. LITERATURE CITED When citing informational references in text, use the form “Erwin (1990)” if the au- thor’s name is part of the sentence and “‘(Er- win 1983)” if it is not. Two articles by one author cited at one time should be written “(Erwin 1989, 1990)’; two articles pub- lished by the same author in the same year, “(Erwin 1990a, 1990b).” “In press” citations in text should be avoided. Use the year of expected publi- cation and end the citation listing in the Literature Cited section with ‘‘(in Press)”’. Citations with more than two authors are cited in the text as: Mathis et al. (1976). Do not underline the “et al.”’. Multiple citations in the text should be organized chronolog- ically and then alphabetically. Names of persons providing unpublished information should include initials when referenced in the text such as: D. White- head, pers. comm. (denotes information ob- tained orally), and D. Whitehead, in litt. (denotes information obtained in a writing). The list of references at the end of the manuscript should have the primary head- ing “Literature Cited.” Only papers referred PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to in the text may be listed and the list must be alphabetically by author. The volume number of a journal or other serial should be cited but the issue number in journals having continuous pagination throughout the year or volume should not be included. Underline scientific names or other words only if italicized in the original title. In citing books, complete collation is not required: give pagination for the body of the work. Complete collation is required only when the auxiliary sections contain the first reference to a taxon under considera- tion. Plates (or Figures), if not included in paginated materials, should be referenced after pagination (e.g., 200 pp. + 10 pls.). Ranges are to be used only for partial col- lation, that is, there is additional material beyond that cited (e.g., 1-200 pp. + 1-10 pls. for works with more than 200 pages and 10 plates). Journal names and book titles are set in Roman type, so do not underline. Abbreviations cannot be used in Litera- ture Cited. All words must be included and spelled out including conjunctions, prepo- sitions, etc. Manuscripts will be returned to the author if journal names are not com- pletely spelled out. CHARGES Authors will be assessed a charge to cover editing and publication expenses. The cur- rent charge to members is $35.00 per print- ed page. These charges are in addition to those for reprints and author’s correction in proof. The charge for long papers, those over 15 printed pages, papers by non-members, and papers for immediate publication is $55.00 per page. Member authors who are retired or not affiliated with an institution may request to have charges waived and should do so when the manuscript is sub- mitted. A waiver may delay publication. Charges for author errors and/or changes in proof, for reprints if ordered, for immediate publication, and for non-members are not waived. Acceptance for papers is based only VOLUME 93, NUMBER 1 on their scientific merit without regard to the author’s financial support. REQUIREMENTS FOR TAXONOMIC PAPERS Taxonomic papers must conform to re- quirements of the International Code of Zoological Nomenclature. In addition, type specimens must be designated for new spe- cies described, type depositories must be clearly indicated, new taxa must be clearly differentiated from existing taxa by means of keys or differential diagnoses. In short, these papers must conform to good taxo- nomic practices. More detailed guidelines for taxonomic papers are available on re- quest from the Systematic Entomology Lab- oratory, ARS, USDA. ACKNOWLEDGMENTS Parts of this manuscript were taken ver- batim or paraphrased from manuscripts on the same subject by H. Larew (1990) and another manuscript by C. B. Robbins and D. B. Lellinger (1990). LITERATURE CITED Larew, H. 1990. Instruction to authors for prepara- tion of manuscripts. Proceedings of the Entomo- logical Society of Washington 92:166-168. Robbins, C. B. and D. B. Lellinger. 1990. Guidelines for manuscripts for publications of the Biological Society of Washington. Proceedings of the Bio- logical Society of Washington, Supplement to Vol- ume 103, No. 1 March 1990. 30 pp. 3G © ein we kine, nile : 1S Pestle “ an omer iii athe er ak y a Nisa * @) ie x tT) vi ; ‘ ’ ed Pore) iT res Pe, Li Fe Ae NG ae 1 AT) (A Te acy Mead it TLL Aba > aah al | ne 4 . ee ee Ask | : Sate apne vs (ata 1 : Th AFIT me Ye Th iP iv PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS Seip salls-of the Eastern United States, by Lewis H. Weld occas scttecneenine eee Sos. Of the Southwest, by Lewis H: Weld cc . SPER ADCESTOMSC VMI IG ptt seme ee ee etn Votre eh ee Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman....-.:0:-:cv:vnovnonensnvnocn Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. | DoS Bes BE NN SN a tek th oa ae ND IO OR ee SITE Oe, Se AL a A Short History of the Entomological Society of Washington, by Ashley B. Gurney... Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. ST Lal a UR Ee th ee ei a NE SE ERP SE eel a alin, AERA MO am Ser Ue dele a ae a Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. SEDO SLIED eS CLs Gi es De SIRE Se Nye OA er 3 Res SAO Ere a. PCN We Nc PA a MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving. 23) [ols UG Ps Reed Eg SR a a Re Se ee ee Ek ie Se a ere a ORT SREP No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. TESS (ByOy, UR ce Bae SN ae IO te CUP Ns SE cart ON Ra AE We we cRinenia: Ue MRE, Ahead rea a No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952.00 No.5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. " Ah anwar ap SLMS CM wine Eel eae Le TNE gs ee eee i ON No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi METH ASI EOS Ose DOO ee ee te we eames PG Wah Lea Smile ee Eola ee a frie Coe No.7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. OE ESD cai gS as Soca SSS ga SURE gl AE ee Aa eRe caer A en i et A ec Ie No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cera- topogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 cc cececesceeeeeeeeeeeeetes No.9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200 (BPD SIRT EES gS is iy Be aan oN SENAY. A le EE SNORE ACR iene EEE Ltr ol Yul Cv an Ne ee No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W. Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982... No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp. neers felts 5s tee ea ee eh AL ee ay eRe ay No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. Saar a tee 8 ee RN, ee TOE ee ee Ne te Ye No. 13. An Identification Manual for the North American Genera of the Family Braconidae (Hy- menoptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987 15.00 11.00 12.00 10.00 11.00 18.00 5.00 18.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $25.00 per volume to non-members and $13.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10 per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian Institution, Washington, D.C. 20560. CONTENTS (Continued from front cover) McDANIEL, B. and A. BOE— Morphological differences in genitalia of Bruchophagus (Hyme- noptera: Chalcidae) that infest alfalfa, red clover, and birdsfoot trefoil seeds (Hymenoptera: Eurytomidae) McKAMEY, STUART H. and LEWIS L. DEITZ—Synonymy in the treehopper genera Hoplo- Dhorion, Metcalfiella, and Ochropepla (Homoptera: Membracidae) McKAMEY, STUART H. and LEWIS L. DEITZ—Nomenclatural changes in the treehopper tribes Hoplophorionini, Smiliini, and Talipedini (Homoptera: Membracidae) MENKE, ARNOLD S. and H. H. EVENHUIS—North American Charipidae: Key to genera, nomenclature, species checklists, and a new species of Di/yta Forster (Hymenoptera: Cy- nipoidea) NORRBOM, ALLEN L.—The species of Anastrepha (Diptera: Tephritidae) with a Grandis-type wing pattern RAY, CHARLES H. JR. and MICHAEL L. WILLIAMS—Two new species of Matsucoccus Cockerell (Homoptera: Margarodidae) similar to Matsucoccus alabamae Morrison SCHMUDE, KURT L. and HARLEY P. BROWN—A new species of Stenelmis (Coleoptera: Elmidae) found west of the Mississippi River WIEGMANN, BRIAN M.—A new species of Japanagromyza (Diptera: Agromyzidae) from Florida, with a key to North American species WIRTH, WILLIS W.—New and little-known species of Forcipomyia (Diptera: Ceratopogonidae) associated with cocoa pollination in Brazil NOTES DAVIS, DONALD R.—Lectotype designation for Opostega heringella Mariani, a synomym of Opostega spatulella Herrich-Schaffer (Lepidoptera: Opostegidae) JENKINS, JOHN—A simple device to clean insect specimens for museums and scanning electron microscopy RIDDICK, ERIC W.—Pseudomethoca simillima (Smith) (Hymenoptera: Mutillidae), a newly discovered parasitoid of Andrena macra Mitchell (Hymenoptera: Andrenidae) TORMOS, J. and S. F. GAYUBO—On intraspecific variation in the Alysiinae (Hymenoptera: Braconidae) OBITUARIES HOFFMANN, CLARENCE H.—William Edwin Hoffmann, 1896-1989 BOOK REVIEWS WHEELER, A. G. JR.— Special Biotic Relationships in the Arid Southwest * ‘ - | VOL. 93 APRIL 1991 NO. 2 (ISSN 0013-8797) my (orp PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED QUARTERLY CONTENTS ANDERSON, ROBERT S.—Larva and pupa of Cleonidius erysimi (Fall) with a discussion of the phylogenetic position of Lixini (Sensu Kuschel) (Coleoptera: Curculionidae) ......... 288 ASHWORTH, ALLAN C. and DONALD P. SCHWERT—On the occurrences of Opisthius richardsoni Kirby and Asaphidion yukonense Wickham (Coleoptera, Carabidae) as late Pleis- (OGENS WORSLS iat Bel ies Se ie eB Oke MR Ser es. ay ae Ona 5 Re ar ee ee Sil BALL, GEORGE E.—Taxonomic notes about the Middle American genus Cyrtolaus Bates CeoleapteranCaranidaesPterOstiehini)\: oat. )ee Se A eae-n Sleek eke ee hice piste eels Sake ue 230 BELLAMY, C. L.—A revision of the genus Eudiadora Obenberger (Coleoptera: Buprestidae) 409 CLARK, WAYNE E.—The Anthonomus guttatus species group (Coleoptera: Curculionidae)... 262 COLONNELLI, ENZO— New World Cnemogonini (Coleoptera: Curculionidae). New species, Mewsconibinations anda type species designation; ..)) 21 4..4. 02 hse. . ee nla srln acc oes 272 FLINT, O. S., JR.—On the identity of Chloronia bogatana Weele (Neuropterida: Megaloptera: CG SPRCTTVG AYE) See eel ata ae Ri Rly GI nM i i aL NO a a es Ot aa or ge 489 { FRANIA, HENRY E.—Displacement of one taxon by another as the cause of certain ecological shifts in Eustilicus Sharp (Coleoptera: Staphylinidae): a test of the evidence ............ 437 GORDON, ROBERT D.— West Indian Coccinellidae IV (Coleoptera): New genera and species OVS UECTN ETC TT US 2) ONS” AEN OC CE san a7 eee eee ee ak a Ge Balke anne eee Ae fe te 298 HENRY, THOMAS J.—Melanotrichus whiteheadi, a new crucifer-feeding plant bug from the southeastern United States, with new records for the genus and a key to the species of eastern North, America (reteroptera; Miridae: Orthotylinae) 25.5. 0.00. be ee a 449 HOEBEKE, E. RICHARD—An Asian ambrosia beetle, Ambrosiodmus lewisi, new to North ted Meme CTA SCO LIdAG)) ye eek. keke lt) oI, Sela RNS BT Rg a 420 HOFFMAN, RICHARD L.—A useful taxonomic distinction between two similar Cychrine NeeHestmOleoplerasG@aravidar eet) sah) eek Fete M eh One GaN obit ye SER ee 407 JOHNSON, PAUL J.— Taxonomic notes, new records, and a key to the adults of North American thes (Os ae Det |e ee Ee OU ee a a A, 0a OE 7 ce Lge a (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1991 DAvID R. SMITH, President NORMAN E. WOODLEY, 7reasurer WAYNE N. MATHIs, President-Elect GARY STECK, Program Chairman M. ALMA SoLis, Recording Secretary RUSSELL B. STEWART, Membership Chairman HOo.uis B. WILLIAMS, Corresponding Secretary JEFFREY R. ALDRICH, Past President JAMES B. STRIBLING, Custodian ROBERT D. Gorpbon, Editor THOMAS J. HENRY, Associate Editor Publications Committee DONALD R. DAVIS TERRY L. ERWIN GEORGE C. STEYSKAL F. CHRISTIAN THOMPSON Honorary President CURTIS W. SABROSKY Honorary Members LoulIsE M. RUSSELL ALAN STONE THEODORE L. BISSELL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, NHB 168, Smithsonian Insti- tution, Washington, D.C. 20560. MEETINGS. — Regular meetings of the Society are held in the Natural History Building, Smithsonian Institution, on the first Thursday of each month from October to June, inclusive, at 8 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.— Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $20.00 (U.S. currency)-of which $18.00 is for a subscription to the Proceedings of the Entomological Society of Washington for one year. PROCEEDINGS. -— The Proceedings are published quarterly beginning in January by The Entomological Society of Washington, % Department of Entomology, NHB-168, Smithsonian Institution, Washington, D.C. Members in good standing receive the Proceedings of the Entomological Society of Washington. Nonmember subscriptions are $50.00 per year, domestic, and $60.00 per year, foreign (U.S. currency), payable in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE P. 218 OF THE JANUARY, 1991 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560. Editor: Robert D. Gordon, Systematic Entomology Laboratory, ARS, % Department of Entomology, Smith- sonian Institution, |Oth and Constitution NW, Washington, D.C. 20560. Books for Review: T. Henry, Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 11 June 1991 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN»PRESS, INC., LAWRENCE, KANSAS 66044, USA | THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. 224 PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 224-225 THE D. R. WHITEHEAD MEMORIAL ISSUE, Editor’s Preface Donald R. Whitehead was a respected member of the entomological profession and a valued member of the Entomological Society of Washington. The Society has chosen to remember him by publication of this memorial issue composed of papers written by his friends and colleagues. Traditionally such issues have been composed of articles on the group of organisms studied by the scientist; however, Don’s interests were so varied as to defy classification, therefore this volume is composed of papers on a wide assemblage of topics. Many new taxa described herein are patronyms in his honor, an irony because Don’s interests lay, not in describing new species, but in utilizing taxa to create classifications useful in developing hypotheses of wide ranging significance. How- ever, time constraints inherent in preparing a volume of this nature precluded much of the cladistic analyses that contributors would normally have utilized. Don influenced many students and colleagues in their scientific thinking and his friendly, always available advice will be missed. The Society is honored to dedicate this volume to his memory. VOLUME 93, NUMBER 2 229 Don Whitehead; in Mexico with snake; in USNM beetle collection; somewhere in Alberta; somewhere in Alberta; in his office at the USNM (left to right). PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 226-229 OBITUARY Donald R. Whitehead 1938-1990 Donald R. Whitehead of the Systematic Entomology Laboratory, Agricultural Re- search Service, Beltsville, Maryland, died on May 4, 1990. His death was a great loss not only to the science of taxonomy but to science in general. Don’s interests were never circumscribed by the confines of a single taxon, rather they were catholic, encompassing a diverse as- semblage that included beetles, wasps, true bugs and millipeds. He was interested in what could be learned from the study of groups of organisms rather than simply ar- ranging them in a logical system. His selec- tion of study groups reflected this; for ex- ample, his interest in millipeds stemmed from his perception that a study of their distribution patterns might result in devel- opment of a hypothesis explaining modern distributions of other organisms. Don’s mil- liped research did in fact result in a hy- pothesis that he termed “mosaic distribu- tion,” published in a volume on milliped systematics. His interest in natural history began early; many were his stories about early adven- tures in the woods and swamps near his home. This interest led him to a B.S. in entomology from Rutgers University, fol- lowed by a two year stint in the Medical Corps of the United States Army. He then entered a Ph.D. program in entomology at the University of Alberta, completing it in 1971. Four years of working as a Research Entomologist, Organization for Tropical Studies, at the U.S. National Museum, led to his final employment in the Systematic Entomology Laboratory, USDA, at the same institution. He was a seeming contradiction in mood and temperament, usually cheerful but of- ten appearing moody or short tempered. His mood was, except in the last couple of years when health was a factor, occasioned by the quality of the conversation or subject matter thereof to which he was exposed. Short of patience with trivial gossip, he would ex- pound for hours on matters he deemed sig- nificant. At these times, we who were his usual companions could begin to realize the depth and breadth of his mental capacities. These capacities enabled him to function as an excellent reviewer and editor of the writ- ten word, especially when asked to partic- ipate in manuscript review. He spent an ex- traordinary amount of time helping authors of scientific papers, in the process changing many scientific contributions from ordinary to excellent. Don was an excellent field person and there were few, if any, phases of his life he enjoyed more. From early New Jersey days through graduate student trips to Mexico and finally West Virginia milliped hunts, he searched for specific groups of arthropods to obtain distributional and host data for his research purposes. In his last years he was deeply involved in milliped collecting in the Appalachian Region, concentrating on the state of West Virginia with its great habitat diversity. As a result, he became deeply attached to those West Virginia hills, spending as much time there as his health and responsibilities would allow. Of all the hours spent in Don’s company, the best re- membered are those spent in the field. Clear are the memories of certain places, Villa Nueve, New Mexico, Panamint Dunes, Cal- ifornia, Bear Heaven, West Virginia, etc. Times that are fully appreciated only in ret- VOLUME 93, NUMBER 2 rospect when there can be no more, when one can only wish that they had not been so brief. No ordinary man this, intelligent beyond his peers, thoughtful, caring, willing to lis- ten, to help, to advise—of all those friends who have passed before me, none will be more sorely missed than Don. Robert D. Gordon Washington, D.C. Dr. DONALD WHITEHEAD’S PUBLICATIONS Whitehead, D. R. 1966. Two remarkable new South American species of Schizogenius Putzeys (Cole- optera: Carabidae). Coleopterists Bulletin 20: 1-7. Whitehead, D. R. 1966. A review of Halocoryza Al- luaud, with notes on its relationship to Schizo- genius Putzeys (Coleoptera: Carabidae). Psyche 73: 217-228. Ball, G. E.and D. R. Whitehead. 1967. Localities for collecting in Mexico. Coleopterists Bulletin 21: 122-138. Whitehead, D. R. 1969. Variation and distribution of the intertidal beetle Halocoryza arenaria (Dar- lington) in Mexico and the United States (Cole- optera: Carabidae). Journal of the New York En- tomological Society 77: 36-39. Whitehead, D. R. 1969. Notes on Dyschirius Bonelli and Akephorus LeConte, with a peculiar new Dys- chirius from Texas (Coleoptera: Carabidae: Scar- itini). Journal of the New York Entomological So- ciety 77: 179-192. Whitehead, D. R. 1972. Classification, phylogeny, and zoogeography of Schizogenius Putzeys (Co- leoptera: Carabidae: Scaritini). Quaestiones En- tomologicae 8: 131-348. Whitehead, D. R. 1973. Annotated key to Platynus, including Mexisphodrus and most “‘Colpodes,” so far described from North America including Mex- ico (Coleoptera: Carabidae, Agonini). Quaestiones Entomologicae 9: 173-217. Whitehead, D. R. 1974. Notes on three Chaudoir species of Platynus from Central America (Carab- idae: Pterostichini: Agoni). Coleopterists Bulletin 28: 103-104. Whitehead, D. R. 1974. An annotated key to de- scribed species of the Neotropical genus G/ypto- lenus (Carabidae: Pterostichini: Agoni). Coleop- terists Bulletin 28: 123-132. Kingsolver, J. M. and D. R. Whitehead. 1974. Bio- systematics of Central American species of Cteno- colum, a new genus of seed beetles (Coleoptera: Bruchidae). Proceedings of the Biological Society of Washington 87: 283-312. 227 Whitehead, D. R. 1974. Variation and synonymy in Hypselonotus (Heteroptera: Coreidae). Journal of the Washington Academy of Sciences 64: 223- 233% Kingsolver, J. M. and D. R. Whitehead. 1974. Clas- sification and comparative biology of the seed beetle genus Caryedes Hummel (Coleoptera: Bru- chidae). Transaction of the American Entomolog- ical Society 100: 341-436. Whitehead, D. R. 1974. Clivina texana LeConte, a synonym of C. planicollis LeConte (Coleoptera: Carabidae: Scaritini). Proceedings of the Ento- mological Society of Washington 76: 454. Whitehead, D. R. 1975. Studies of Conotrachelus Schonherr and Microscapus Lima (Coleoptera: Curculionidae: Cryptorhynchinae) associated with Hymenaea courbaril Linnaeus in Central Ameri- ca, with notes on the Cristatus Group of Cono- trachelus. Journal of the Washington Academy of Sciences 65: 36—40. Whitehead, D.R. 1975. Additions to “Annotated key to Platynus” (Coleoptera: Carabidae: Agonini). Quaestiones Entomologicae 11: 6. Whitehead, D. R. 1975. Parasitic Hymenoptera as- sociated with bruchid-infested fruits in Costa Rica. Journal of the Washington Academy of Sciences 65: 108-116. Whitehead, D. R. and J. M. Kingsolver. 1975. Bio- systematics of the North and Central American species of Gibbobruchus (Coleoptera: Bruchidae: Bruchinae). Transaction of the American Ento- mological Society 101: 167-225. Whitehead, D. R. and J. M. Kingsolver. 1975. Mega- sennius, a new genus for Acanscelides muricatus (Sharp) (Coleoptera: Bruchidae) a seed predator of Cassia grandis L. (Caesalpiniaceae) in Central America. Proceedings of the Entomological So- ciety of Washington 77: 460-465. Whitehead, D. R. andG. E. Ball. 1975. Classification of the Middle American genus Cyrtolaus Bates (Coleoptera: Carabidae: Pterostichini). Quaes- tiones Entomologicae 11: 591-619. Whitehead, D. R. and J. M. Kingsolver. 1976. Beetles and wasps associated with Cassia biflora L. (Caes- alpiniaceae) fruits in Costa Rica, with a new spe- cies of Sennius (Coleoptera: Bruchidae). Journal of the Washington Academy of Sciences 65: 154— 157. Kingsolver, J. M. and D. R. Whitehead. 1976. The North and Central American species of Meibo- meus (Coleoptera: Bruchidae: Bruchinae). ARS USDA Tech. Bulletin No. 1523: 1-54. Whitehead, D. R. 1976. Classification and evolution of Rhinochenus Lucas (Coleoptera: Curculionidae: Cryptorhynchinae), and Quaternary Middle American zoogeography. Quaestiones Entomolo- gicae 12: 118-201. 228 Whitehead, D. R. 1976. Note to contributors and readers. Coleopterists Bulletin 30: 112. (Editorial) Whitehead, D. R. 1976. Editorial. Coleopterists Bul- letin 30: 212. Whitehead, D. R. 1976. Collecting beetles in exotic places without leaving home: The herbarium. Co- leopterists Bulletin 30: 249-250. (Editorial) Whitehead, D. R. 1976. Admissions and acknowl- edgements. Coleopterists Bulletin 30: 367. (Edi- torial) Clark, W. E., D. R. Whitehead, and R. E. Warner. 1977. Classification of the weevil subfamily Tychiinae, with a new genus and species, new com- binations, and new synonymy in Lignyodini (Co- leoptera: Curculionidae). Coleopterists Bulletin 31: 1-18. Whitehead, D. R. 1977. Cerambycidae of North America. Coleopterists Bulletin 31: 64. (Book re- view) Whitehead, D. R. 1977. Subtribes Dyschiriina and Clivinina. Jn Reichardt, H., ed., A synopsis of the genera of Neotropical Carabidae (Insecta: Cole- optera). Quaestiones Entomologicae 13: 390-392. (Taxonomic notes) Whitehead, D. R. 1977. Diagnosis. In Pests Not Known to Occur in the United States or of Limited Distribution. A Weevil, Baris lepidii Germar (Co- leoptera: Curculionidae). Cooperative Plant Pest Report 2: 681-682. (Identification Aid) Whitehead, D. R. 1977. New Apion (Coelocephala- pion) species (Curculionidae: Apioninae) from fruits of Pterocarpus (Leguminosae: Faboidea) in Central America. Coleopterists Bulletin 31: 165— 172. Whitehead, D. R. and H. Reichardt. 1977. Classifi- cation of Listropus Putzeys, a subgenus of Schi- zogenius Putzeys (Coleoptera: Carabidae: Scari- tini). Coleopterists Bulletin 31: 239-250. Erwin, T. L., D. R. Whitehead, and G. E. Ball. 1977. North American Beetle Fauna Project, Checklist, Yellow Version: Family 4, Carabidae, the Ground Beetles. World Digest Publications, Kinderhook, New York. 68 pp. Batra, L. R., D. R. Whitehead, E. E. Terrell, A. M. Golden, and J. R. Lichtenfels. 1978. Overview of predictiveness of agricultural biosystematics, pp. 275-310. In Romberger, J. A., R. H. Foote, L. Knutson, and P. L. Lentz, eds., Beltsville Sym- posia in Agricultural Research. [2] Biosystematics in Agriculture. Allanheld, Osmun & Co. Publish- ers, Inc., Montclair, New Jersey. xii + 340 pp. (Book chapter) Whitehead, D. R. 1978. Apion (Trichapion) candyae new species (Coleoptera: Curculionidae), a gall- maker of leaf petioles of Erythrina berteroana Ur- ban (Fabaceae) in El Salvador. Coleopterists Bul- letin 32: 193-201. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Whitehead, D. R. 1978. What should be published in The Coleopterists Bulletin? Coleopterists Bul- letin 32: 204. (Editorial) Whitehead, D.R. 1979. Speciation patterns and what they mean, pp. 23-33. Jn Erwin, T. L., G. E. Ball, D. R. Whitehead, and A. L. Halpern, eds., Carabid Beetles: Their Evolution, Natural History, and Classification. Proceedings of the First Interna- tional Symposium of Carabidology, Smithsonian Institution, Washington, DC, August 21, 23, and 25, 1976. Dr. W. Junk bv, Publishers, The Hague. 635 pp. (Book chapter) Whitehead, D. R. 1979. Recognition characters and distribution records for species of Conotrachelus (Coleoptera: Curculionidae) that damage avocado fruits in Mexico and Central America. Proceedings of the Entomological Society of Washington 81: 105-107. Whitehead, D.R. 1979. Notes on Apteromechus Faust of America north of Mexico (Coleoptera: Curcu- lionidae: Cryptorhynchinae). Proceedings of the Entomological Society of Washington 81: 230-233. Hoebeke, E. R. and D. R. Whitehead. 1980. New records of Rhinoncus bruchoides (Herbst) for the Western Hemisphere and a revised key to the North American species of the genus Rhinoncus (Cole- optera: Curculionidae: Ceutorhynchinae). Pro- ceedings of the Entomological Society of Wash- ington 82: 556-561. Whitehead, D.R. 1980. Probable evolution and mor- phological variation in South American disonych- ine flea beetles (Coleoptera: Chrysomelidae) and their amaranthaceous hosts, by G. B. Vogt, J. U. McGuire, Jr., and A. D. Cushman. Coleopterists Bulletin 34: 352-354. (Book review) Whitehead, D.R. 1981. The weevil Apion (Perapion) curtirostre Germar in North America, and the sys- tematic position of Perapion Wagner with a review of the North American species (Coleoptera: Cur- culionoidea: Apionidae). Coleopterists Bulletin 34: 397-400. Anderson, D. M., R. D. Gordon, J. M. Kingsolver, T. J. Spilman, R. E. White, and D. R. Whitehead. 1981. Illustrated keys to adult and larval beetles (Coleoptera) found associated with wood at inter- ception points. Printed and distributed by APHIS Training Center, Frederick, Maryland. 25 pp. Whitehead, D. R. 1982. Foods of Caulophilus spp., particularly the broadnosed grain weevil, C. ory- zae (Gyllenhal), based on interception records (Coleoptera: Curculionidae: Cossoninae). Pro- ceedings of the Entomological Society of Wash- ington 84: 81-84. Whitehead, D. R. and R. M. Duffield. 1982. An un- usual specialized predator prey association (Co- leoptera: Coccinellidae, Chrysomelidae): Failure VOLUME 93, NUMBER 2 of a chemical defense and possible practical ap- plication. Coleopterists Bulletin 36: 96-97. Whitehead, D. R. 1983. Pests Not Known to Occur in the United States or of Limited Distribution, No. 31: Apple Blossom Weevil. APHIS 81-43, 10 pp. September 1983. Whitehead, D. R. and D. M. Anderson. 1983. Pests Not Known to Occur in the United States or of Limited Distribution, No. 32: European Chestnut Weevil. APHIS 81-43, 10 pp. September 1983. Alonso Zarazaga, M. A. and D. R. Whitehead. 1983. Nomenclatural notes on Apionidae (Coleoptera: Curculionidae). Proceedings of the Entomological Society of Washington 85: 626-627. Wheeler, A. G. and D. R. Whitehead. 1985. Larinus planus (F.) in North America (Coleoptera: Cur- culionidae: Cleoninae) and comments on biolog- ical control of Canada thistle. Proceedings of the Entomological Society of Washington 87(1): 207- PANSY Whitehead, D. R. 1985. Notes on Schizogenius put- zeysi Kirsch (Coleoptera: Carabidae: Scaritini). Coleopterists Bulletin 38: 226. Whitehead, D.R. 1985. Systematics of Apion (Trich- apion) johnschmitti, anew species from Costa Rica (Coleoptera: Curculionoidae: Apionidae). Cole- opterists Bulletin 39: 111-118. Whitehead, D. R. and K. Whittle. 1985. Pests Not Known to Occur in the United States or of Limited Distribution, No. 64: A Curculionid Weevil. APHIS 81-46, 15 pp. September 1985. Whitehead, D. R., D. M. Anderson, and K. Whittle. 1985. Pests Not Known to Occur in the United States or of Limited Distribution, No. 65: Banded 729 Pine Weevil. APHIS 81-46, 13 pp. September 1985. Shelley, R. M. and D. R. Whitehead. 1986. Part V. Colors and color patterns. Jn Shelley, R. M. and D. R. Whitehead, eds., A reconsideration of the milliped genus Sigmoria, with a revision of Del- totaria and an analysis of the genera in the tribe Apheloriiini (Polydesmida: Xystodesmidae). Memoir of the American Entomological Society 35: 169-174. Whitehead, D. R. and R. M. Shelley. 1986. Part VI. Relationships within Sigmoria. In Shelley, R. M. and D. R. Whitehead, eds., A reconsideration of the milliped genus Sigmoria, with a revision of Deltotaria and an analysis of the genera 1n the tribe Apheloriiini (Polydesmida: Xystodesmidae). Memoir of the American Entomological Society 35: 174-204. Shelley, R. M. and D. R. Whitehead. 1986. Part VIII. Relationships in the tribe Apheloriini. /n Shelley, R. M. and D. R. Whitehead, eds., A reconsider- ation of the milliped genus Sigmoria, with a re- vision of Deltotaria and an analysis of the genera in the tribe Apheloriiini (Polydesmida: Xystodes- midae). Memoir of the American Entomological Society 35: 212-218. Whitehead, D. R. and A. G. Wheeler, Jr. 1990. What is an immigrant arthropod? Annals of the Ento- mological Society of America 83: 9-14. Colonnelli, E.and D. R. Whitehead. 1990. A revision of the genus Bangasternus (Coleoptera, Curcu- lionidae). Fragmenta Entomologica 22: 185-217. Whitehead, D. R. (In press). Weevils (Curculionidae, Coleoptera). Jn Gorham, R. D., ed., Insect and Mite Pests in Food: An Illustrated Key. (Book chapter; September 1992) PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 230-239 TAXONOMIC NOTES ABOUT THE MIDDLE AMERICAN GENUS CYRTOLAUS BATES (COLEOPTERA: CARABIDAE: PTEROSTICHINI) GEORGE E. BALL Department of Entomology, University of Alberta T6G 2E3, Edmonton, Alberta, Can- ada Abstract. — Based on features of adults and on chorological considerations, 2 new species of Cyrtolaus (s. str.) are described from Oaxaca, México: C. oaxacanus (type locality— Sierra Zempoal, 10.4 km S. Totontepec), and C. whiteheadi (type locality —Sierra Madre de Oaxaca, 11.7 km E. Huautla de Jimenez). Character states and localities of specimens not recorded previously of the Chiapan-Guatemalan C. subiridescens Whitehead and Ball, C. ricardo Whitehead and Ball, C. furculifer Bates, and C. grumufer Whitehead and Ball confirm generally the published treatments of these taxa. Key Words: In August, 1965, toward the beginning of the 14-month period that Don Whitehead and I spent in México, we found in a cloud forest in central Chiapas 2 carabid adults of rather remarkable form (heavy-bodied, but with strikingly long mandibles, narrow head, and markedly vaulted elytra) that we thought represented some aberrant species of the platynine genus Colpodes (sensu latissimi). Some years later, after studying many of the types that H. W. Bates had described in writing the carabid part of the Biologia Cen- trali-Americana, we concluded that our specimens were members of the genus Cyr- tolaus Bates (1882: 99). We described this species and four others as new, in a revision of Cyrtolaus (Whitehead and Ball 1975). In that treatment, we combined in a single ge- nus /thytolus Bates and Cyrtolaus, ranking each group as a subgenus. Because the range of elytral features of Cyrtolaus (sensu lato) seemed to bridge the range of elytral features of the tribes Ptero- stichini and Agonini (= Platynini), we con- cluded that this genus belonged in neither tribe in the restricted sense, that they should new species, phylogeny, biogeography, Mexico, Coleoptera, Carabidae be ranked as subtribes of Pterostichini, and that Cyrtolaus should go in its own subtribe, Cyrtolaina, which we proposed. The last de- cision was based on both the seeming in- termediate position of Cyrtolaus in the Pterostichini, and its seeming relict status — the genus seemed not to have close phylo- genetic relatives. Liebherr (1986) discovered that 2 Antil- lean species, described originally as mem- bers of Colpodes, in fact represented an un- described genus, which he named Barylaus. Further, Liebherr determined that Barylaus and Cyrtolaus, though phylogenetically iso- lated, were endemic New World represen- tatives of the pterostichine subtribe Coe- lostomina, a group that had been thought to be confined to the Old World (except for an introduced species of Coelostomus in the Antilles). Thus, the name Cyrtolaina White- head and Ball, 1975 is a junior synonym of Coelostomina. Also, Liebherr argued that the platynines should be ranked as a tribe, and thus equivalent to the Pterostichini, and not as a component of the latter. I regard as elegant Liebherr’s discovery VOLUME 93, NUMBER 2 of the subtribal relationships of Cyrtolaus and Barylaus. However, on the basis of cho- rological considerations, I do not accept his conclusion that Barylaus is more closely re- lated to the geographically distant Afro- tropical lineage of Coelostomina than to the more proximal Middle American Cyrto- laus. As knowledge of relationships of Cyrto- laus was increasing, so was additional ma- terial accumulating of the species. Below, I report and analyze the new information thus acquired, to seek validation of the structural and chorologically diagnostic features of taxa previously described, and to make known taxa not known previously. MATERIAL AND METHODS Material This study is based on 36 adults of the subgenus Cyrtolaus, collected in the Mexi- can states of Oaxaca and Chiapas, and in southwestern Guatemala. Four-letter cod- ens used in the text to indicate sources or depositories of study material are: CASC_ Department of Entomology, Cal- ifornia Academy of Sciences; UASM __ Department of Entomology, Uni- versity of Alberta; and USNM Department of Entomology, United States National Museum of Natural History. METHODS Recognition. — Species were identified us- ing the key and illustrations in Whitehead and Ball (1975), and identity was confirmed by examination of the male genitalia, which were prepared using standard techniques. Specimens the combination of features of which did not fit the couplets and regarded initially as taxonomically distinct entities were examined in detail. If such examina- tion showed them to be distinctive in few and varied features, the specimens were 1n- cluded as members of the described species to which each was most similar. If the more 231 detailed examination suggested otherwise, the specimens were regarded as taxonomi- cally distinct, and were ranked as species. For specific ranking of C. oaxacanus, new species, primacy was given to chorological features. See below for details. Measurements. — The following measure- ments were made as part of the descriptions of taxa and to compare with those reported by Whitehead and Ball (1975: Tables 1-3). HL = length of head from base of dorsal condyle of left mandible to poste- rior margin of eye; ML = length of left mandible, from apex to base of dorsal condyle; PL = length of pronotum along mid-line, from anterior to posterior margin; EL = length of elytra, along suture, from basal ridge to apex. Size was expressed as Standardized Body Length (SBL) and is the sum of HL + PL + EL. Two ratios were computed: ML/PL and PL/EL. Not reported in detail are values for the ratio Antennomere 3: width/length. Values are between 0.26 and 0.32, thus well within the range reported previously (0.26-0.38) for Cyrtolaus (s. str.), excluding C. lobipen- nis. These relatively low values indicate the relatively elongate antennae that are char- acteristic of the most derived lineage of Cyr- tolaus. Chorology.— Localities reported are in the following mountain ranges: Sierra Madre de Oaxaca and Sierra Zempoal, in the Mexican state of Oaxaca (see outline map of the re- gion in Ball and Roughley [1982: 364, Fig. 63]); and Mesa Central, in central and northwestern Chiapas, and Sierra Madre de Chiapas, in southeastern Chiapas and southwestern Guatemala (see outline map of region in Whitehead and Ball [1975: 606, Figs. 22, 23)). KEY TO SPECIES OF CYRTOLAUS BATES, BASED ON CHARACTERISTICS OF ADULTS This key is modified from the one pre- sented by Whitehead and Ball (1975: 598), and includes all known species of Cyrtolaus. 232 Ol. Ol’. 02(01’). 02’. 03(02’). 03’. 04(03’). 04’. 05(04). OS: 06(05’). 06’. 07(04'). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pronotum with anterior marginal bead complete; postero-lateral setae near pos- terior angles. Elytron with plica clearly developed, epipleuron interrupted (““crossed’’); interneurs impunctate. Ab- dominal sternum VII with 2 setae in male, 4 in female Subgenus /thytolus Bates C. orizabae Bates Pronotum without anterior marginal bead; postero-lateral setae distinctly an- terad posterior angles (Figs. 2-4). Elytron with epipleuron interrupted or not; in- terneurs more or less punctate (Figs. SA— 7A). Abdominal sternum VII with 4 setae in male, 6 to 8 setae in female iSO Oe Subgenus Cyrtolaus Elytron with epipleuron interrupted by plica. Labrum with 4 marginal setae C. lobipennis Bates Elytral epipleuron not interrupted. La- brum with 6 marginal setae 03 Elytron with microsculpture of trans- verse lines, surface faintly iridescent; in- terneurs grossly punctate; intervals cari- nate; apex spined or not, spines shorter or longer (Figs, '6ASB)i eee ee Elytron with microsculpture mesh pat: tern isodiametric; apex spined ........ 04 Head with 2 (single pair) or 3 supraorbital SetIgeTOUS PUNCHUTES. 44. hcl OS Head with 4 (2 pairs) supraorbital setig- CLOUSIPUNCHINES tes ete see 07 Elytron with interval 2 apically more or less flat, not distinctly swollen (Figs. 5A, B). Geographical range—Huautla Pla- teau, Oaxaca ips. Payson C. whiteheadi, new species, p. 236. Elytron with interval 2 swollen apically. Geographical range—Mesa Central and Sierra Madre de Chiapas, Chiapas, and Guatemalan. ta: ae erm sciais ctars toate Elytron with interval 3 markedly raised apically. Disc of pronotum with micro- sculpture normal, surface without seri- ceous luster. Geographical range —Sierra Madre de Chiapas, Chiapas and Guate- mala C. furculifer Bates, p. 238. Elytron with interval 3 not raised api- cally. Disc of pronotum with microlines dense, surface with sericeous luster. Geo- graphical range—western part of Mesa Central, Chiapas 06 treat tee C. grumufer Whd. & Ball, p. 238. Elytron with interneurs grossly punctate. Pronotum with hind angles rounded, lo- C. subiridescens Whd. & Ball, p. 237. 07’. 08(07’'). 08’. 09(08’). 09’. 10(09’). 10’. bate, lateral margins broadly rounded, not sinuate; anterior angles produced. Geo- graphical range—central Guatemala ... C. spinicauda Bates Elytron with interneurs hardly punctate to distinctly so. Pronotum with hind an- gles various; lateral margins less round- ed, or sinuate posteriorly (Figs. 2, 4); an- terior angles not produced Geographical range—Sierra Madre de Chiapas, southeastern Chiapas and southwestern Guatemala. Elytron with apical declivity gradual (Fig. 7B), apical spine longer or shorter (Fig. 7A) gs nance C. ricardo Whd. & Ball, p. 238. Geographical range—Mesa Central of Chiapas, or mountains of Oaxaca. Ely- tron with apical declivity various; apical spine short (Fig. 5A) Pronotum with posterior angles rectan- gular. Elytron with interneurs distinctly punctate on disc; slope of elytral declivity more abrupt. Geographical range— Mesa Central, Chiapas 08 09 Pronotum with posterior angles acute. Elytron with interneurs very obscurely punctate on disc; apical declivity sloped gradually (Fig. 5B) Larger (SBL more than 11.0 mm); prono- tum relatively shorter (ML/PL 0.66-0.74). Male with apex of median lobe pointed (Fig. 1A). Geographical range—Sierra Zempoal (vicinity of Volcan Zempoal- tepetl, Oaxaca) C. oaxacanus, new species, p. 232. Smaller (SBL less than 10.0 mm); prono- tum relatively longer (ML/PL 0.60). Geographical range—Sierra Madre de Oaxaca (near Valle Nacional, Oaxaca) . C. newtoni Whd. & Ball New TAxA, NEw LocALity RECORDS, AND VARIATION IN DIAGNOSTIC FEATURES Cyrtolaus oaxacanus, NEW SPECIES (Figs. 1 A—-C) Type material.— HOLOTYPE male and ALLOTYPE female, each labelled: MEX. OAXACA 10.4 km./ s. Totontepec cloud/ forest: tree ferns/ in litter 2480 m./ June 17, 1979 79-38; MEXICAN EXP. 1979/ J. S. Ashe, G. E. Ball/ and D. Shpeley/ Collec- VOLUME 93, NUMBER 2 233 dh Fig. 1. Male genitalia of Cyrtolaus oaxacanus, new species: A, median lobe, apical portion, dorsal aspect; B and C, median lobe, with internal sac everted—B, left lateral aspect; C, right lateral aspect. Scale bar = 1.0 mm. Legend (for features of internal sac): 1, 2, and 3, spinose sclerites 1, 2, and 3, respectively; C-scl, C-sclerite; V.1., ventral lobe. tors. The holotype is in USNM, and allo- type in UASM. Type locality.—México, Oaxaca, Sierra Zempoal, 10.4 km S. Totontepec. Derivation of specific epithet.—The Lat- inized adjectival form of “Oaxaca,” the name of the Mexican state that includes the type locality of this species. Recognition.—The type specimens re- semble closely the holotype of C. newtoni Whitehead and Ball. However, they are larger (Table 1), and the pronotum is rela- tively shorter, as reflected by higher values for the ratio ML/PL (Table 2). These 2 taxa occur in different but adjacent mountain ranges (C. oaxacanus in the Sierra Zempoal, at 2400 m; C. newtoni in the Sierra Madre de Oaxaca, at 1700 m). The 2 pairs of supraorbital setae distin- guish adults of C. oaxacanus from C. white- headi, the third species of Cyrtolaus known from Oaxaca, the adults of which have only a single pair of such setae. The virtually impunctate elytral inter- neurs, angulate elytral apices and isodia- metric mesh pattern of the elytra distinguish adults of C. oaxacanus from those in the mountains of Chiapas and Guatemala. Males are distinguished by details of the genitalia (Figs. 1A—C): apex of median lobe acute (Fig. 1A); internal sac with sclerite | hardly evident (Fig. 1B), and spinose scler- ites 2 and 3 small, and widely separated from one another (Figs. 1B, C). Description. — Adults with character states of subgenus Cyrtolaus, and following. Data PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 234 9-0-0 —0-0—0-0-9-0-0-0~— 0-9 — 0 —0— 0-9 — 0 ©~0—e-0~0—0—o— 0—0— 0-2 ~o Oe oo 6.., 1, C. whitehead: Left elytron, A, 2, dorsal aspect of: > Pronotum 4 2-4 4, C. ricardo Wh Pronota and left elytra of species of Cyrtolaus. 7 = Figs new species dorsal aspect > 7 = 1eS itehead and Ball iteheadi, new spec > head and Ball f left lateral aspect, and 7, C. ricardo Wh ite subiridescens Whi Gs 3 3 ’ subiridescens G 6, > C. wh 3; f: ly, o itehead and Ball respective outline o > B 3 > Whitehead and Ball VOLUME 93, NUMBER 2 235 Table 1. Data about variation in Standardized Body Length (SBL—mm) among samples of the species of Cyrtolaus (s. str.). Males Females Species and Localities N Range Mean N Range Mean C. oaxacanus Toton., Oax. 1 1M 2 1 11.26 C. whiteheadi Rio Sant., Oax. 2 11.12-12.19 11.65 C. subiridescens Mitont., Chis. 7 10.92-11.39 E27 3 10.99-11.99 11.39 Tenejapa, Chis. 1 10.72 Las Marg., Chis. 2 10.38-11.26 10.82 DD 11.06-11.32 11.19 Las Rosas, Chis. 1 11.93 C. ricardo Tacana, Chis. 7 10.72-11.86 11.14 l 10.99 San Marc, Guat. l 10.65 2 10.65-10.99 10.82 C. furculifer Prov., Chis. 1 18333 1 WES 2 Muxbal, Chis. 2 12.73-13.33 13.03 C. grumufer Rizo Oro, Chis. 1 12.66 Table 2. Data about variation in values for the ratio ML/PL among sample of the species of Cyrtolaus (s. SET)» Males Females Species and Localities N Range Mean N Range Mean C. oaxacanus Toton., Oax. 1 0.66 l 0.74 C. whiteheadi Rio Sant., Oax. 2 0.59-0.61 0.60 C. subiridescens Mitont., Chis. a 0.64-0.69 0.65 3 0.65-0.71 0.67 Tenejapa, Chis. 1 0.67 Las Marg., Chis. D) 0.67-0.69 0.68 2 0.64-0.65 0.645 Las Rosas, Chis. 1 0.59 C. ricardo Tacana, Chis. 7 0.65-0.68 0.67 l O72 San Marc, Guat. 1 0.73 2 ONS 0.75 C. furculifer Prov., Chis. 1 0.55 0.63 Muxbal, Chis. p 0.59-0.63 0.61 C. grumufer Rizo Oro, Chis. 1 0.56 236 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Data about variation in values for the ratio PL/EL among sample of the species of Cyrtolaus (s. str.). Males Species and Localities N Range C. oaxacanus Toton., Oax. 1 0.42 C. whiteheadi Rio Sant., Oax. C. subiridescens Mitont., Chis. 7 0.44—0.48 Tenejapa, Chis. 0.46 Las Marg., Chis. 2 0.44-0.46 Las Rosas, Chis. 1 0.47 C. ricardo Tacana, Chis. 7 0.41-0.44 San Marc, Guat. 1 0.42 C. furculifer Prov., Chis. 1 0.46 Muxbal, Chis. C. grumufer Rizo Oro, Chis. about variation in Standardized Body Length, and in values for ratios ML/PL and PL/EL, in Tables 1-3. Microsculpture. Average for Cyrtolaus (s. str.), elytra with mesh pattern isodiametric. Fixed setae. Average for Cyrtolaus (s. str.) Head, antennae and mouthparts average. Pronotum (slightly damaged in holotype; somewhat distorted in allotype) probably normally similar to that of C. newtoni and C. whiteheadi (Fig. 2; description below). Elytra. In form, similar to Figs. 5A, and B, but apical angles less projected. Inter- neurs on disc nearly impunctate, but punc- tures on apical declivity of moderate size. Legs. Average for Cyrtolaus (s. str.). Male genitalia (Figs. 1! A—C). Median lobe in dorsal aspect with apical portion (Fig. 1 A) moderately long, tapered to a point. En- dophallus (internal sac) with sclerite | hard- ly evident (Fig. 1B), spinose sclerites 2 and 3 of moderate size, prebasal in position, widely separated from one another; C-scler- Females Mean N Range Mean 1 0.38 2 0.44—0.46 0.45 0.46 3 0.41-0.46 0.44 0.45 2 0.43-0.44 0.435 0.43 1 0.40 2 0.40 0.40 1 0.44 2 0.41-0.44 0.42 1 0.50 ite evident, on right side; ventral lobe (V.1.) prominent. Habitat.—The type specimens were col- lected in cloud forest, under bark of a fallen tree, in the vicinity of tree ferns, growing on a moderately steep slope. Geographical distribution.— Known from the type locality only. Chorological affinities.—This species is geographically isolated from all other species of Cyrtolaus. The geographically closest species is C. newton. Phylogenetic relationships. — Probably this species is related to C. newtoni and C. whiteheadi. A more detailed resolution can- not be offered now. Cyrtolaus whiteheadi, NEw SPECIES (Figs. 2, 5A, B) Type material. -HOLOTYPE female and PARATYPE female, each labelled: MEX. Oaxaca/ Rio Santiago, 11.7/ km. e. Huautla de/ Jiminez 1150 m,/ cloud forest/ July 13, VOLUME 93, NUMBER 2 1975; MEX. EXP. 1975/ G.E. Ball &/ H.E. Frania/ collectors. Holotype in USNM and paratype in UASM. Type locality.—México, Oaxaca, Sierra Madre de Oaxaca, 11.7 km E. Huautla de Jiminez. Derivation of specific epithet. — Latinized genitive form of the surname of the late Donald R. Whitehead, in whose honor and memory this species is named. Recognition. — Adults of this species have a single pair of supraorbital setae, a feature distinguishing them from all other species of Cyrtolaus, except C. furculifer Bates and C. grumufer Whitehead and Ball. Adults of the latter 2 species have the apical portions of one or more elytral intervals elevated (cf. Whitehead and Ball 1975: 600, Figs. 12, 13A, B), whereas in C. whiteheadi, the api- cal portion of interval 2 is not more convex than is the basal portion of this interval (Figs. 5A, B). Description. — With features of subgenus Cyrtolaus and following. See Tables 1-3 for data about variation in Standardized Body Length and in values for ratios ML/PL and REVEL. Head, antennae and mouthparts average for subgenus Cyrtolaus. Pronotum. Form as in Fig. 2, anterior angles obtuse, narrow, protruded slightly; posterior angles subspinose, protruded pos- teriorly; lateral areas elevated, more broadly so posteriorly; lateral margins slightly sin- uate posteriorly. Elytra. In form, as in Figs. 5A, B, vaulted. Humerus narrowly rounded; lateral margin sinuate preapically, apex with short spine about opposite termination of interval 2. Intervals slightly convex; interneurs finely and sparsely punctate on disc, punctures larger on apical declivity. Legs. Average for subgenus Cyrtolaus. Habitat.—The type material was found in damp leaf litter on a gentle slope in cloud forest. Geographical distribution.— Known from the type locality only. 237 Chorological affinities.—The locality for this species is the northernmost for the sub- genus Cyrtolaus. It is 80 km west of the locality of the next nearest species (C. new- toni) and is on the western rim of the Rio Grande basin, which is probably one of the major barriers for montane taxa in eastern México. Phylogenetic relationships. — Evidently, this species is related to C. newtoni and C. oaxacanus, based on the derived form of the posterior angles of the pronotum, and on geographical proximity but disjunct dis- tribution of the three species. Cyrtolaus subiridescens Whitehead and Ball (Figs. 3, 6A, B) Material examined.—Sixteen specimens, as follows. All localities are in México, in the state of Chiapas, on the Mesa Central. Municipio Mitontik, 11 km N. Tzonte- huitz, on road to Pueblo Viejo, 1981 m, X.22.1976 (7 males, 3 females—CASC). Municipio Tenejapa, Tenejapa Center, 2042 m, III.15.1975 (male—CASC). Municipio Las Margaritas, 48 km NE Las Margaritas on road to Campo Alegre, 2134 m, X.25.1976 (2 males, 2 females—CASC). Municipio Las Rosas, Villa Las Rosas, 1371 m, X.30.1976 (female —CASC). Taxonomic notes.—In body size (SBL) and in proportions (ML/PL and PL/EL) all of the specimens are within the ranges re- ported previously (Tables 1-3; cf. White- head and Ball 1975: Tables 1-3). The ma- terial from Mitontik and Tenejapa is typical in all respects, as determined by comparison with paratypes of C. subiridescens. The specimens collected at Las Margar- itas and Villa Las Rosas, however, are dis- tinctive in having fairly long apical elytral spines (Figs. 6A, B). In pronotal form (Fig. 3) and in details of the male genitalia, these specimens are typical of C. subiridescens. The difference in elytral spination, in the genus as a whole, is important phylogenet- ically, and one might be tempted on this basis to propose taxonomic recognition, at 238 the subspecies level. The localities, how- ever, are within the known geographical range of C. subiridescens (Whitehead and Ball 1975: Fig. 23). Under this circum- stance, I doubt that the difference in elytral spination 1s indicative of anything more than minor intraspecific geographical variation. Cyrtolaus ricardo Whitehead and Ball (Figs. 4, 7A, B) Material examined.—Eleven specimens, as follows. The localities are in the Sierra Madre de Chiapas, on the Pacific versant of Middle America. MEXICO, Chiapas. Vol- can Tacana, NE slope, 1830 m, XII.20.1976 (seven males, female—UASM). GUATE- MALA. San Marcos Sivinal, cloud forest, 2621 m, XII.31.1976 (male, 2 females— UASM). Taxonomic notes.—Tables 1-3 provide data about variation in Standardized Body Length and in values for the ratios ML/PL and PL/EL. The specimens collected on Ta- cana are indistinguishable from the type se- ries, which was collected in the same general area. The specimens collected at San Mar- cos Sivinal, on the slopes of Volcan Taju- mulco, adjacent to and within a few hours walk from Tacana, are similar to the Tacana material in pronotal form (Fig. 4), and in details of the male genitalia (2 males dis- sected). In proportions, the 2 groups differ slightly in values for the ratio ML/PL (Table 2). Also, the apical spines of the elytra of the San Marcos specimens are shorter (Figs. TA, B). Elevation.—The new material collected on Volcan Tacana extends the known alti- tudinal range of C. ricardo downward by ca. 900 m. However, this species and C. fur- culifer, the only other species of Cyrtolaus inhabiting the slopes of this volcano, remain allopatric, though it seems likely that their ranges are (or were) in contact, at least mar- ginally. Conclusion.— The new material is within the general geographical range that we ascribed to C. ricardo (Whitehead and Ball PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1975: 610). The range of morphological variation is extended slightly, but not tax- onomically significantly so. Cyrtolaus furculifer Bates Material examined.— Five specimens, as follows. All were collected on the north- eastern slopes of the Volcan Tacana, in southeastern Chiapas, México. Union Juar- ez, Barranca Providencia, montane tropical forest, in leaf litter, 1500 m, XII.15.1975 (male and female, UASM). Union Juarez, Finca Muxbal, ex pile of mouldy wood, 1430 m, XII.26.1975 (2 females—UASM). Rio Muxbal, 1463 m, XII.21.1976 (female, UASM). Union Juarez, agricultural land, 1720 m, XII.14—26.1975 (set of elytra— UASM). Taxonomic notes. — The specimens noted above, most of them collected in or near the Barranca Providencia, from where White- head and Ball (1975: 611) reported this species, are typical members of C. furculifer, as shown by Standardized Body Length and proportions (Tables 1-3). The individual collected at 1720 m extends the known ver- tical range of this species upward by ca. 200 m, but still beneath the known altitudinal range of C. ricardo (see comments for that species, above). Cyrtolaus grumufer Whitehead and Ball Material examined.—One female, from Chiapas, México, in the Mesa Central: Cer- ro Baul, NW slope, west of Rizo de Oro, 1768 m, X.2.1979 (CASC). Taxonomic notes.—This specimen is within the morphological range of the type material (Whitehead and Ball 1975: 612) as shown by Standardized Body Length and body proportions (Tables 1-3). It is also within the reported vertical range (1524—- 1829 m), and from northwestern Chiapas. However, Cerro Baul is about 160 km from the only area reported previously for C. gru- mufer. This is a marked but not surprising range extension for this species. VOLUME 93, NUMBER 2 NOTES ABOUT PHYLOGENETIC ASPECTS A reconstructed phylogeny of the species of Cyrtolaus was offered by Whitehead and Ball (1975: 615, Fig. 25). The new data sug- gest that the Oaxacan species (C. oaxacan- us, C. newtoni, and C. whiteheadi) form a clade apart from a Chiapan-Guatemalan complex (C. brevispina, C. ricardo, C. spini- cauda, C. furculifer, and C. grumufer). The longer apical elytral spines reported for 2 samples of C. subiridescens render equivocal the position of this taxon. If these spines were developed independently of the Chiapan-Guatemalan complex noted above, C. subiridescens could continue to occupy a relatively basal position, as postulated pre- viously by Whitehead and Ball (1975). If, however, the longer spines are indicative of relationship (and their absence from most specimens of C. subiridescens a loss), then this species could be placed with the Chia- pan-Guatemalan complex. I favor the latter alternative on chorological grounds. This resolution postulates a monophyletic Oaxa- can assemblage and a monophyletic Chia- pan-Guatemalan assemblage, separated from one another by the Isthmus of Te- huantepec. In part, such a hypothesis depends upon the relationships of the Oaxacan species of Cyrtolaus. Until males are known for C. newtoni and C. whiteheadi, their relation- ships to one another and to C. oaxacanus will remain equivocal. Consequently, it seems premature to pursue this topic further at this time. ACKNOWLEDGMENTS I take pleasure in thanking D. H. Kava- naugh, Department of Entomology, Cali- fornia Academy of Sciences, for making available for study material of Cyrtolaus that was collected in Chiapas by Dennis E. 239 Breedlove. It is a distinct pleasure, also, to acknowledge the continuing efforts of Dr. Breedlove, a distinguished systematist in the Academy’s Department of Botany, to en- rich the entomological holdings of his in- stitution. Finding specimens of Cyrtolaus is both a challenge and a delight. I am grateful to H. E. Frania and D. Shpeley, who accompanied me on the trips when the new Oaxacan species of Cyrtolaus were taken, and who collected the specimens of C. whiteheadi and C. oaxacanus, respectively. I thank also Henry Frania, J. Belicek, and D. R. Procter for their efforts to obtain material of Cyr- tolaus during Christmas time sojourns on the chilly slopes of Volcan Tacana. The careful illustrations for this contri- bution were prepared by Diane Hollingdale. D. Shpeley, Assistant Curator of the Strick- land Museum, provided the curatorial as- sistance on which I depend. Funding for field and museum work and for publication costs was provided by Grant A- 1399, Natural Sciences and Engineering Research Council of Canada. LITERATURE CITED Ball, G. E.and R. E. Roughley. 1982. The Hypherpes- like taxa of southern Mexico: Classification, and evolutionary considerations (Coleoptera: Carabi- dae: Pterostichus). Transactions of the American Entomological Society 108: 315-399. Bates, H. W. 1882. Insecta, Coleoptera, Carabidae, Vol. 1, Part 1, pp. 40-152, plates iii—v. In God- man, F. D., and O. Salvin, eds., Biologia Centrali- Americana. 1879-1911. Coleoptera, 7 volumes in 17 parts. London. Liebherr, J. K. 1986. Barylaus, new genus (Coleop- tera: Carabidae) endemic to the West Indies with Old World affinities. Journal of the New York Entomological Society 94: 83-97. Whitehead, D. R. and G. E. Ball. 1975. Classification of the Middle American genus Cyrtolaus Bates (Coleoptera: Carabidae: Pterostichini). Quaes- tiones Entomologicae 11: 591-619. PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 240-243 THE “MISSING” FEMALES OF ALBERTISIUS GESTROI (PASCOE) (COLEOPTERA: CURCULIONIDAE), AN ENIGMA OF NATURAL HISTORY, WITH A NOTE ON DR H. JAMES RICHARD T. THOMPSON Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, England. Abstract. —The hitherto unknown female of Albertisius gestroi (Pascoe) (Curculionidae: Entiminae, Celeuthetini) is described and figured and its genitalia compared with those of A. excellens (Faust). Attempts to account for the apparent dearth of females in A. gestroi are followed by a note on Dr. H. James, an American who collected one of the specimens described. Key Words: The New Guinea weevil genus 4/bertisius contains two species, A. gestroi (Pascoe) known only from Yule I. and A. excellens (Faust) which occurs around Port Moresby. When A/bertisius was described (Thompson 1977: 253) both sexes of A. excellens were known (from at least 12 males and 9 fe- males) whereas A. gestroi was known only from 26 males, nearly all collected by the Italian explorer L. M. D’Albertis. Later, a single female was found among undeter- mined material in the Natural History Mu- seum and a check of the Genoa Museum’s 27 D’Albertis specimens by the curator, Dr. Poggi, revealed two more. The sex ratio for the available specimens is thus 51:3 or 17:1 in favour of males. The newly discovered female of A. gestroi is described below. Albertisius gestroi (Pascoe), female Apirocalus gestroi Pascoe 1885: 209; pl. 1, Fig. 3. Heteroglymma gestroi (Pascoe); Marshall 1938: 96. Curculionidae, sex ratio, genitalia, New Guinea, explorers Apirocalus gestroi Pascoe; Marshall 1956: es Albertisius gestroi (Pascoe); Thompson 1977: 253) Differs from male (Fig. 1) as follows: el- ytra with dorso-lateral carinae reduced but sides strongly angulate two-thirds of length from base (Fig. 2); covered with stiff erect brown setae. Legs with all femora less swol- len; hind tibiae less swollen but with similar dense brush of semi-recumbent setae on in- ner aspect. Female genitalia (Fig. 6) similar to those of A. excellens (Thompson 1977: fig. 23). Spermatheca (Fig. 5) clearly of same very distinctive type as that of A. excellens (Figs. 7, 8) but gland-lobe free (not fused laterally with body) and duct-lobe much shorter. Material examined.—1 2, with ‘N. Guin- ea/ Isola Yule/ T.F. V.1875/ L. M. D’AI- bertis’ (mostly printed) (Genoa Museum); 12, with ‘New/ Guinea/ [B.M. 18]76.32’ (handwritten, with orange line)(Natural History Museum, London). The register en- VOLUME 93, NUMBER 2 241 Figs. 1-4. Albertisius species. 1, A. gestroi (Pascoe), male. 2, idem, female. 3, A. excellens (Faust), male. 4, idem, female. Scale-line = 2.0 mm. try for this specimen reads: “‘Purchased of Mr Higgins 20/10/76. Collected by Dr James.” The second Genoa specimen was not ex- amined. Remarks. —Albertisius gestroi is a fairly typical celeuthetine weevil, apterous and showing moderate sexual dimorphism (mainly affecting the shape of the elytra). The most likely explanation for a predom- 242 Figs. 5-8. 8, A. excellens (Faust), spermathecae. Scale-lines = 0.2 mm (Figs. 5, 7 and 8) and 2.0 mm (Fig. 6). inance of males would be asynchronous emergence of the sexes but D’Albertis was on the island from March to October and moreover collected some 250 specimens of a related species (Apirocalus cornutus Pas- coe) in April, May and June while his 4. gestroi specimens were all taken in May. From his journal (D’Albertis 1880) it is known that his movements, even within the island, were restricted so it is possible that the specimens were collected by one of the natives (that he records having employed to collect for him) from a site to which he him- self did not have access. It is worth noting that James’s specimen must have been taken between October and January, the wet sea- son, a time of year which D’Albertis did not spend on the island. The few other records of A. gestroi PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bs Albertisius species, genitalia. 5, A. gestroi (Pascoe), spermatheca. 6, idem, female genitalia. 7 and (Thompson 1977: 256) are undated but none is recent. As Yule I. is only about five miles long and much visited, it may well be that the species is extinct and the mystery of its anomalous sex ratio will never be solved. HISTORICAL NOTE Little is known of Dr. H. James who col- lected the Natural History Museum female of A. gestroi (Fig. 2). He was a young Amer- ican recruited by William Macleay as sur- geon on the barque Chevert which was used by Macleay for his New Guinea expedition of 1875. He was described as “‘of pleasant manners” (D’Albertis 1880, 1: 363) and“... had been instructed in the art of bird skin- ning and was a first-rate shot” (Goode 1977: 106). He first met D’Albertis on Yule I. on 30 August 1875 when the expedition stopped VOLUME 93, NUMBER 2 there on its way to the New Guinea main- land and again in late October on his way back to Port Moresby on the missionary supply steamer E//engowan. He returned to Yule I. with a botanical collector named Knight on 6 November “‘to set up a house and make collections” (D’Albertis 1880, 1: 387). On the same visit the E//engowan took D’Albertis off, so James and Knight were the only Europeans on the island. After three months both were so ill they returned to Somerset (Cape York) on the E//lengowan. James returned to Yule I. yet again on 23 August 1876 with a Swedish trading partner but both were unexpectedly murdered by natives upon their arrival. Some of the beetles James collected (in- cluding the A. gestroi female) were sent to E. T. Higgins, a collector and dealer, who sold them to the Natural History Museum on 20 October 1876 by which time James was dead. A cetoniine beetle, Jschiopsopha jJamesii (Waterhouse), from the same lot of specimens, was named after him “‘in accor- dance with a wish expressed by Mr Higgins” (Waterhouse 1876: 473). ACKNOWLEDGMENTS I am most grateful to Dr. R. B. Madge (International Institute of Entomology) for finding the book by Goode, with its account of Dr. James and to Dr. R. Poggi of the 243 Museo Civico di Storia Naturale ““Giacomo Doria,”’ Genoa, Italy for his help with ma- terial and information. Figs. 1, 7 and 8 are reproduced from Thompson (1977) by permission of the Trustees of the Natural History Museum; Figs. 1—4 were prepared by the Natural His- tory Museum Photographic Unit. LITERATURE CITED D’Albertis, L. M. 1880. New Guinea: What I Did and What I Saw. 2 vols. Sampson Low, Marston, Searle and Rivington, London. x + 424 and x + 406 pp. Goode, J. 1977. Rape of the Fly. Thomas Nelson (Australia) Limited, Melbourne. xi + 272 pp. Marshall, Sir Guy A. K. 1938. On Elytrurus (Col. Curcul.) and an allied new genus. Transactions of the Royal Entomological Society of London 87: 69-101. . 1956. The Otiorrhynchine Curculionidae of the Tribe Celeuthetini (Col.). British Museum, London. 134 pp. Pascoe, F. P. 1885. List of the Curculionidae of the Malay Archipelago collected by Dr Odoardo Bec- cari, L. M. D’Albertis, and others. Annali del Mu- seo Civico di Storia Naturale di Genova (2)2: 201- B38 Thompson, R. T. 1977. A revision of the New Guin- ea weevil genus Apirocalus Pascoe (Coleoptera: Curculionidae). Bulletin of the British Museum (Natural History) (Entomology) 36: 193-280. Waterhouse, C. O. 1876. Descriptions of two new species of Cetoniidae. Annals and Magazine of Natural History (4)18: 422-423. PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 244-247 A FOURTH GENUS OF SMALL-BODIED XYSTODESMID MILLIPEDS FROM THE SOUTHEASTERN COASTAL PLAIN (POLYDESMIDA) ROWLAND M. SHELLEY North Carolina State Museum of Natural Sciences, Raleigh, North Carolina 27611. Abstract. —The minute xystodesmid milliped, Lourdesia minuscula, n. gen., n. sp., 1s described from the Coastal Plain of southern Alabama. The principal diagnostic features are ridges on the sides of the metazonites near the pleural/sternal junctures, the absence of a cyphopod receptacle in females, and a minute, simple gonopod telopodite that lacks a prefemoral process. Since the milliped also lacks sternal hairs and lobes, and the sternal remnant between the gonopodal coxae, it cannot be assigned to the Rhysodesmini, and its tribal position is uncertain. Key Words: The Coastal Plain of the southeastern United States harbors a diverse xystodes- mid diplopod fauna, among which are three endemic small-bodied genera tentatively assigned to the Rhysodesmini— Caralinda, Parvulodesmus, and Gonoessa—with four, one, and five species, respectively (Hoffman 1978, Shelley 1979, 1983a, b, 1984). These millipeds are substantially smaller than sympatric forms of Sigmoria, Dicellarius, and Pachydesmus, which range from two to seven times as large, and hence are a dis- tinctive component of the regional fauna. Caralinda, with moderately long, complex gonopods, occurs in the contiguous sections of Alabama, Georgia, and Florida. Parvu- lodesmus and Gonoessa have extremely long gonopods, overlapping three segments an- terior to the 7th, and may represent a single genus. The former is known only from Ab- beville County, South Carolina, and the lat- ter occurs broadly across southern Ala- bama. While recently sorting specimens loaned by the Florida State Collection of Arthropods, Gainesville (FSCA), I discov- ered three samples of a minute, undescribed xystodesmid with very small gonopods that Lourdesia, Rhysodesmini, Xystodesmidae clearly requires a new genus and possibly also a new tribe. It lacks such traditional rhysodesmine features as sternal hairs and a sternal remnant between the gonopods, but the species also does not fit in the en- demic east Nearctic tribes (Apheloriini, Nannariini, and Pachydesmini). It attests to a substantial fauna of small-bodied xysto- desmids in the southeast, and one or more new tribes may be warranted when more material is available and the diversity of these forms is better known. These small millipeds are prevalent in cool weather. Most have been taken from November—March, and they are poorly represented in collec- tions because little field activity tradition- ally occurs during this time of year. Con- sequently, anyone conducting winter field trips to the southeast, particularly in the Florida panhandle and southern Alabama and Georgia, may find his efforts rewarded with the unveiling of a largely unknown dip- lopod fauna. Two other small xystodes- mids, definite members of the Rhysodes- mini, inhabit this region—Pleuroloma pinicola Shelley, in southeastern North Car- olina and coastal South Carolina, and P. VOLUME 93, NUMBER 2 cala (Chamberlin), in peninsular Florida. However, their congeners—P. flavipes Ra- finesque, occurring across the northern, cen- tral, and midwestern states, and P. plana Shelley, ranging from the Florida panhandle to central South Carolina—are considerably larger, so small size is not a feature of this genus (Shelley 1980). Lourdesia, NEw GENUS Type species. — Lourdesia minuscula, new species. Description.—A genus of small-bodied rhysodesmine xystodesmids with the fol- lowing characteristics: Body composed of head and 20 segments in both sexes; adults ranging from 12-16 mm long and 2-3 mm wide, W/L ratio from 17-18%. Head of normal appearance, smooth; epicranial suture faint, not bifid; facial setae reduced, epicranial, interanten- nal, and frontal absent. Antennae relatively short, with 4 small apical sensory cones, no other sensory structures apparent. Terga smooth, polished; strictures dis- tinct, impressed. Collum broad. Paranota strongly depressed; peritremata distinct, ozopores opening laterad. Sides of meta- zonites with variably elevated ridges just below coxae at pleural/sternal junctures. All sterna of males and females glabrous and essentially flat and plate-like, at most with only very slight indentations, without lobes or spiniform projections from caudal margins. Gonapophyses moderately long, apically expanded. Coxae and prefemora without tubercles or spines. Gonopodal aperture relatively small, ovoid. Gonopods with very short telopo- dites, lying nearly entirely over aperture, only slightly overhanging anterior margin. Coxae large, without apophyses, with clus- ters of 6-8 setae lateral to cannula, con- nected by membrane only, no detectable sclerotized sternal remnant; apodemes rel- atively long and narrow. Prefemur short, without prefemoral process. Acropodite not divided, not demarcated from prefemur, 245 short and blade-like, curving bisinuately and bent mediad near midlength, tapering throughout length, without lobes or projec- tions. Cyphopodal aperture narrow, encircling 2nd legs. Cyphopods relatively large. Re- ceptacle absent. Valves large, subequal. Operculum large, located laterad to valves. Species. — One is known; others probably await discovery in southern Alabama and the Florida panhandle. Remarks. — Though it shares the absence of the cyphopodal receptacle with Cherokia (Hoffman 1960) and that of a gonopodal prefemoral process with three species of Gonoessa (Shelley 1984), Lourdesia cannot be assigned to the Rhysodesmini as cur- rently understood because of the absence of a sternal remnant between the gonopods, sternal hairs, particularly on postgonopodal sterna of males, and lobes or projections from the caudal sternal margins. With short, simple gonopodal telopodites that are un- divided and lack flanges, lobes, and lamel- lae, Lourdesia also does not seem closely related to either Caralinda or Gonoessa. It shares small size, glabrous sterna, and lat- eral metazonal ridges with Gyalostethus, a primarily montane genus that ranges east- ward onto the Piedmont Plateau and west- ward onto the Ridge and Valley, Appala- chian Plateaus, and Interior Low Plateaus physiographic provinces. Hoffman (1965) assigned Gyalostethus to the Rhysodesmini because of its acicular prefemoral process and the right angle between the coxa and telopodite; however it also apparently lacks the sternal remnant between the coxae. The small-bodied southeastern xystodesmids thus present major difficulties in tribal placement. They are even anomalous in the Rhysodesmini, the only established tribe that can accommodate them, and the enig- matic status of Lourdesia is underscored by the absence of ventrodistal spines on the ambulatory prefemora. In this regard, it re- sembles western Nearctic genera of the tribe Chonaphini. 246 Lourdesia minuscula Shelley, NEw SPECIES Figs. 1-4 Type specimens.— Male holotype and 1 male, 5 female, and 3 juvenile paratypes (FSCA) collected by N. B. Causey, 26 Jan- uary 1965, along U.S. highway 84, 5.5 mi. E Elba, Coffee Co., AL. Diagnosis.—With the characters of the genus. Color in life. —Unknown. All specimens completely blanched in preservative, with- out trace of pigmentation pattern. Holotype.—Length approximately 12.2 mm, maximum width 2.1 mm, W/L ratio 17.2%, depth/width ratio 90.0%. Head capsule smooth, polished; epicra- nial suture faint, terminating in interanten- nal region. Antennae reaching back to just beyond caudal margin of 2nd tergite, be- coming progressively more hirsute distally; first antennomere subglobose, 2-6 clavate, 7 short and truncate, relative lengths of an- tennomeres 6>5>4>2>3>1>7. Genae not margined laterally, with faint medial impressions, ends broadly rounded and projecting slightly beyond adjacent cranial margins. Facial setae as follows: epicranial, interantennal, and frontal absent, genal 2-2, clypeal about 9-9, labral about 14-14. Collum broad, ends extending slightly be- yond those of following tergite. Paranota angled sharply ventrad and continuing slope of dorsum; anterior corners rounded, cau- dolateral corners blunt on all segments. Peritremata relatively broad, strongly ele- vated above paranotal surface; ozopores lo- cated just caudal to midlength of peritre- mata, opening laterad. Sides of metazonites smooth, polished, with elevated ridge varying from slightly notched or scalloped to smooth and slightly longer caudad (Fig. 1). Strictures distinct ventrally. Sterna of segments 4—6 glabrous, without distinct modifications, with only slight, barely perceptible concavities. Post- gonopodal sterna also glabrous, flat and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON | | \| 4 ma J x (, th 4 / | a\ arAN \ \ \ A, | ; wee eases g : / i) 2. '3 Figs. 1-4. Lourdesia minuscula. 1-3, holotype. 1, caudal view of 7th segment showing profiles of ridges on sides of metazonites. 2, left gonopod, medial view. 3, the same, lateral view. 4, left cyphopod of female paratype, anterior view. Setation is omitted from all drawings. Scale line = 0.5 mm for Figs. 2, 3, 1.9 mm for Figs. 1, 4. plate-like, without grooves, elevations, or projections from caudal margins. Coxae and prefemora without projections; tarsal claws of normal length on all legs, variably hooked to sublinear. Hypoproct broadly rounded; paraprocts with margins strongly thickened. Gonopodal aperture ovoid, without in- dentations, margins not thickened, caudal margin and caudal halves of sides slightly elevated above metazonal surface. Gono- pods in situ with coxae protruding through aperture, telopodites projecting ventrad and curving toward midline, apices overlapping and extending slightly beyond anterior mar- gin of aperture. Gonopod structure as fol- lows (Figs. 2, 3): Coxae closely appressed, filling nearly entire aperture opening. Pre- femur lightly hirsute, without prefemoral process. Acropodite relatively short, con- tinuous with prefemur, in form of progres- sively narrowing, undivided blade, curving gently and bisinuately, bending strongly mediad distal to midlength and curving bi- sinuately to subacuminate tip. Prostatic groove arising in pit in base of prefemur, VOLUME 93, NUMBER 2 running along inner surface of acropodite to terminal opening. Male paratype.—The male paratype agrees with the holotype in all particulars. Female paratypes.—Length approxi- mately 15.9 mm, maximum width 2.8 mm, W/L ratio 17.6%, depth/width ratio 75.0%. Agreeing essentially with males in all struc- tural features; paranota only slightly more strongly depressed; ridges on sides of meta- zonites smaller and more rounded. Cypho- pods in situ with valves projecting through aperture. Valves (Fig. 4) subequal, relatively large, oriented transversely. Receptacle ab- sent. Operculum relatively large, located laterad below free end of valves. Variation.—The Baldwin County male agrees closely with the holotype. Ecology.—The only habitat information on the vial labels is the notation “bluff mixed woods” with the Baldwin County male. Pre- sumably this refers to a mixed pine/hard- wood association. I also found light-colored debris in the guts of two individuals, sug- gesting a sandy substrate. Both samples were collected in January, indicating a preference for cool weather. Distribution.— Known only from the type locality and the following site, also in south- ern Alabama, which straddle the Florida panhandle suggesting probable occurrence between Pensacola and DeFuniak Springs, Florida. ALABAMA. Baldwin Co., ca. 10 mi W Loxley, near junction of US highways 90/ 98, M, 22 January 1965, N. B. Causey (FSCA). Remarks.—Lourdesia minuscula and Parvulodesmus prolixogonus Shelley are the smallest xystodesmids in the eastern Ne- arctic faunal region. With its narrow width and distinct peritremata, L. minuscula ap- pears more like a large paradoxosomatid, for example like a large Oxidus gracilis (C. L. Koch), than a xystodesmid. It clearly is 247 not a paradoxosomatid, lacking the dorsal metazonal grooves and the characteristic se- tal arrangement on the paraprocts, but its general facies differs markedly from those of other eastern xystodesmids. ACKNOWLEDGMENTS Special thanks are extended to G. B. Ed- wards, FSCA, for loan of the milliped ship- ment containing this new genus, and to R. L. Hoffman for advice on its systematic po- sition. I also thank R. E. Gordon for the invitation to publish a paper in this me- morial issue for Don Whitehead, whose dis- cerning mind and keen insight greatly en- hanced my understanding of milliped evolution and relationships within the Xys- todesmidae. LITERATURE CITED Hoffman, Richard L. 1960. Revision of the milliped genus Cherokia (Polydesmida: Xystodesmidae). Proceedings of the United States National Mu- seum 112: 227-264. 1965. Revision of the milliped genera Bo- raria and Gyalostethus (Polydesmida: Xystodes- midae). Proceedings of the United States National Museum 117: 305-348. . 1978. A new genus and species of rhysodes- mine milliped from southern Georgia (Polydes- mida: Xystodesmidae). Proceedings of the Bio- logical Society of Washington 91: 365-373. Shelley, Rowland M. 1979. A new milliped of the genus Caralinda from north Florida (Polydesmi- da: Xystodesmidae). Florida Entomologist 62: 183- 187. 1980. Revision of the milliped genus Pleu- roloma (Polydesmida: Xystodesmidae). Canadian Journal of Zoology 58: 129-168. . 1983a. Parvulodesmus prolixogonus, n. gen., n. sp., a new xystodesmid milliped from South Carolina (Polydesmida). Proceedings of the Bio- logical Society of Washington 96: 121-126. . 1983b. New records and species of the mil- liped genus Caralinda (Polydesmida: Xystodes- midae). Florida Entomologist 66: 407-415. . 1984. Anew xystodesmid milliped genus and five new species from the Coastal Plain of Ala- bama (Polydesmida). Florida Entomologist 67: 453-464. PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 248-261 THE FLOWER FLY GENUS ORNIDIA (DIPTERA: SYRPHIDAE) F. CHRISTIAN THOMPSON Systematic Entomology Laboratory, ARS, USDA, NHB-168, Smithsonian Institution, Washington, D.C. 20560. Abstract.—The flower fly genus Ornidia (Diptera: Syrphidae) is revised. The genus is redescribed; a key to species is presented; the phylogenetic relationships of the genus and species are hypothesized; the included species are redescribed; with a new species, white- headi, described from Panama (type) and Colombia; and the critical characters are illus- trated. Key Words: Ornidia is small group of brilliant metal- lic green or purple flies found mainly in the New World tropics: Anyone who has spent time in these tropics knows these beautiful flies as they are common around human habitations. The genus contains only 4 spe- cies: One common species found every- where in the New World tropics and that has spread extensively through the Old World tropics during the last century with commerce, two others which are less com- mon but widespread, and one new species, presently known from a few specimens from Panama and Colombia. This paper presents a revision of the genus, with complete syn- onymies (except only major references giv- en for obesa), descriptions, and distribu- tional and biological data for all taxa. Genus Ornidia Lepeletier and Serville Ornidia Lepeletier & Serville, 1828: 786. Type-species, Syrphus obesus Fabricius (orig. des.). Curran 1930: 2 (key); Val 1972 (key, biometry, evolution); Thompson 1972: 106 (descr., relationships). Volucella, subg. Ornidia: Hull 1949: 348 (description); Hardy 1964: 403 (descrip- tion). Head: face concave beneath antenna, key, phylogenetic relationships, neotropical straight below median tubercle, with dis- tinct median tubercle and smaller lateral tu- bercle; frontal prominence distinct, low, above middle of head; frontal triangle short, ’y as long as eye contiguity, slightly puffed out; front of female narrow, about twice as long as wide at antennal base, as long as face, with convergent sides dorsally, puffed out, with a transverse depression about '4 of frontal length above antenna, with short sublateral longitudinal depression extend- ing about '4 the length of front above trans- verse depression. Eye pilose, holoptic in male. Antenna short, shorter than face; ba- soflagellomere elongate, twice as long as broad at base; arista plumose, as long as antenna. Thorax: about as long as broad; noto- pleuron enlarged, swollen and elongate pos- teriorly on lateral ‘2; meso-anepisternum with anterior portion bare; meso-katepister- num with posterior ’2 completely pilose; meso-anepimeron with posterior portion pilose; meropleuron with barrette pilose; with a patch of pile anterobasal to meta- thoracic spiracle; scutellum with a pre-api- cal depression, without ventral pile fringe. Wing: without microtrichia; marginal cell closed, petiolate, with apical portion an- VOLUME 93, NUMBER 2 gulate posteriorly; apical crossvein reces- sive. Abdomen: suboval, convex, without bris- tles. Ornidia belongs to the tribe Volucellini and is the sister of Copesty/um [Volucellini = Graptomyza + (Volucella + (Ornidia + Copestylum)), see Thompson and White- head (1986) for explanation of cladistic for- mulae] (Thompson 1972). The genus is de- fined (synapomorphy) by its facial and notopleural structure, the arrangement of a large medial and smaller sublateral tuber- cles on the face (Figs. 1-3), and the enlarged, posteriorly produced notopleuron (Fig. 4), states which are unique among flower flies. The cladistic relationships are derived from the following characters (see Table 1), with the polarity determined by outgroup com- parison (tribe Rhinigiini, genus Ferdinan- dea): 1) Male eyes holoptic (0) or dichoptic (1). This character varies within the genus Co- pestylum, one species group has dichoptic eyes, the rest holoptic, which is accepted as the ground plan condition. 2) Arista bare (0), sparsely and short pi- lose (1), or plumose (densely and long pi- lose) (2). The aristal pilosity varies greatly in both Graptomyza and Copestylum. In Graptomyza most species have the arista sparsely pilose with short hairs, which I ac- cept as the ground plan condition (as inclu- sa), some species have only a few very short aristal hairs, one undescribed species from Australia has no aristal hairs, and some spe- cies have long and numerous hairs (as in longirostris). In Copestylum, there are a few species groups with distinctive aristal pi- losity, but these are all clearly derived from the basic plumose condition. One species, pseudotachina, has the arista bare, a sec- ondary reduction. 3) Meso-anepisternum bare (0) or pilose (i): 4) Meso-katepisternum continuously pi- lose (0) or bare medially (1). 249 5) Meso-anepimeron bare posteriorly (0) or pilose (1). 6) Meso-katepimeron (barrette) bare (0) or pilose (1). 7) Postalar wall bare (0) or pilose poste- riorly (1). 8) Scutellum without (0) or with pre-api- cal depression (1). 9) Medius with apical portion (apical crossvein) processive (0), arcuate and strongly recurrent (1) or straight (2). 10) Cell R4+5 (apical cell) open (0), pet- iolate (1) or bulbous apically (2). Within Copestylum, the apical cell varies from widely open, to petiolate and bulbous. I ac- cept the widely open state as the ground plan condition for Copestylum. 11) M2 present (0) or absent (1). 12) Larvae saprophagous (0) or special- ized inquilines in nests of social Hymenop- tera: (1): The taxa examined were: Copestylum (apicalis Loew, compactus Curran, fornax Townsend, gibbera Schiner, hirtipes Macquart, marginatum Say, mexi- canum Macquart, trituberculatum Thomp- son, tympanitis Fabricius); Ferdinandea (cupreus Scopol1); Graptomyza (alabeta Mutin, doddi Fer- guson, flavicollis Ferguson, inclusa Walker, liberia Greene, longirostris Wiedemann, maculipennis de Meijere, microdon Osten Sacken, nigripes Brunetti, plumifer Fergu- son, signata Walker); Ornidia (all species); Volucella (bombylans Linnaeus, decolo- rata Walker, elegans Loew, inanis Linnae- us, inflata Fabricius, jeddona Bigot, linearis Walker, nigricans Coquillett, pellucens Lin- naeus, rotundata Edwards, tabanoides Mot- schulsky, trifasciata Wiedemann, and zo- naria Poda). I now accept the diphyletic origin of the Old World volucellines as Graptomyza are now known to be saprophagous, not spe- cialized inquilines. Also, Tachinosyrphus is no longer accepted as a genus distinct from Copestylum (new synonymy) as that ar- 250 6 Figs. 1-8. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mesonotum 1st 2nd sternum Features of Ornidia. 1, 2. Facial profiles. 1, aemula Williston. 2, major Curran. 3. obesa Fabricius, head, lateral. 4. obesa Fabricius, notopleuron and adjacent structures, dorsal view. 5, 6. Wing, apical half. 5, aemula Williston. 6, major Curran. 7, 8. Abdominal sternum, lateral profile. 7, major Curran. 8, aemula Williston. rangement left Copestylum as undefined. The above cladistic formula, which is the same as my 1972 phylogeny, is derived when the type species are used as exemplars (CI = 87, RI = 71, | tree). The possibility that Ornidia is a specialized subgroup of Cope- stylum rests on the evaluation of one char- acter, the pilosity of the posterior portion of the anepimeron. In my previous analysis, I considered the pilose condition to be prim- itive in all situations. Hence, Copestylum was defined by a bare posterior anepimeron. However, under a strict outgroup criterion, the bare condition is primitive within the VOLUME 93, NUMBER 2 Table 1. Characters of volucelline taxa. 251 to oe) Taxa a Characters 6 7 n oo ve) oO i) Ferdinandea cuprea Graptomyza inclusa Graptomyza longiventris Volucella pellucens Ornidia obesa Copestylum marginatum Copestylum mexicanum Copestylum nasicum Copestylum tympanitis Copestylum apicale Copestylum pseudotachina Copestylum trituberculatum Copestylum fornax Copestylum hirtipes Copestylum gibbera Copestylum compactus Copestylum ground plan NNNNKFNNNNNNNNNN — © SSeS} (SS COO SS) OS] LDS) SOL OnOeoro ro Ore) Grutor SS SOOO Or OrOrO Or Ory Or Oi NN © SOTO. OO O19 OO Ore ORS | OrOre) — Sooo =O OO Oo OO @ ee o.9o Oo © OOH = O'O © = Ot =" SE ee ea ee ee ONKONORrF OR RF KFNNRK OOO eee ee ee ee eee EE OOO oO O19 Oo © OC: O:. Oo 1C 'O = Oo O'S See text for descriptions of the characters. tribe Volucellini. Copesty/um is a large ge- nus (350+ species) with many distinct groups. When exemplars of these groups are added to the analysis and strict outgroup criterion is used, a far different statement of relationship results (Volucellini = Grap- tomyza + (Volucella + Ornidia + Cope- stylum), consensus of some 100 trees). Thus, until Copestylum can be revised, the rela- tionships of Ornidia will remain uncertain. Ornidia is an endemic New World group consisting of 4 known species. One species (obesa), however, is hemisynanthropic and has spread extensively in the Pacific and across the Orient to the east coast of Africa. The cladistic relationships among the spe- cies (Ornidia = aemula + (obesa + (major + whiteheadi)))are tentative as they rest on only a few characters (size, scutellar de- pression, male genitalia), and without knowing the outgroup the polarity of these characters is uncertain. KEY TO THE SPECIES OF ORNIDIA 1. Apical wing spot large (Fig. 5); 2nd sternum with basomedial digitate process (Fig. 8). Mesonotum and 3rd tergum black pilose; oc- Apical wing spot small (Fig. 6); 2nd sternum WITHOUT PILOCESSH (10h) meter tes cyeteicusie the eile 2. Mesonotum and 3rd tergum extensively pale pilose; occiput shiny. Scutellar depression di- vided medially whiteheadi Thompson Mesonotum and 3rd tergum black pilose; oc- ciput extensively white pollinose 3. Scutellar depression divided medially; with prescutellar bristles major Curran Scutellar depression continuous medially; without prescutellar bristles obesa Fabricius Ornidia aemula Williston Figs. i, 5.18, 9s Volucella aemula Williston 1888: 272. Type- locality: Bolivia, Santa Cruz, Piedra Blan- ca [4 km west of Corumba, Mato Grosso, Brazil]. Lectotype 2 AMNH here desig- nated. Kertesz 1910: 184 (catalog cita- tion); Fluke 1957: 58 (catalog citation). Ornidia aemula: Curran 1930: 2 (key ref- erence); Doesburg 1963: 17, 1966: 96 (Su- rinam); Val 1972 (biometry, distribution, male genitalia figured); Baez 1985: 74 (Venezuela). Head: dark purplish black; face shiny, 252 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9-12. Male genitalia of Ornidia, lateral view. 9, aemula Williston. 10, obesa Fabricius. 11, major Curran. 12, whiteheadi Thompson. VOLUME 93, NUMBER 2 253 L oa i: SS TAK iS EL gpil\\ fa) GY i) WAN SS ale . Figs. 13-16. Male genitalia of Ornidia, dorsal view. 13, aemula Williston. 14, obesa Fabricius. 15, major Curran. 16, whiteheadi Thompson. black pilose except white pollinose under antenna; cheek shiny, black pilose; frontal triangle shiny, black pilose; frontal lunule brown; antenna brownish black, black pi- lose; arista orange, with aristal hairs black; vertical triangle shiny, black pilose; eye brown pilose on dorsal ' only; occiput shiny on ventral %, white pollinose elsewhere, white and black pilose. Thorax: metallic purplish black, shiny ex- cept narrowly grayish-white pollinose an- teriorly; pleuron black pilose; mesonotum black pilose, without pre-scutellar bristles; scutellum with pre-apical depression broad, shallow and not distinctly divided medially, black pilose; squama and plumula black; halter white except stem brownish. Wing: hyaline except for maculae; apical macula large, covering all of apex of cell R1+2. Legs: metallic purplish black, black pilose. Abdomen: metallic purplish black, shiny; venter black pilose except white pilose on basal 4 of 2nd sternum; 4th sternum with- out bristle apicolaterally; dorsum black pi- 254 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 17. Habitus of Ornidia whiteheadi Thompson. VOLUME 93, NUMBER 2 lose except with long tawny pile on basal 7 of 4th tergum; male genitalic segments black pilose. Size.—Mesonotal length, 3.9-4.4; wing length, 12.4-14.3 (see Val 1972). Types. — Volucella aemula was based on two female specimens, both of which are in the American Museum of Natural History with identical labels (““Brazil, Piedra, HH Smith” “‘Volucella aemula Williston’’). One is here designated lectotype and has been so labeled, the other paralectotype. Distribution.—Costa Rica*, Colombia’*, Venezuela*, Surinam, Brazil, Bolivia* (* in- dicates specimens examined from these countries). Ornidia aemula is easily distinguished from the other species of the genus by it large apical wing spot (Fig. 5), digitate pro- cess on the second sternum (Fig. 8) and the more pronounced facial tubercle (Fig. 1). Ornidia major Curran Pigs 1, 14315 Volucella violacea Macquart 1842: 23, pl. 6, fig. 3 (habitus), 3a (head). Type-local- ity: Brazil. Lectotypte 2 MNHN, Paris here designated. Preoccupied by Volucel- la violacea Say 1830. New synonym. Ornidia major Curran 1930: 2. Type-local- ity: Brazil. Holotype 6 AMNH. Val 1972 (biometry, distribution, male genitalia figured); Baez 1985: 74 (Venezuela). Metallic green or purplish blue flies. Head: face shiny, black pilose except white pollinose under antenna; cheek shiny, black pilose; frontal triangle shiny, black pilose; frontal lunule orange; front shiny, tawny and black pilose; antenna brownish black, black pilose; arista orange, with aristal hairs black; vertical triangle shiny, black pilose; eye black pilose; occiput shiny on ventral 4, white pollinose dorsally, white pilose with a few black hairs dorsally. Thorax: shiny except narrowly grayish- white pollinose anteriorly; pleuron white pi- lose except black pilose on anepisternum 255 and anepimeron; mesonotum black pilose, with short weak pre-scutellar black bristles; scutellum with pre-apical depression divid- ed medially, black pilose except tawny pi- lose on margins; squama and plumula black; halter white except stem brownish orange. Wing: hyaline except for maculae; apical macula small, only covering area around R1+2. Legs: front and mid coxae and tro- chanters metallic green, shiny; hind coxa metallic green, grayish-white pollinose; middle coxa black pilose; front and hind coxae white pilose basally, black pilose api- cally; femora shiny, dark metallic green to bluish black, becoming black apically, black pilose; tibiae metallic purplish black, shiny, black pilose; tarsi brownish black, black pi- lose. Abdomen: shiny except extensively black pollinose on 2nd tergum; venter black pilose except black on basal '4 of 2nd sternum; 4th sternum without bristles apicolaterally; Ist tergum tawny pilose; 2nd and 3rd terga short black pilose; 4th tergum long tawny pilose; male genitalic segments white pilose, except 8th tergum black pilose. Size.—Mesonotal length, 3.6—4.2; wing length, 10.6-12.5 (see Val 1972). Types. — Volucella violacea Macquart was based on an unspecified number of female specimens. A single female with the appro- priate Macquart determination label (“V. violacea, Mr. Sylvira Bresil’’) and museum labels (“Sylvair ... [unreadable], Brasil, 1839” “No 1094, Volucella, violacea’’) was found in Paris, is here designated lectotype and has so been labeled. Giglio-Tos (1892b: 64) synonymized violacea with obesa; this synonymy was accepted by subsequent workers. The holotype of major Curran was examined and found to correspond to the current concept of the name. Distribution.— Mexico (Vera Cruz)*, Guatemala*, Costa Rica*, Panama*, Co- lombia*, Venezuela*, Ecuador*, Peru*, Bra- zil*, Bolivia* and Paraguay”*. Ornidia major is distinguished from obesa by its larger size (see Val 1972), more or 256 less divided preapical scutellar depression and the presence of short prescutellar bris- tles; major differs from aemula and white- headi as noted under those species. Earlier authors, such as Williston (1891), misiden- tified major as obesa. Ornidia obesa Fabricius Figs. 3, 4, 6, 10, 14 Syrphus obesus Fabricius 1775: 763. Type- locality: “America” [= Virgin Islands]. Lectotype 6 UZM, Copenhagen (Thomp- son 1981: 195). Musca obesa: Gmelin 1790: 2868 (descrip- tion). Volucella obesa: Wiedemann 1830: 199 (re- descr.); Macquart 1842: 21 (distribution; Chile); Walker 1849: 637 (distribution, Mauritius, synonymy); Bigot 1859: 431 (Madagascar; descriptive note; breeds in latrines in Mexico); Macquart 1850: 311 (note on distribution); Williston 1887: 143 (description, head figured), 1891: 50 (syn- onymy, distribution); Giglio-Tos 1892b: 64 (distribution; synonymy), 1895: 360 (Seychelles); Aldrich 1905: 380 (catalog citation); Banks 1907: 450 (Virginia); Kertesz 1910: 763 (catalog citation); Bez- zi 1915: 4 (questionable record from Af- rica); Banks et al. 1916: 186 (Virginia); Sack 1921: 138 (Paraquay, larva figured); Jones 1922: 33, 53 (Colorado); Hull 1923: 297 (New Jersey); Curran 1925: 206 (male genitalia figured); Johnson 1925: 172 (Massachusetts); Johannsen 1926: 798 (New York); Bezzi 1928: 77 (Tahiti); Wil- liams 1931: 284, 1939: 284 (life history, Hawaii); Shannon & Aubertin 1933: 168 (Argentina; not in Chile); Doesburg 1963: 18, 1966: 96 (Surinam); Hardy 1964: 404 (Hawau, description, head & male geni- talia figured); Greenberg & Povolny 1971: 83 (public health importance); Greenberg 1971: 201 (biotic associates). Ornidia obesa: Lepeletier & Serville 1828: 786 (descr.; Asia, Mauritius); Val 1972 (biometry, geographic variation, male genitalia figured); Knutson et al. 1974: 335 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Oriental distribution); Smith & Vock- eroth 1980: 497 (Afrotropical distribu- tion); Thompson 1981: 120 (West Indian references, status); Baez 1985: 75 (Ven- ezuela); Thompson & Vockeroth 1989: 447 (Australian distribution); Bullock 1990: 253 (ex rotting Jacaratia mexicana (Caricaceae)). Volucella obesoides Giglio-Tos 1892a: 4. Type-locality: Mexico. Holotype ¢ IMZ. Giglio-Tos 1892b: 65 (description), 1895: 360 (Paraguay, Argentina); Godman & Salvin 1903: 90 (citation); Aldrich 1905: 380 (catalog citation); Kertesz 1910: 197 (catalog citation); Curran 1930: 2 (key). Synonymy by Thompson et al. 1976: 70. Ornidia obesoides: Curran 1930: 2 (key); Doesburg 1963: 18, 1966: 97 (Surinam). Metallic green or purplish blue flies. Head: face shiny, white pilose except white pollinose under antenna; cheek shiny, white pilose; frontal triangle shiny, white and black pilose; frontal lunule orange; front shiny, tawny and black pilose; antenna brownish black, black pilose; arista orange, with aris- tal hairs black; vertical triangle shiny, black pilose; eye brown pilose; occiput shiny on ventral “4, white pollinose dorsally, white pilose with a few black hairs dorsally. Thorax: shiny except narrowly grayish- white pollinose anteriorly and with black pollinose medial vitta on anterior 2 of mesonotum; pleuron tawny pilose with a few black hairs intermixed; mesonotum black pilose; scutellum with pre-apical de- pression continuous, not divided medially, black pilose except tawny pilose on margins; squama and plumula black; halter white ex- cept stem brownish orange. Wing: hyaline except for maculae; apical macula small, only covering area around R1 +2. Legs: front and mid coxae and trochanters metallic green, shiny; hind coxa metallic green, gray- ish-white pollinose; middle coxa black pi- lose; front and hind coxae white pilose ba- sally, black pilose apically; femora shiny, dark metallic green to bluish black, becom- VOLUME 93, NUMBER 2 ing black apically, black pilose; tibiae me- tallic purplish black, shiny, black pilose; tar- si brownish black, black pilose. Abdomen: shiny except black pollinose on most of 2nd tergum in male; venter yellow pilose except black on apical edge of 3rd and apical 3 of 4th sterna; 4th sternum without bristles apicolaterally; Ist tergum tawny pilose; 2nd and 3rd terga short black pilose; 4th tergum long tawny pilose; male genitalic segments tawny pilose, except 8th tergum black pilose; female genitalic seg- ments tawny pilose. Size.—Mesonotal length, 2.3-3.6; wing length, 7.4-11.2 (see Val 1972). Types.—The syntypes of obesa Fabricius were examined, and a lectotype designated (Thompson 1981). Vockeroth (in Thomp- son et al. 1976) examined the holotype of obesoides Giglio-Tos and identified it as a specimen of obesa Fabricius. Distribution. — Massachusetts, New York, New Jersey, Colorado, Arizona, New Mex- ico*, Texas*, Florida*, Bahamas*, Cuba’, Jamaica*, Dominican Republic*, Haiti*, Puerto Rico*, Lesser Antilles*, Trinidad*, Mexico*, Guatemala*, El Salvador*, Be- lize*, Honduras*, Nicaragua*, Costa Rica*, Panama*, Colombia*, Venezuela*, Suri- nam*, Guyana*, French Guiana*, Ecua- dor*, Peru*, Brazil*, Paraguay*, Bolivia*, Argentina*, Hawaii*, Marquesas*, Society Islands*, Marshall Islands*, Samoa*, Niue*, Solomon Islands*, New Caledonia*, Java, Seychelles, Madagascar, Mauritius, Re- union, Tanzania, South Africa. Ornidia obesa is a ubiquitous fly in the New World tropics and once was more widespread in the Old World tropics and the north of the New World. In the New World, the species have been recorded as far north as Colorado, Massachusetts and New York, and in the Old World tropics as far west as mainland Africa, but there are no modern records from these areas [in the above synonymy I have tried to give all the exotic citations to obesa, but only the prin- cipal New World ones]. Ornidia obesa is 257 known to breed in human latrines and other semiliquid wastes, and back when sanita- tion was not as good as today, Ornidia obesa apparently had a more extensive range. As Macquart (1850: 311) first noted, Ornidia obesa is the only New World fly that has spread to the Old World tropics. Ornidia obesa is known to carry bacteria of public health importance (Sa/monella, Shigella) (Alcivar and Campos 1946) and Mycobac- terium (Currie 1910). The species is also beneficial as the maggots can convert coffee- production waste products into useful pro- tein sources for cattle feed (Larde 1989). Ornidia whiteheadi Thompson, NEw SPECIES Figs. 12, 16 Male.— Head: metallic green; face shiny, black and tawny pilose except white polli- nose under antenna; cheek shiny, white pi- lose; frontal triangle shiny, tawny and black pilose; frontal lunule orange; antenna brownish black, black pilose; arista orange, with aristal hairs black; vertical triangle shiny, black pilose; eye brown pilose; oc- ciput shiny, white pilose with a few black hairs dorsally. Thorax: metallic green, shiny except nar- rowly grayish-white pollinose anteriorly; pleuron tawny pilose with a few black hairs intermixed; mesonotum tawny pilose ex- cept medial '4 black pilose, with a single weak pre-scutellar black bristle on left side; scutellum with pre-apical depression divid- ed medially, black pilose except tawny pi- lose on margins; squama and plumula taw- ny; halter orange except stem brownish. Wing: hyaline except for maculae; apical macula small, only covering area around R1 +2. Legs: coxae and trochanters metallic green, shiny, white pilose; femora shiny, bright metallic green and tawny pilose on basal 13, becoming dark apically, with apex black, black pilose on apical 7; tibiae me- tallic purplish black, shiny, black pilose; tar- si brownish black, black pilose. Abdomen: metallic green, shiny; venter 258 white pilose except bare on medial 4 of 3rd and *%3 of 4th sterna; 4th sternum with ap- icolateral black pile fused together to give appearance of strong black bristles; Ist ter- gum tawny pilose; 2nd tergum black pilose on apical ¥4, tawny pilose basally; 3rd ter- gum short black pilose on apical '3 and nar- row medially, elsewhere tawny pilose; 4th tergum tawny pilose; male genitalic seg- ments white pilose, except 8th tergum black pilose; male genitalia as figured (Figs. 12, 16), with a large postanal process posterior to cerci and between surstyles (see Fig. 12b). Female similar to male except for normal sexual dimorphism and more extensively pale tawny pilose: mesonotum almost en- tirely tawny pilose, 2nd and 3rd terga tawny pilose except for black pilose on apical '/. Size.—Mesonotal length, 3.0-3.2; wing length, 9.5-10.2. Holotype.— Male, PANAMA: Cerro Campana, 15 July 1963, W. G. Real, de- posited in California Academy of Sciences, San Francisco. Paratypes: 3 females in Unit- ed States National Museum. Panama: Canal Zone, Barro Colorado Island, 13 May 1979, Silberglied & Aiello, ovipositing on Afta dung. COLOMBIA: Antioquia, Aljibes, Providencia, c. 33 km sw Zaragosa; 29 De- cember 1970, Richard W. Pinger, “in ham- ster trap’; same locality and collector, 13 October 1970. Ornidia whiteheadi is similar in size and appearance to major, but is readily distin- guished from all other Ornidia species by the extensive tawny pile on the mesonotum, pleuron and 3rd tergum. The structure of the male genitalia is also unique among vol- ucellines. Ornidia whiteheadi is named for Donald R. Whitehead (see the appended eulogy). When I first saw the unique male (back in 1974) I asked Don, who knew Central America well, about the type locality —was it unique in any way? Don thought not. We then discussed the ethics of describing spe- cies from unique specimens. Our conclu- sion was that such new species should only PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON be described when the more inclusive taxa (genus, tribe or family) are revised. Don as- sured me that more specimens of my new species would appear some day as Central America was an area of active exploration, but, like the man for which the species is named, whiteheadi remains represented by a unique male! So, to remain true to his convictions, I have herewith revised the more inclusive taxon, Ornidia! ACKNOWLEDGMENTS I thank Adrian Pont, the Natural History Museum (formerly the British Museum (Natural History)), London (BMNH); Neal Evenhuis, Bishop Museum, Honolulu; Paul Arnaud, Jr., California Academy of Sci- ences, San Francisco (CAS); David A. Gri- maldi, the American Museum of Natural History, New York (AMNH); and Loic Ma- tile, Museum National d’Histoire Naturelle, Paris (MNHN); for permission to study ma- terial in their care. I also thank Neal Evenhuis (vide supra); Lynn Carroll, Douglass R. Miller, Alma So- lis and Norman E. Woodley of the System- atic Entomology Laboratory, USDA, Washington, D.C.; and Greg Courtney of the Smithsonian Institution (USNM), Washington, D.C., for their critical reviews of the manuscript. The color plate was pre- pared by Gustavo Hormiga of the Maryland Center for Systematic Entomology, Uni- versity of Maryland, College Park; the line drawings by Britt Griswold of Arlington, Virginia, and Mary Lou Cooley of the Sys- tematic Entomology Laboratory, USDA, Washington, D.C. LITERATURE CITED Aldrich, J. M. 1905. A catalogue of North American Diptera. Smithsonian Miscellaneous Collections 46(2). 680 pp. Alcivar, Z. C. and R. F. Campos. 1946. Las moscas, como agentes vectores de enfermedades entericas en Guayaquil. Revista Ecuatoriana de Higiene y Medicina Tropical 3: 3-14. Baez, M. 1985. Datos sobre la distribucion del genero VOLUME 93, NUMBER 2 Ornidia en Venezuela (Diptera: Syrphidae). Bo- letin de Entomologia Venezolana 4(9): 73-76. Banks, N. 1907. Captures of Diptera. Entomological News 18: 450. Banks, N., C. T. Greene, W. L. McAtee, and R. C. Shannon. 1916. District of Columbia Diptera: Syrphidae. Proceedings of the Biological Society of Washington 29: 173-203. 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Lille 1841(1): 65-200, 22 pls.) 1850. Dipteres exotiques nouveaux ou peu connus. 4.e. supplement. 77 pp. Paris. (Also pub- lished in Mem. Soc. Sci. Agric. Lille 1847(2): 161- 23i=) Sack, P. 1921. Dr. L. Zurcher’s Dipteren-Ausbeute aus Paraguay: Syrphiden. Archiv fur Naturge- schichte (A)87: 127-149. Shannon, R. C. and D. Aubertin. 1933. Syrphidae. Part VI, pp. 120-170. /n Diptera of Patagonia and South Chile Based Mainly on Material in the Brit- ish Museum (Natural History). British Museum, London. Smith, K. G. V. and J. R. Vockeroth. 1980. 38. Fam- ily Syrphidae, pp. 488-510. Jn Crosskey, R. W., ed., Catalogue of the Diptera of the Afrotropical Region. British Museum (Natural History), Lon- don. 1437 pp. Thompson, F. C. 1972. A contribution to a generic revision of the Neotropical Milesinae (Diptera: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Syrphidae). Arquivos de Zoologica, Sao Paulo 23: 73-215. 1981. The flower flies of the West Indies (Diptera: Syrphidae). Memoirs of the Entomolog- ical Society of Washington 9. 200 pp. Thompson, F. C. and J. R. Vockeroth. 1989. 51. Family Syrphidae, pp. 437-458. Jn Evenhuis, N. L., ed., Catalog of the Diptera of the Australasian and Oceanian Regions. Bishop Museum Press and E. J. Brill, Honolulu and Leiden. 1154 pp. Thompson, F. C., J. R. Vockeroth, and Y. S. Sedman. 1976. Syrphidae. Catalogue of the Diptera of the Americas South of the United States 46. 195 pp. Thompson, F. C. and D. R. Whitehead. 1986. Cla- distic statements as cladistic formulae or how to hate cladistics and love mathematics. Biologie und Naturwissenshaft der Kafer 3: 27-32. Val, F.C. do. 1972. On the biometry and evolution of the genus Ornidia (Diptera, Syrphidae). Papeis Avulsos Zoologica 26(1). 28 pp. Walker, F. 1849. List of the Specimens of Dipterous Insects in the Collection of the British Museum. Part III. British Museum, London. [3] + 485-687 pp. Wiedemann, C. R. W. 1830. Aussereuropaische zweiflugelige Insekten. Vol. 2, 684 pp. Hamm. Williams, F. X. 1931. Handbook of the Insects and Other Invertebrates of Hawaiian Sugar Cane Fields. 400 pp. Experiment Station, Hawaiian Sugar Planters’ Association, Honolulu. 1939. Biological studies of Hawaiian water- loving insects. Part III. Diptera or flies. B. Astei- idae, Syrphidae and Dolichopodidae. Proceedings of the Hawaiian Entomological Society 10: 281- Shey Williston, S. W. 1887. Synopsis of the North Amer- ican Syrphidae. Bulletin. United States National Museum (1886) 31, 335 pp. 1888. Diptera Brasiliana, ab H. H. Smith collecta. Part I—Stratiomyidae, Syrphidae. Trans- action of the American Entomological Society 15: 243-292. 1891-1892. Fam. Syrphidae, pp. 1-56 (De- cember 1891), 57-79 (February 1892). In God- man, F. D., and O. Salvin, eds., Biologia Centrali- Americana, or, Contributions to the Knowledge of the Fauna and Flora of Mexico and Central America. Zoologia. Diptera. Vol. HI. Taylor & Francis, London. VOLUME 93, NUMBER 2 261 Don Whitehead Remembered (1938-1990) I don’t know when I first met Don, but I first came to know him when I joined the staff of the Systematic Entomology Laboratory back in 1974. Don was then a Research Fellow working for Dan Janzen. Dan is interested in saving the tropics and was then trying to figure out how tropical dry forests, such as those in Costa Rica, really work. Seeds and the insects associated with them were the key components to this puzzle. Don was Dan’s insect specialist. And there has never been a specialist as versatile and competent as Don. While he worked mainly with John Kingsolver deciphering the patterns of evolution and distribution of seed weevils, Don also tackled some parasitic wasps and seed bugs. Don had worked on ground beetles for his graduate work and his real love was for millipeds. From his first major paper, which received unusual praises in a rare review (for a Ph.D. thesis) in Systematic Zoology, every paper of Don’s was carefully crafted to present the findings and always a unique interpretation of their significance. Don had the best mind in the laboratory. When I needed an answer Don was my source. In the beginning it was cladistics, statements of how things were interrelated. Back then there were no PCs, one either had access to mainframe computers or one worked it out by hand. Don was better than either approach! Once I received a manuscript for review. I knew the cladogram, the tree of relationships, was wrong despite the author’s assertion otherwise, for I saw a few misplaced characters. However, I wanted to be certain as I was reviewing someone else’s work. So, with manuscript in hand I went off to see if Don was ready for a beer. It was late in the afternoon, and Don and I did our best thinking over a beer or two, in those days. With a pencil and couple of napkins, Don quickly found a number of shorter trees, with one significantly so. Today kids with their PCs and Hennig86s can do this, but then and now only Donald Whitehead could do it the old-fashioned way faster! Later the questions became more mundane, like what is the best way to prioritize research in SEL. But regardless of topic, I always sought Don’s advice on virtually everything. Don was quiet, a careful thinker. Hence, he wasn’t always speaking out at meetings. He would instead dissect the problem, consider the logical alternatives, and formulate the best argument. So, frequently his input came later, and hence many will never know how critical Don was in shaping his Science and organization, SEL. I can only say what Newton said before, if I have seen further and achieved more, it’s because I have stood on the shoulders of giants. Don was my giant. Don was always a support for me; from providing the little things like reading my manuscript to critical things like sharing thoughts on life. Don’s last act for me was to re-direct me to the essentials. Last summer when Don discovered he had cancer, he quietly re-assessed his life, ordered his affairs, and turned his mind back to critical questions. There is one unique charac- teristic of life on this earth and that is that it is diverse. ““Why is life so diverse?” is the essential question for systematists. Don began with that question, providing insights to its answer with his studies on ground beetles, seed weevils and the like. Don had always been fascinated by the strange and delightful many-legged ““worms,” millipeds, that are found in our mountains. The diverse array of colors of these millipeds didn’t seem to correlate with where Don found them. So over the last year, Don searched to formulate an answer to this puzzle. And in October at Providence, he told the Eastern Branch of the Entomological Society of America that he had an answer. He likewise told his colleagues here at a meeting of the Entomological Society of Wash- ington. I was with him at both places. Don was having fun for he was again reveling in the excitement of his science. He was struggling with that essential question of why life is diverse. All the while I was lost in the strange world of government red-tape trying to get a computer system for the laboratory. Don saved me, his example being an inspiration— Order your life, seek the essential, and you will find your rewards. Don’s devotion to Science will always live. His colleagues have and will continue to establish monuments to it. And the uniqueness of some of these monuments is such that all generations will know his name. Years ago and in the mountains of Mexico, George Ball and Don pursued ground beetles. Don also collected a stray milliped. Today that creature is known as Geoballus caputalbus, or in English, Whitehead’s milliped. No one will ever forget that name! PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 262-271 THE ANTHONOMUS GUTTATUS SPECIES GROUP (COLEOPTERA: CURCULIONIDAE) WAYNE E. CLARK Department of Entomology and Alabama Agricultural Experiment Station, Auburn University, Alabama 36849-5413. Abstract. —Four species of neotropical Anthonomini are assigned to the Anthonomus guttatus group. Adults of two of these species have been collected on Casearia sylvestris Sw. (Flacourtiaceae). Characters diagnostic of the A. guttatus group and of each of the species are described and some are illustrated. A key to the species is presented. Three new species are described: A. argocephale (Brazil), A. albocapitis (British Virgin Islands, Cuba, Haiti, Puerto Rico, Republica Dominicana) and A. leucocephale (Cuba). A lectotype is designated for A. guttatus (Champion). Relationships of the 4. guttatus group to A. sallei Burke and the A. triensis group are discussed. Key Words: The Anthonomus guttatus group contains four species that occur in México, Central and South America and the West Indies, two of which have been collected on Cas- earia sylvestris Sw. (Flacourtiaceae). Cham- pion (1903) placed A. guttatus (Champion), with the other Central American Anthono- mini known by him to have 6 antennal funicular articles, in the genus Pseudan- thonomus Dietz. As Burke (1979) asserted, however, A. guttatus does not belong in that genus. The species was recently transferred to Anthonomus (Clark 1990), along with several of the other species Champion (1903, 1910) had assigned to Pseudanthonomus but in which the upper margin of the lateral rostral groove 1s directed to the upper mar- gin rather than to or below the lower margin of the eye. This paper includes descriptions, illustrations and a key to A. guttatus and three previously undescribed species in the A. guttatus group and discussions of the re- lationships of the group to A. sallei Burke and to the A. triensis group. It is dedicated to the memory of the late Donald R. White- Anthonomus, caesaria, Anthonomini, Curculionidae head. The specific epithets, Latin or Greek combinations of words meaning “white” and “head,” join Anthonomus whiteheadi (Clark In Press) as tokens of my apprecia- tion to Don for his help and encouragement over the years. MATERIALS AND METHODS Specimens of 78 adult weevils, including the types of the previously described spe- cies, were examined. These were from the collections of the following individuals and institutions (codens identify the collections in the text): AMNH_ The American Museum of Nat- ural History, New York, New York: USA, L: B. Hermann: AUEM_ Auburn University Entomologi- cal Collections, Auburn, Ala- bama, USA, W. E. Clark; BMNH The British Museum (Natural History), London, England, R. T. Thompson; VOLUME 93, NUMBER 2 CWOB C. W. O’Brien Collection, Tal- lahassee, Florida, USA; Universidade Federal do Parana, Curitiba, Brazil, G. H. Rosado- Neto; H. and A. Howden Collection, Ottawa, Canada; Museum of Comparative Zool- ogy, Cambridge, Massachusetts, USA, D. G. Furth; Museo Nacional de la Historia Natural, Santo Domingo, Re- publica Dominicana, A. Zaglul; Museu de Zoologia, Universi- dade de Sao Paulo, Sao Paulo, Brazil, U. Martins; Texas A&M University, College Station,’ Texas. WiSA, HH: OR: Burke; National Museum of Natural History, Washington, D.C., USA, D. R. Whitehead; Museum fiir Naturkunde der Humboldt-Universitat, Berlin, DDR, F. Hieke. Measurements were made with the aid of an ocular micrometer in a dissecting micro- scope as follows: total length from anterior margin of eye to elytral apex in lateral view; width across elytra at widest point; length of pronotum, dorsally, from anterior to pos- terior margins; length of rostrum from an- teroventral margin of eye to apex, across arc, in lateral view; length of distal portion of rostrum from antennal insertions to apex in lateral view; width of frons at narrowest point between eyes; width of base of ros- trum just distad of eyes in dorsal view; and width of pro- and metafemora, in anterior view, excluding the inner marginal teeth. The range and, in parentheses, the mean and sample size of each measurement are given for each species. DZUP HAHC MCZC MHND MZSP TAMU USNM ZMHB The Anthonomus guttatus Species Group Recognition Characters.—The A. gutta- tus group includes Anthonomini with 6 an- 263 tennal articles; a single profemoral tooth; broad, leucine scales in postscutellar patch- es on sutural interstriae, in a basal patch on interstria 6, and in a diagonal median row and a transverse declivital row of patches (Figs. 1-8); abdominal tergum 7 of male (Fig 9) with integument thickened posteri- orly, with deeply emarginate internal phrag- ma; pygidium of female (Fig. 10) with an inverted v-shaped median sulcus anterior to similarly shaped median carina with deeply emarginate internal phragma and with a bilobed, apicomedian prominence; endophallus (Figs. 15-18) with a median sclerite bounded by two lateral sclerites; and tegmen without parameres. The species are further characterized as follows: Length: 2.00—3.04 mm. Width: 1.02-1.70 mm. Head: vertex with elongate, narrow, pallid scales; venter with broader, imbri- cated, lacteous scales; eyes strongly convex posteriorly. Rostrum: proximal portion ru- gulose; upper margin of lateral rostral groove directed to upper mid-portion of eye; distal portion rugulose, glabrous. Prothorax: each puncture with an elongate, apically round- ed, leucine scale or a narrower, aeneus scale; broad scales abundant in narrow middorsal vitta. Elytra: interstriae with apically rounded, leucine scales and narrower, ae- neus scales; slightly more convex, darker in color and glabrous in alternating patches on even-numbered interstriae anterior and posterior to most prominent patches of pal- lid scales; strial punctures small, with mi- nute setae. Abdomen: posterior margin of sternum 5 of male broadly, subquadrately emarginate; abdominal sternum 5 of female with shallow apicolateral emarginations that receive apicolateral prominences of pygid- ium; endophallus minutely denticulate proximally. Legs: profemur about as wide as metafemur; protibia straight, with inner- marginal prominence; metatibial mucro of male short, slightly curved; metatibial mu- cro of female minute. Plant associations. — Adults of two of the 264 species in the A. guttatus group, A. argo- cephale in Brazil and A. albocapitis in the Republica Dominicana, have been collected on Casearia sylvestris Sw. (Flacourtiaceae). One specimen of the latter species was taken, according to label data, on Faramea occi- dentalis (L.) A. Rich. (Rubiaceae). The im- mature stages and developmental sites of the species are unknown, so the host asso- ciations of the species cannot be ascer- tained. It seems likely, however, that C. syl- vestris will be found to be a host because of the large number of specimens collected on the plant. Relationships. — The thickened posterior margin with the emarginate internal phrag- ma of abdominal tergum 7 of the male (Fig. 9) and the similar, corresponding, inverted v-shaped median sulcus with emarginate i1n- ternal phragma of the pygidium (tergum 7) of the female (Fig. 10) are similar to the *“*... Inverted v-shaped to broadly u-shaped line(s) . . .”’ found in the species of Anthon- omus in the A. triensis group (Clark In Press). It seems possible that the bilobed, apico- median prominence of the pygidium of the female of the species in the A. guttatus group might be homologous to the apicolateral py- gidial prominences observed in the females of the A. triensis group (Clark In Press). It likewise seems reasonable that the “pair of spines and . . . more distal median sclerite”’ (Clark In Press) of the species in the A. trien- sis group are homologous to the three large endophallic sclerites in the A. guttatus group. It should be noted, however, that the api- colateral pygidial prominences are also shared with the species in the 4. alboan- nulatus group and others (Clark In Press), and that the endophallic sclerites are similar to those in species in the genus Atractomerus (Clark 1989). The species in the A. triensis group are somewhat similar to the species in the A. guttatus group in having variegated lighter and darker ferruginous to piceous elytral and pronotal integument, and in pos- session of a large, posterolateral, elytral macula bounded by patches of pallid scales. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The species in the 4. guttatus group exhibit nothing comparable, however, to the en- larged profemur with the large, bi-emargin- ate tooth of the species in the A. triensis group. As noted by Burke (1979), A. sallei Burke “superficially” resembles A. guttatus. That Mexican and Central American species is like the species in the A. guttatus group in possession of 6 antennal funicular articles and in having a similar pattern of patches of broad, pallid scales on the elytra. Ab- dominal tergum 7 of A. sallei lacks the mod- ifications described above for the species in the A. guttatus group, however, and the en- dophallus lacks the three elongate sclerites that characterize the species in that group. It is intriguing that A. sallei, like some of the species in the A. guttatus group, is known to be associated with plants in the genus Casearia (Burke 1979). KEY TO SPECIES OF ANTHONOMUS IN THE A. GUTTATUS SPECIES GROUP 1. Glabrous sections of elytral interstriae 2 and 4 between anterior and declivital patches of broad, pallid scales interrupted medially by diffuse patch of similar scales (Fig. 4); meta- tibia of male (Fig. 12) with well-developed, median, inner-marginal prominence Scaratere Oe Se ee ET Ee A. argocephale 1’. Glabrous sections of elytral interstriae 2 and 4 between anterior and declivital patches of broad, pallid scales continuous, not interrupt- ed medially by diffuse patch of such scales (Figs. 1, 2, 5-8); metatibia of male without inner-marginal prominence (Fig. 11), OR, in- ner marginal prominence at apical ' (Figs. 13, VPP Reena uta RYAN B bia Board dodo bs ob & 2 Elytral interstriae 3 and 7 without significant patches of pallid scales, interstria 5 without declivital patch (Figs. 1, 2); metatibia of male (Fig. 11) curved, without inner-marginal prominence; profemoral tooth uncinate .... NPSL EE ool te ATO Ee aaa oO oO eB A. guttatus 2'. Elytral interstriae 3 and 5 with significant an- terior and declivital patches of pallid scales, interstria 7 with small anterior patch (Figs. 5S— 8); metatibia of male (Figs. 13, 14) not curved, with inner-marginal prominence at apical 1; profemoral tooth straight 3. Metatibia of male with well-developed inner- marginal prominence (Fig. 13); rostrum of fe- N VOLUME 93, NUMBER 2 male long, strongly, evenly curved (Fig. 5) .. EGE Bice d PON Oba CODE See o ore. 4. albocapitis 3’. Metatibia of male with feebly developed in- ner-marginal prominence (Fig. 14); rostrum of female shorter, most strongly curved at point of antennal insertion ........... A. leucocephale Anthonomus guttatus (Champion) igs 2) ble 15 Pseudanthonomus guttatus Champion 1903: 195. Lectotype (here designated). PAN- AMA. Chiriqui: male [Sp. figured] [Bu- gaba,/ Panama./ Champion.] [é9] [B.C.A.Col.IV.4/ Pseudanthonomus/ guttatus,/ Champ.] [Type] (BMNH). Paralectotypes. PANAMA. Chiriqui: 1 female, mounted with the lectotype; 1 male [32] [Bugaba,/ Panama./ Champi- on.] [B.C.A.Col.IV.4/ Pseudanthono- mus/ guttatus,/ Champ.]; 1 male, 5 fe- males [Bugaba,/ Panama./ Champion.] [B.C.A. Col.IV.4/ Pseudanthonomus/ guttatus,/ Champ.] (BMNH). Blackweld- er 1947: 840. O’Brien and Wibmer 1982: 113. Burke 1979: 203. Anthonomus guttatus (Champion). Clark 1990: 657. Recognition characters (Figs. 1, 2).—An- thonomus guttatus is distinguished from the other members of the 4. guttatus group by the following combination of characters: El- ytra (Figs. 1, 2) glabrous between median and declivital patches of pallid scales on interstriae 2 and 4, without significant an- terior and declivital patches of pallid scales on interstriae 3 and 7, but with fairly dis- tinct anterior patch on interstria 5; meta- tibia of male (Fig. 11) curved; profemoral tooth curved; broad, pallid scales on lower portion of propleuron replaced on lower mid-portion by narrower scales. The eyes are less strongly prominent in 4. guttatus than in the other members of the A. guttatus group. Male.— Length: 2.24—2.64 mm (mean = 2.43, n = 6). Width: 1.26-1.42 mm (mean = 1.33, n = 6). Head: eyes not prominent, 265 separated by distance approximately 0.7 x width of rostrum at base. Rostrum: length 1.38-1.57 (mean = 1.50, n = 6) x pronotal length; distal portion 34-38% (mean = 36, n = 6) of total rostral length. Prothorax: interspaces between punctures narrow; broad, pallid scales forming broad postoc- ular vitta on pleuron. E/ytra: leucine scales elongate, dense in diagonal median and de- clivital rows of patches on interstriae 2, 4, 6, 8 and 10. Abdomen: aedeagus (Fig. 15) subparallel sided, with slight apical prom- inence; endophallus (Fig. 15) with median sclerite long, lanceolate, lateral sclerites shorter, stouter. Legs: profemoral tooth slender, acute, curved; protibia with inner- marginal prominence feebly developed; protibial uncus short, nearly straight. Female.— Length: 2.00-2.72 mm (mean = 2.46,n= 10). Width: 1.10-1.52 mm (mean = 1.34, n = 10). Rostrum: length 1.56-1.84 (mean = 1.71, n = 10) x pronotal length; length of distal portion 43-50% (mean = 47, n = 10) of total rostral length. Legs: metatibia nearly straight, without inner- marginal prominence. Discussion.—Champion (1903: 195) considered A. guttatus to be closely related to A. curvicrus (Champion), noting that the femoral tooth “‘arises from near the middle” in both species. The metatibia is curved in the male of both species (the female of A. curvicrus is unknown), but all of the femora are curved, and more strongly so, in A. cur- VICTUS. Burke (1979) stated that A. sallei differs from A. guttatus “‘in its larger size, stouter legs, shorter, stouter rostrum and the con- stricted appearance of the head.” Plant associations. — Specimens of A. gut- tatus from Turrialba, Cartago, Costa Rica, labelled ‘“‘moss on tree trunks” were ex- amined, but there is no indication of the host relationships of the species. Distribution.—In addition to the 9 syn- types from Panama, 13 specimens of A. gut- tatus from the following localities were ex- amined. BRAZIL. Sao Paulo: Fazhenda Pau 266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-8. Anthonomus guttatus group members, habitus, lateral and dorsal views. 1, 2. A. guttatus, 2, Turrialba, Cartago, Costa Rica; 3, 4. A. argocephale, , Chapec6, Santa Catarina, Brazil; 5, 6. A. albocapitis, °, 10.5 km E Sabana de la Mar, El Seibo, Republica Dominicana; 7, 8. A. /leucocephale, 6, holotype. VOLUME 93, NUMBER 2 d’alho, Ita (1 MZSP). COSTA RICA. Car- tago: Turrialba (6 HAHC). San José: 2 km S Col6n (1 HAHC). MEXICO. San Luis Potosi: Tamazunchale (2 USNM). VENE- ZUELA. Lara: Terepaima (1 AUEM). Anthonomus argocephale, NEw SPECIES Figs. 4, 12, 16 Type series.— Holotype. BRAZIL. Santa Catarina: male [Brasilien/ Nova Teutonia/ 27°11'B - 52°23’L/ Fritz Plaumann/ 14 XI 1949/ 300 W. 500 m.] [Pflanze/ 640] (MZSP). Paratypes. BRAZIL. Parana: | fe- male [Parana/ Guarauna/ 12-40] [4877] [Colecao/ F. Justus Jor] [DPT°/ ZOOL/ UF- PARANA] 1 female [DPT°/ ZOOL/ UF- PARANA] [PIRACUARA - PARANA/ BRASIL - 20/ 11/ 1970/ Marinoni & Moure]. Rio Grande do Sul: | female [75 28/ Pio Buek] [Porto Alegre/ 73.9.44] [Cas- earia/ sylvestris Bery] [4990] [Gregorio Bondar/ Collection/ David Rockefeller/ Donor]. Santa Catarina: 2 males [CHA- PECO’'/ 27°07 52'33/ 600 m] [F. Plaumann/ Nov. 1962]. 1 male, 1 female [Brasilien/ Nova Teutonia/ 27°11'B - 52°23'L/ Fritz Plaumann/ IX 1954/ 300 W. 500 m.]; 1 male [Brasilien/ Nova Teutonia/ 27°11'B - 52°23’L/ Fritz Plaumann/ X 1962/ 300 W. 500 m.]. Total paratypes, 8 (AMNH, DZUP, MZSP). Recognition characters (Figs. 3, 4).—An- thonomus argocephale is distinguished from the other members of the A. guttatus group by the following combination of characters: Elytra (Figs. 3, 4) with glabrous sections be- tween median and declivital patches of broad, pallid scales on interstriae 2 and 4 interrupted medially by diffuse patches of similar broad, pallid scales, without signif- icant declivital patches of pallid scales on interstriae 3 and 7, but with small anterior patch on interstria 5; metatibia of male (Fig. 12) with well-developed inner-marginal prominence; broad, pallid scales on lower portion of propleuron replaced on lower mid-portion by narrower scales. It is most likely to be confused with A. guttatus, but 267 differs from that species by the above listed characters as well as by having more prom- inent eyes and by the metatibia of the female which has the outer margin broadly concave and has a broad inner-marginal promi- nence. Male.— Length: 2.24—2.52 mm (mean = 2.39, n = 4). Width: 1.22-1.46 mm (mean = 1.32, n = 4). Head: eyes prominent, sep- arated by distance approximately 0.7 x width of rostrum at base. Rostrum: length 1.39-1.59 (mean = 1.46, n = 4) x pronotal length; distal portion 30-37% (mean = 34, n = 4) of total rostral length. Prothorax: interspaces between punctures narrow; broad scales forming broad postocular vitta on pleuron. E/ytra: leucine scales elongate, dense in diagonal median, posteromedian and declivital rows of patches on interstriae 2, 4, 6, 8 and 10. Abdomen: aedeagus (Fig. 16) constricted in distal '4, narrowed to bluntly rounded apex; endophallus with median sclerite long, slender, spatulate, lat- eral sclerites shorter, slender. Legs: profem- oral tooth slender, acute, conical; protibia with well-developed inner-marginal prom- inence; protibial uncus short, slightly curved. Female.— Length: 2.24-2.48 mm (mean = 2.41,n=4). Width: 1.28-1.44 mm (mean = 1.40, n = 4). Rostrum: length 1.68-1.92 (mean = 1.78, n = 4) x pronotal length; length of distal portion 42-54% (mean = 46, n = 4) of total rostral length. Legs: meta- tibia with slight inner-marginal promi- nence, outer margin slightly concave. Plant associations.— Label data indicate that a paratype of A. argocephale from Porto Alegre, Rio Grande do Sul, Brazil, was col- lected on Casearia sylvestris. Distribution. —Anthonomus argocephale is known only from the type series from Brazil. Anthonomus albocapitis, NEw SPECIES Figs:.5;6;.9, 10) fea Type series.— Holotype. REPUBLICA DOMINICANA. E/ Seibo: male [REPUB- LICA Dominicana/ El Seibo: 10.5 km. E/ 268 Sabana de la Mar/ 11 Sept. 1983/ W. E. Clark] [on Casearia/ sylvestris Sw./ (Fla- courtiaceae)/ dt. S. McDaniel] [27213/ host] (MHND). Paratypes. BRITISH VIRGIN ISLANDS. Tortola: 1 male [British Virgin Is./ Tortola, Meyers/ 18-19 Aug. 1982/ R. S. Miller, colr.]. CUBA. 1 male [Cuba,/ G. Wright]; 1 male [Cayamas/ 5.2 Cuba] [EA Schwarz/ Collector]; 2 males, 2 females [Cayamas/ 24.2 Cuba] [EA Schwarz/ Col- lector]; 1 male, 1 female [Cayamas/ 26.2 Cuba] [EA Schwarz/ Collector]; 1 male [Cayamas/ 2.3 Cuba] [EA Schwarz/ Collec- tor]; 1 female [Cayamas/ 4.3 Cuba] [EA Schwarz/ Collector]; 1 female [Cayamas/ 4.3 Cuba] [EA Schwarz/ Collector] [404.]; 1 fe- male [Cayamas/ 6.3 Cuba] [EA Schwarz/ Collector]. Camaguey: 1 male [CUBA: Monte Imias/ nr. California,/ Camaguey Prov./ June 7, 1959/ M. W. Sanderson/ C59- 20]. HAITI. ile de la Tortue: 1 male [Tortue Is]./ Haiti EC&/ GM Leonard] [Bisse Terre/ Apr. 29/ #20]. REPUBLICA DOMINI- CANA. El Seibo: 13 males, 15 females [RE- PUBLICA Dominicana/ El Seibo: 10.5 km. E/ Sabana de la Mar/ 11 Sept. 1983/ W. E. Clark]. Santiago: | male [fthills Cord. Cent./ S. of Santiago/ June ’38, Dom. Rep./ Dar- lington]. PUERTO RICO. 1 female [Por- torico/ Krug S. G.] [259./ Anthonomus./ Germ.] [Zool. Mus./ Berlin]; 1 female [Por- torico/ Krug S. G.]. Aguadilla: 1 male [Aguadilla/ P.R. 6-26-50/ Faraemea/ occi- dentalis] [Stringer/ San Juan/ 10,279/ 5G- 15661] [Anthonomus/ sp./ REW/ II-51]; 1 female [PUERTO RICO/ Barriomora,/ Is- abella, VIII-4-58/ A. F. A. Sanderson]. Are- Figs. 9, 10. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cibo: 2 males [PUERTO RICO:/ Camba- lache For./ Arecibo/ XI-6-1959/ A. M. Nadler]. Guayama: 1 female [Aibonito, P.R./ July 14-17, °14]. Mayagtiez: 2 males [Mayaguez, P.R./ VI-2-1932/ Coll: F. Mora] [Stuart/ T. Danforth/ Collection]; 1 male, 1 female [Mayaguez, P.R./ VI-4-1932/ Coll: F. Mora] [Stuart/ T. Danforth/ Collection]; 1 male [Mayaguez, P.R./ July 24-29, °14] [Anthonomini]. Ponce: 1 female [5 mi. N. E. Jayuya/ P.R. VII 23, 1969/H. & A. How- den]. Total paratypes, 57 (AMNH, AUEM, CWOB, MCZC, MHND, TAMU, USNM, ZMHB). Recognition characters (Figs. 5, 6).—An- thonomus albocapitis is distinguished from the other members of the 4. guttatus group by the following combination of characters: Elytra (Figs. 5, 6) glabrous between median and declivital patches of broad, pallid scales on interstriae 2 and 4, with significant an- terior and declivital patches of pallid scales on interstriae 3 and 5, and with anterior patch on interstria 7; metatibia of male (Fig. 13) with inner-marginal prominence dis- placed distally; scales on vertex of head dis- tinctly narrower than scales around dorsal margins of eyes and on frons. Male. — Length: 2.04-2.64 mm (mean = 2.30, n = 10). Width: 1.02-1.44 mm (mean = 1.24, n = 10). Head: with short, broad, lacteous scales around dorsal margins of eyes and on frons; eyes prominent, separated by distance approximately 0.7 x width of ros- trum at base. Rostrum: length 1.41-1.58 (mean = 1.48, n = 10) xX pronotal length; distal portion 31-38% (mean = 36, n = 10) — Anthonomus albocapitis, abdominal terga, dorsal view. 9. terga 7 and 8 (pygidium), ¢ holotype; 10. tergum 7 (pygidium), 2 paratype, 10.5 km E Sabana de la Mar, El Seibo, Republica Dominicana. Anthonomus guttatus group members, 4 right metathoracic legs, posterior views. 11. A. guttatus, Turrialba, Cartago, Costa Rica; 12. A. argocephale, holotype; 13. A. albocapitis, holotype; 14. A. leucocephale, Figs. 11-14. holotype. Figs. 15-18. Anthonomus guttatus group members, aedeagus, dorsal views. 15. A. guttatus, Turrialba, Car- tago, Costa Rica; 16. A. argocephale, holotype; 17. A. albocapitis, holotype; 18. A. leucocephale, holotype. VOLUME 93, NUMBER 2 269 270 of total rostral length. Prothorax: interspac- es between punctures narrow; broad scales forming small anterolateral patches, present on lower portion of pleuron. E/ytra: leucine scales short, dense in median patch on in- terstria 2, in anteromedian row of patches on interstriae 4-8, and in declivital fascia across interstriae 2-10; median patches on even-numbered interstriae longer than patches on odd-numbered interstriae; sec- tions of declivital fascia on interstriae 4 and 5 longer than sections on other interstriae. Abdomen: aedeagus (Fig. 17) slightly wid- ened in distal 3, narrowed to subtruncate apex; endophallus (Fig. 17) with median sclerite short, slender, lateral sclerites long. Legs: profemoral tooth conical, acute; pro- tibia with well-developed inner-marginal prominence; protibial uncus with slight bas- al prominence, nearly straight. Female.— Length: 2.32-3.04 mm (mean = 2.56,n= 10). Width: 1.24—1.70 mm (mean = 1.42, n = 10). Rostrum: length 1.50-1.70 (mean = 1.59, n = 10) X pronotal length; length of distal portion 38-52% (mean = 46, n = 10) of total rostral length. Abdomen: pygidium with shallow apicodorsal depres- sion. Legs: metatibia nearly straight. Plant associations.—Specimens of A. al- bocapitis were collected at the type locality on Casearia sylvestris. Label data on a para- type from Aguadilla, Puerto Rico, indicate that the specimen was collected on Faramea occidentalis. Distribution.—Anthonomus — albocapitis is known from the type series from the West Indian islands of Cuba, Hispanola, Puerto Rico and Tortola. Anthonomus leucocephale, NEw SPECIES Figs. 7, 8, 14, 18 Type series.— Holotype. CUBA. male [Cayamas/ 10.6 Cuba] [EA Schwarz/ Col- lector] (USNM). Paratype. 1 female [Cuba] [Wickham/ Collection/ 1933] (USNM). Recognition characters (Figs. 7, 8).—The near absence of an inner-marginal metatib- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ial prominence in the male (Fig. 14), the narrower, more strongly tapered aedeagus (Fig. 18), and the different relative lengths of the endophallic sclerites (the median sclerite is much longer and the lateral scler- ites are much shorter in A. /eucocephale, cf. Figs. 17, 18) distinguish A. /eucocephale from A. albocapitis. The two species are obvi- ously very closely allied. Although the metatibial prominence is only feebly de- veloped in A. leucocephale, the slight prom- inence that does exist is in the same unique position as that in A. albocapitis. It is no- table that the metatibial prominence is less well developed in Cuban specimens of A. albocapitis than in the other specimens in the type series. Furthermore, the patches of pallid scales on the elytra, though less ex- tensive in A. leucocephale, are arranged in the same pattern as the ones in A. albocapitis (cf. Figs. 5-8). Similarly, the aedeagus, though narrower in A. /eucocephale, has es- sentially the same form as that in 4. albo- capitis (cf. Figs. 17, 18). Male. — Length: 1.96 mm (n = 1). Width: 1.02 mm (n = 1). Head: eyes prominent, separated by distance approximately 0.8 x width of rostrum at base. Rostrum: length 1.17 (n = 1) X pronotal length; distal por- tion 32% (n = 1) of total rostral length. Pro- thorax: interspaces between punctures broad, convex; broad scales present on low- er portion of pleuron. E/ytra: leucine scales short, dense in median patch on interstria 2, in anteromedian row of patches on in- terstriae 4-8, and in declivital fascia across interstriae 2-10; median patches on even- numbered interstriae longer than patches on odd-numbered interstriae; sections of de- clivital fascia on interstriae 4 and 5 longer than sections on other interstriae. Abdomen: aedeagus (Fig. 18) narrowed to subtruncate apex; endophallus (Fig. 18) with median sclerite long, slender, lateral sclerites short. Legs: profemoral tooth conical, acute; pro- tibia with inner-marginal prominence fee- bly developed; protibial uncus slender, curved, acute. VOLUME 93, NUMBER 2 Female.— Length: 2.04 mm (n = 1). Width: 1.16 mm (n = 1). Rostrum: length 1.34 (n = 1) X pronotal length; length of distal portion 41% (n = 1) of total rostral length. Legs: metatibia nearly straight. Plant associations.— Unknown. Distribution. — Anthonomus leucocephale is known from the type series from Cuba. ACKNOWLEDGMENTS Thanks are extended to the individuals and institutions listed in the Materials and Methods section for the loan of specimens. The manuscript was reviewed by H. R. Burke, G. L. Miller, and M. L. Williams. This paper is published as Alabama Agri- cultural Experiment Station Journal Series No. 17-902722P. LITERATURE CITED Blackwelder, R. E. 1947. Checklist of the coleopte- rous insects of Mexico, Central America, the West Indies and South America, Part 5. Bulletin. United States National Museum (185): I-IV, 765-925. Burke, H.R. 1979. New species of Mexican and Cen- tral American Anthonomus (Coleoptera: Curcu- Pap lionidae). The Southwestern Entomologist 4(3): 201-208. Champion, G. C. 1903. Curculionidae: Curculioni- nae, Volume 4, Part 4, pp. 145-312. Jn F. D. Godman and O. Salvin, eds., 1879-1911. Biologia Centrali-Americana, Insecta, Coleoptera, 7 vol- umes in 17 parts. Dulau, London. 1910. Biologia Centrali-Americana. Insecta. Coleoptera. Rhynchophora. Curculionidae. Cur- culioninae (concluded) and Calandrinae, vol. 4, pt. 7, pp. 79-221. In F. D. Godman and O. Salvin, eds., 1879-1911. Biologia Centrali-Americana, Insecta, Coleoptera, 7 volumes in 17 parts. Dulau, London. Clark, W. E. 1989. Revision of the Neotropical wee- vil genus Atractomerus Duponchel and Chevrolat (Coleoptera: Curculionidae). Transactions of the American Entomological Society 114: 313-413. 1990. The Neotropical species of Pseudan- thonomus Dietz (Coleoptera: Curculionidae). Transactions of the American Entomological So- ciety 116(3): 655-695. In press. Revision of the Anthonomus al- boannulatus and Anthonomus triensis species groups (Coleoptera: Curculionidae). Coleopterists Bulletin. O’Brien, C. W. and G. J. Wibmer. 1982. Annotated checklist of the weevils (Curculionidae sensu lato) of North America, Central America, and the West Indies (Coleoptera: Curculionoidea). Memoirs of the American Entomological Institute 34: i-ix + 1-382. PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 272-277 NEW WORLD CNEMOGONINI (COLEOPTERA: CURCULIONIDAE). NEW SPECIES, NEW COMBINATIONS AND A TYPE SPECIES DESIGNATION ENZO COLONNELLI USDA-ARS Biological Control of Weeds Laboratory—Europe, via Gastone Monaldi, 34-00128 Rome, Italy. Abstract.—Two new species of weevils from Arizona are described and illustrated: Auleutes donaldi and Orchestomerus whiteheadi. Orchestomerus Dietz is removed from synonymy of Hypocoeliodes Faust. New combinations are: Orchestomerus bicarinatus (Champion), O. chiriquensis (Champion), O. gibbicollis (Champion), O. modestus (Hus- tache), O. phytobioides (Champion), O. pleurostigma (Faust), O. suturalis (Hustache), O. ulkei Dietz, O. wickhami Dietz, all from Hypocoeliodes. The type species of Orchestomerus, O. wickhami Dietz, is designated. Auleutes marionis Fall is newly placed in synonymy with Orchestomerus ulkei. Key Words: The tribe Cnemogonini Colonnelli, 1979, comprising 12 genera and 116 species (Co- lonnelli 1979, O’Brien and Wibmer 1982, Wibmer and O’Brien 1986), is the only tribe in the subfamily Ceutorhynchinae whose distribution is mainly in the New World; the genera Augustinus Korotyaev, 1981 (10 species) and Phytobiomorphus Wagner, 1937 (2 species), both from east Asia, and the holarctic Auleutes epilobii (Paykull, 1800) are not known to occur in the Western Hemisphere. The Cnemogonini are in need of revision, and a paper on the taxonomy of this tribe with descriptions of new genera and new species is in preparation. It is not my intention here to attempt the resolution of the many phylogenetic problems still open; I take today the opportunity to pay homage to the memory of my good col- league and an outstanding taxonomist, Dr. Donald R. Whitehead, naming after him two new species from Arizona, one of which was submitted to me, with a label ‘‘?Cra- Coleoptera, Curculionidae, Arizona, new combinations, new synonymy ponius n. sp.,”” by Donald himself some time ago. Auleutes donaldi, NEw SPECIES Holotype male.—Length: 2.67 mm. Width: 1.78 mm. Pitchy-brown, somewhat shining; antenna (club excepted), tarsi and tibial mucros reddish brown. Upper surface with sparse intermixed blackish and white semi-erect hairlike scales and white appli- cate narrowly lanceolate scales; the latter form an undulate fascia just before the mid- dle of the elytra and are condensed into a periscutellar spot and a few scattered spots in the posterior half of the elytra. Whitish sparse lanceolate recumbent scales on under surface. Rostrum 0.75 times shorter than prothorax, gently curved, coarsely punc- tured, and with a trace of smooth longitu- dinal carina. Antenna short, inserted just before middle of rostrum; scape with apical hook; funicular joint 1 much thicker than others; joints 6 and 7 transverse; club large, acuminate oval. Frons depressed, vertex VOLUME 93, NUMBER 2 f 4 \ s) 6 Figs. 1-8. 213 0,5 mm 0,5 mm 3 7 ) Auleutes donaldi n. sp., holotype: 1, dorsal view; 3, metatibia; 4, dorsal, 6, lateral view, 5, apex of aedeagus. 4. instabilis Champion, male from Costa Rica: 2, dorsal view; 7, dorsal and 8, lateral view of aedeagus. Schematic drawings. with carina. Prothorax transverse (length/ width = 0.8/1), coarsely punctured, strongly constricted anteriorly, widest at bisinuate base, apical margin shallowly incised at middle and with faint trace of obtuse tu- bercle at each side of incision; disc before middle with two obtuse tubercles separated by shallow furrow; lateral tubercles mod- erately prominent. Elytra slightly transverse (I/w = 0.94/1), widest just behind humeri; humeral and preapical calli not very prom- inent. Striae formed by large deep punc- tures, each of which bears a thin recumbent seta. Intervals not much wider than striae, the odd-numbered intervals more raised and wider than the others. Legs slender; femora unarmed, tibia almost straight, meso and metatibia with obvious spinelike mucro; tarsi slender; claws bifid. Rostral channel very deep, base of metasternum emarginate; abdominal segment 5 with shallow central fovea. Aedeagus: Figs. 4, 5 and 6. See also Figs. 1 and 3. Paratypes.— Males are very similar to the holotype. Females differ in the absence of tibial mucros and abdominal fovea; their rostrum is moreover much smoother in its apical half. One specimen, slightly imma- ture, has integumental color reddish brown. Length: 2.66-—2.70 mm. Type series. — Holotype é: Arizona, Santa Rita Mts., Madera Canyon, Bog Springs Campground, 5000’, 24. VIII.1988, coll. R. Baranowsky, by evening sweeping in oak forest. Paratypes: same data as holotype, | 2; Madera Canyon near Bog Springs Camp- ground, 5000’, 22.VIII.1988, 4 6 and 2 9, coll. R. Baranowsky, by sweeping in mixed forest. Holotype and 4 paratypes in the Mu- seum of the University, Lund; | paratype in the U.S. National Museum, Washington, D.C.; 2 in the author’s collection, Rome. Remarks. —Auleutes donaldi is quite iso- lated among the species of the genus. It ex- hibits only a superficial similarity with A. curvipes Dietz, 1896 from Texas; the single male specimen at present known of the lat- ter has however the fore tibia curved and bearing a small hook at its inner apical an- gle, a basal longitudinal sulcus on the pro- 274 notum, and the white scales on the elytra are not arranged to form a sinuate fascia (Dietz 1896). The new species cannot be confused with any of Auleutes described to date; it has only a slight resemblance to the Central American A. instabilis Champion, 1907, which has no or a very faint sulcus on the pronotum, odd-numbered intervals not wider than the others, only the male mid tibia with a mucro, and a differently shaped aedeagus (see Figs. 2, 7, 8). Orchestomerus whiteheadi, NEw SPECIES Holotype male.—Length: 2.45 mm. Width: 1.78 mm. Reddish brown, dull; an- tenna and tarsi reddish. Upper surface with white and brown recumbent hairlike scales forming very faint irregular fascia on apical half of elytra; velvety brown small round scales are condensed into a periscutellar spot; traces of the rustlike pollen which cover the integument in living specimens are evident. Scattered, yellowish, lanceolate, recumbent scales on under surface. Rostrum about as long as prothorax, slightly curved, coarsely punctured except at apex, and with sharp longitudinal carina. Antenna inserted 0.43 times the length of rostrum from apex of beak; scape with apical spine; joint | of fu- niculus thicker than others; joint 6 round, 7 transverse; club acuminate oval. Frons with shallow depression, vertex finely car- inate. Prothorax transverse (l/w = 0.75/1), coarsely punctured, constricted in front, widest just before sinuate base, apical mar- gin shallowly incised at middle, the incision limited at each side by obtuse tubercle; disc convex, two central obtuse tubercles sepa- rated by longitudinal complete furrow, lat- eral tubercles prominent. Elytra transverse (I1/w = 0.875/1), widest just behind promi- nent humeral calli, preapical tubercles very faint. Striae with deep punctures each bear- ing a recumbent extremely thin seta. Inter- vals wider than striae; odd-numbered in- tervals wider than others; interval 3 with elongate tubercle just before middle; 5 with similar elevation starting just behind base; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tubercle of interval 7 situated in basal third of the elytra. Legs short; femora unarmed; fore and middle tibia angulate at base, mid- dle tibia with obvious apical mucro, very small apical hook of hind tibia very difficult to see. Rostral channel deep on prosternum and mesosternum, metasternal excavation broad and shallow; segment 5 of abdomen with shallow central fovea. Aedeagus: Figs. 22, 23 and 24; see also Fig. 13. Paratypes. — The general appearance of the males does not differ from that of the ho- lotype. Females differ in their abdomen be- ing uniformly convex and their tibiae lack- ing apical mucros. The reddish brown integumental color can be more or less dark, independently from the degree of matura- tion of the specimen. Length: 2.44—2.79 mm. See also Figs. 14, 18, 20, 31 and 33. Type series. — Holotype 6: Arizona, Santa Rita Mts., 10.VI, coll. Hubbard and Schwarz. Paratypes: same data as holotype, 7 6 and 2 2; same locality, 20.V, 2 6 and 1 O22 TEV,. 6s 27 SVs 29 INES 3 ONVeine: 31.V, 4 éand 2 9; 5.VI, 3 2; 9.VI, 1 6; 14. VI, Io 65. VI, vo and! 1992-21 Vie. coll Hubbard and Schwarz. Arizona, Chiricahua Mts., 1. VII, 1 2, coll. Hubbard and Schwarz; Chiricahua Mts., 6200’, 20. VI.1926, 1 4, coll. A. A. Nichol. Arizona, Santa Catalina Mts., 5000’, 7.VI.1926, 1 4, coll. A. A. Nichol. Holotype and 25 paratypes in the U.S. Na- tional Museum, Washington, D.C.; 8 in the author’s collection, Rome. Remarks.—Champion (1907) synony- mized Hypocoeliodes Faust, 1896, type spe- cies H. coronatus Faust designated by Champion (1907) (Wibmer and O’Brien 1986) and Orchestomerus Dietz, 1896, type species by present designation O. wickhami Dietz, 1986. Study of about 3000 specimens and of most of the types of the Cnemogonin1 (as a part of the revision of the tribe now in progress) revealed that this synonymy is incorrect. Although both genera are unde- niably closely related, diagnostic characters for Orchestomerus (resurrected name) are VOLUME 93, NUMBER 2 18 19 275 12 0,5 mm 13 esas 6 17 | ) 0,5 mm Figs. 9-21. Hypocoeliodes coronatus Faust, male from Venezuela: 9, lateral view of pronotum and head; 12, left front femur; 10, lateral and 11, dorsal view of aedeagus. Orchestomerus whiteheadi n. sp., holotype: 13, left front femur. O. whiteheadi, paratypes: 14, dorsal view; 18, lateral view of pronotum and head; 20, head and rostrum. O. chiriquensis Champion from Chiapas, Mexico: 15, dorsal view; 19, lateral view of pronotum and head. O. ulkei Dietz from North Carolina: 16, dorsal view; 21, head and rostrum. O. wickhami Dietz from Texas: 17, dorsal view. Schematic drawings. the unarmed femora, the lack of obvious punctures on the sides of the prothorax, and the ligulate apex of the aedeagus (Figs. 13, 18, 19, 22-30). In Hypocoeliodes the femora are toothed (in some species very feebly), the sides of the pronotum show obvious deep punctures much larger than the coarse punc- tation which covers the rest of the protho- rax, and the aedeagus exhibits a broadly truncated apex (Figs. 9-12). In addition, the size of most species of Hypocoeliodes is greater and the integumental color is darker than those of Orchestomerus. An alphabetic list of all the hitherto described species be- longing to Orchestomerus (all are new com- binations from Hypocoeliodes) follows: O. bicarinatus (Champion, 1907), O. chiri- quensis (Champion, 1907), O. gibbicollis 276 Aa Ag] PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0,5 mm cq 0,5 mm 33 34 Figs. 22-34. Orchestomerus whiteheadi, holotype: 22, lateral, 23, dorsal view, 24, apex of aedeagus. O. whiteheadi, paratypes: 31, tarsus; 33, male hind tibia. O. chiriquensis: 25, lateral, 26, dorsal view, 27, apex of aedeagus; 32, tarsus. O. ul/kei: 28, lateral, 29, dorsal, 30, apex of aedeagus; 34, male hind tibia. Schematic drawings. (Champion, 1907), O. modestus (Hustache, 1947), O. phytobioides (Champion, 1907), O. pleurostigma (Faust, 1896), O. suturalis (Hustache, 1947), O. ulkei Dietz, 1896 (= Auleutes marionis Fall, 1913; new synony- my), and O. wickhami Dietz, 1896. For the distribution of these species see O’Brien and Wibmer (1982) and Wibmer and O’Brien (1986). The new species is closely related to the Central American O. chiriquensis, from which it can be differentiated by the shorter rostrum, the shallow incision of the fore margin of the pronotum, the broader third joint of the tarsus, and the different shape of the aedeagus (Figs. 14, 15, 18, 19, 22- 27). O. whiteheadi cannot be confused with any other Orchestomerus so far described. In particular it is easily separated from the North American O. ulkei by the reddish brown rather than piceous integumental color, the thicker rostrum, the lack of an evident acute mucro at the inner apical an- gle of the male metatibia, and the different shape of the aedeagus (Figs. 14, 16, 20, 21, 22-24, 28-30, 33, 34). Study of quite a long series of specimens from the eastern United States revealed that Au/eutes marionis Fall is a synonym of O. ulkei Dietz. Although I have not examined the type specimens (both species were described on the basis of one specimen each), I have no doubt in pro- posing the synonymy. The compressed an- tennal joints of the species do not allow easy observation of the separation between the third and fourth joint when the antenna is examined from its compressed side. The other North American species, O. wick- hami, from Texas is smaller, its integument is reddish (at least on elytra), the recumbent scales of the prothorax and elytra are nu- merous and white, the inner tubercles of the pronotum are in the form of an oblique ca- rina (Fig. 17), and the male middle and hind tibia have a minute hook at their inner api- cal angles (Dietz 1896). ACKNOWLEDGMENTS I wish to express my sincere thanks to Roy Danielsson, Museum of the Univer- sity, Lund and Robert D. Gordon, System- atic Entomology Laboratory, USDA, U.S. VOLUME 93, NUMBER 2 National Museum, Washington, D.C. for the loan of specimens. I also thank Héléne Per- rin, National Museum of Natural History, Paris and Richard T. Thompson, British Museum (Natural History), London for as- sisting during the visit. Valuable material was also loaned to me by Anne T. Howden, Carleton University, Ottawa. Particular thanks go to Lloyd Knutson, Director, USDA-ARS Biological Control of Weeds Laboratory, Rome for reading the English manuscript. Useful comments were also provided by Wayne E. Clark. LITERATURE CITED Champion, G. C. 1907. Biologia Centrali-Ameri- cana. Insecta, Coleoptera. Curculionidae Curcu- lioninae. Vol. IV, part 5, pp. 137-240. London. 277 Colonnelli, E. 1979. Note su Panophthalmus e generi affini, con descrizione di una nuova tribu’ e di una nuova specie della sottofamiglia Ceutorhynchinae (Coleoptera, Curculionidae). Bollettino dell’ As- sociazione Romana di Entomologia 34: 1-9. Dietz, W.G. 1896. Revision of the genera and species of Ceutorhynchini inhabiting North America. Transaction of the American Entomological So- ciety 23: 387-480. O’Brien, C. W. and G. J. Wibmer. 1982. Annotated checklist of the weevils (Curculionidae sensu lato) of North America, Central America, and the West Indies (Coleoptera: Curculionoidea). Memoirs of the American Entomological Institute 34: i-ix + 1-382. Wibmer, G. J. and C. W. O’Brien. 1986. Annotated checklist of the weevils (Curulionidae sensu lato) of South America (Coleoptera: Curculionoidea). Memoirs of the American Entomological Institute 39: i-xvil + 1-563. PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 278-287 CLADISTIC ANALYSIS OF PRIOCYPHIS HUSTACHE AND RELATED GENERA (COLEOPTERA: CURCULIONIDAE) ANALIA A. LANTERI AND JUAN J. MORRONE Division Entomologia and Laboratorio de Sistematica y Biologia Evolutiva, Museo de La Plata, Paseo del Bosque, 1900 La Plata, Argentina. Abstract.—This paper presents the cladistic analysis and relationships of Priocyphus Hustache and related genera to test their monophyly and determines the taxonomic placement of some critical species. The cladistic analysis was carried out using 45 trans- formation series from the external morphology (including mouthparts) and male and female genitalia. Polarity decisions were based on the outgroup comparison with the genera Thoracocyphus Emden and Lamprocyphus Marshall. The most parsimonious cladogram obtained shows the following phylogenetic sequence: Lamprocyphopsis Lanteri [L. viri- dinitens (Kuschel) and L. paraguayensis Lanteri], Priocyphopsis Lanteri [P. humeridens (Hustache)], Mendozella Hustache [M. curvispinis (Hustache)], Cyrtomon Schoenherr [C. hirsutus (Hustache) nov. comb, C. ovalipennis (Hustache), C. glaucus (Bovie), and C. gibber species group Lanteri], and Priocyphus Hustache (P. bosqi species group Lanteri and P. inops species group Lanteri). Characters from the female and male genitalia are the most important to elucidate the relationships of the taxa under study. Key Words: Cladistic analysis, Priocyphus, Cyrtomon, Priocyphopsis, Lamprocyphopsis, Mendozella, Curculionidae, Naupactini The genus Priocyphus Hustache belongs to the tribe Naupactini and is restricted to the grasslands and savannahs of the Cha- coan domain (sensu Cabrera and Willink 1973) from Paraguay to central Argentina. This genus was established by Hustache (1939) and included three species, P. bosqi Hustache 1939, P. hirsutus Hustache 1939 and P. humeridens (Hustache 1926) (trans- ferred from Trichocyphus Heller). Kuschel (1950) described Priocyphus inops, C. hus- tachei, and P. viridinitens. He also added two species originally assigned to Neocy- phus Bovie (= Cyrtomon Schoenherr and Cyphus Germar), P. glaucus (Bovie 1907) and P. ovalipennis (Hustache 1938). Lanteri (1990a) hypothesized that Prio- cyphus sensu Kuschel (1950) was not nat- ural and split the group into three genera: Priocyphus (Priocyphus bosqi, P. hustachei, P. inops, and P. kuscheli Lanteri, 1990), Priocyphopsis (Priocyphopsis humeridens and P. hirsutus), and Lamprocyphopsis (Lamprocyphopsis viridinitens and L. par- aguayensis Lanteri, 1990a). Moreover, she placed P. glaucus and P. ovalipennis again in Cyrtomon (Lanteri 1990b) and suggested that the genus Mendozella Hustache is re- lated to both Priocyphus and Cyrtomon (Lanteri 1989). We consider the genera Lamprocyphop- sis, Priocyphopsis, Mendozella, Cyrtomon (sensu Lanteri 1990b), and Priocyphus (sen- su Lanteri 1990a) to be a monophyletic group (Figs. 1-5). Synapomorphies that jus- tify this group are as follows: VOLUME 93, NUMBER 2 Figs. 1-5. Female, dorsal view. 1, Lamprocyphopsis viridinitens (Kuschel); 2, Priocyphopsis humeridens (Hustache); 3, Mendozella curvispinis (Hustache); 4, Cyrtomon gibber (Pallas); 5, Priocyphus bosqi Hustache. —maxillae with subrectangular mala, hav- ing long lacinial teeth, and with truncate- conical palpal article 3; —dorsal comb of hind tibiae as long as to longer than apical comb; and -sclerites in the internal sac present. The relationships and monophyly of these genera are uncertain, as well as the place- ment of some critical species (Priocyphopsis hirsutus, Cyrtomon glaucus, and C. ovali- pennis). A cladistic analysis seems to be the most appropriate resolution to these prob- lems and to developing a phylogenetic clas- sification of the group. MATERIALS AND METHODS The study was based on the revisionary works of Lanteri (1989, 1990a, b), and the examination of specimens borrowed from the following collections: CWOB Charles W. O’Brien Collection, Tallahassee, Florida, USA Departamento de Zoologia da Universidade Federal do Parana, Curitiba, Brazil Fundacion e Instituto Miguel Lillo, San Miguel de Tucuman, gentina Museo Argentino de Ciencias Na- turales Bernardino Rivadavia, Buenos Aires, Argentina Museo de La Plata, La Plata, Ar- gentina DZUP FIML MACN MLPC MZSP Museu de Zoologia da Univer- sidade de S. Paulo, S. Paulo, Bra- zil MNHN Museum National d’Histoire Naturelle, Paris, France NZAC_ New Zealand Arthropod Collec- tion, Auckland, New Zealand The cladistic methodology here applied is summarized in Nelson and Platnick (1981), and Wiley (1981). Terminal taxa.—We selected nine ter- minal units (Table 1): the genus Lampro- cyphopsis, Cyrtomon gibber species group, Priocyphus bosqi and P. inops species groups, and the single species Mendozella curvi- spinis, Priocyphopsis humeridens, P. hirsu- tus, Cyrtomon glaucus, and C. ovalipennis. Lamprocyphopsis 1s a very homogeneous genus, including two species. Cyrtomon gib- ber species group comprises four taxa (Lan- teri 1990b), that according to Kuschel (1958) should be considered as a single species. The two species groups of Priocyphus are ho- mogeneous and clearly monophyletic (Lan- teri 1990a). Mendozella is a monotypic ge- nus. The two species of the genus Priocyphopsis were considered as separate units because P. hirsutus has a doubtful tax- onomic placement; this species was origi- nally described in Priocyphus (Hustache 1939) and tentatively assigned to Priocy- phopsis (Lanteri 1990a). Cyrtomon glaucus and C. ovalipennis are critical species that 280 Table 1. Species Groups 1 Cyrtomon gibber species group Lanteri 1990b Species C. gibber (Pallas 1781) C. luridus (Boheman 1840) C. pistor (Boheman 1833) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Terminal units, including species and geographical distribution. Geographical Distribution From central-eastern Brazil to Bo- livia, Paraguay, and central Ar- gentina. C. inhalatus (Germar 1824) tO Cyrtomon glaucus (Bovie 1907) 3 Cyrtomon ovalipennis (Hustache 1938) 4 Lamprocyphopsis Lanteri 1990a 5 Mendozella curvispinis (Hustache 1926) 6 Priocyphopsis hirsutus (Hustache 1939) 7 Priocyphopsis humeridens (Hustache 1926) 8 Priocyphus bosqi species group Lanteri 1990a 9 Priocyphus inops species group Lanter 1990a were described in Cyphus and Neocyphus, transferred to Priocyphus (Kuschel 1950), and included in Cyrtomon (Lanteri 1990b). Transformation series.—A detailed mor- phological study indicated 45 transforma- tion series, 29 from the external morphol- ogy (head, mouthparts, pronotum, elytra, and tibiae) and 16 from the genitalia (ster- num 8 of female, ovipositor, spermatheca, and aedeagus). Morphometric features with overlapping ranges and characters variable within a taxon were excluded. Male and female genitalia were not eval- uated in much detail or were ignored by earlier specialists (e.g. Germar 1824, Schoenherr 1833, Marshall 1922, Emden 1936, Hustache 1939, Kuschel 1950). The last revisionary works (Lanteri 1989, 1990a, b) considered them for diagnosing the gen- era of the group under study. Recent con- tributions (Diaz et al. 1990a, b) showed the importance of mouthparts to separate some taxa of the group, and several characters from this source were incorporated in this analysis. Identification of apomorphic characters L. viridinitens (Kuschel 1950) L. paraguayensis Lanteri 1990a P. bosqi Hustache 1939 P. hustachei Kuschel 1950 P. inops Kuschel 1950 P. kuscheli Lanteri 1990a Central-eastern Argentina. Central-northern Argentina. Northeastern Argentina and Para- guay. Central-western Argentina. Central-western Argentina. Northwestern Argentina. Central Argentina. Northeastern Argentina and Para- guay. was based on the outgroup comparison (Watrous and Wheeler 1981) with 7hora- chocyphus Emden as the first outgroup and Lamprocyphus Marshall as the second out- group. Binary coding was applied, except for twelve transformation series with more than one apomorphic character. Seven of these series (1, 7, 9, 15, 16, 22, 39, and 44) were treated as additive, and five (3, 22, 31, 40, and 43) as non-additive. The transformation series and characters are listed in Table 2. In the data matrix (Table 3), plesio- morphic characters were coded with 0, apo- morphic ones with numbers | and 2 (ad- ditive) or letters a, b, and c (non-additive), and unknown characters with question marks. Data analysis.—The data set was ana- lyzed using the phylogenetic program HENNIG86 version 1.5 (Farris 1988), ap- plying the implicit enumeration option. The consistency (Kluge and Farris 1969) and re- tention (Farris 1989) indices were calculat- ed excluding autapomorphies. VOLUME 93, NUMBER 2 Table 2. Series of transformation and characters. Plesiomorphic Characters (0) Apomorphic Characters (1, 2, or a, b, c) — Rostrum not truncate-conical. Rostral groove narrow to moderately wide. Rostral lateral carinae indistinct. Scrobes extending below eyes. Preocular depression distinct. Supraocular lobes absent. Eyes strongly convex. Antennae robust. Scape not reaching hind margin of eyes. Funicular article 2 longer than 1. Mandibular pharyngeal process short and wide. Maxillary mala suboval, with short lacinial teeth. Article 3 of maxillary palpus subcylindrical. Prementum hexagonal. Prementum with 15-25 short fine setae. Disc of pronotum flat to depressed, lacking groove. Disc of pronotum rugose. Pronotum lacking lateral tubercles. Base of pronotum not to slightly thickened. Base of pronotum and elytra bisinuate. Humeri strongly prominent. Humeri not tuberculate. Elytral declivity not abrupt. Supernumerary striae of even intervals ab- sent. Elytral setae recumbent. Elytral scales round to round-oval. Tibial denticles present. Number of tibial denticles 6-13. Dorsal comb of hind tibiae shorter than api- cal comb. Female sternum 8 elongate. Apodeme of sternum 8 1.25-2.5 as long as plate. Ovipositor lacking sclerotized dorsal plates. Coxites of ovipositor slightly sclerotized. Membrane between coxites with three folds. Styli not concealed by coxites. Spermatheca subglobose. Nodulus of spermatheca truncate-conical. Nodulus of spermatheca short. slightly truncate-conical (WF/WR = 1.15-1.45) (1). moderately truncate-conical (WF/WR = 1.50-1.65) (2). very wide (1). distinct, reaching fore margin of eyes (a). distinct, reaching hind margin of eyes (b). evanescent (1). indistinct (1). present (1). slightly to moderately convex (1). flat (2). slender (1). reaching or slightly exceeding hind margin of eyes (1). largely exceeding hind margin of eyes (2). about as long as | (1). long and narrow (1). subrectangular, with long lacinial teeth (1). truncate-conical (1). subcircular (1). with less than 11 long coarse setae (1). lacking setae (2). convex, with distinct groove (1). smooth (1). with lateral tubercles (1). conspicuously thickened (1). straight (1). moderately to slightly prominent (1). moderately tuberculate postero-laterally (a). moderately tuberculate antero-laterally (b). strongly tuberculate antero-laterally (c). abrupt (1). present (1). erect (1). lanceolate (1). absent (1). 4-5 (1). as long as, to longer than apical comb (1). transversal (1). about as long as plate (a). about 3.5 x as long as plate (b). with a pair of sclerotized dorsal plates (1). strongly sclerotized (1). with several folds (1). concealed by coxites (1). subcylindrical (1). tubular (1). medium-length (1). long (2). 282 Table 2. Continued. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Plesiomorphic Characters (0) Apomorphic Characters (1, 2, or a, b, c) 39 Nodulus of spermatheca not curved. slightly curved (1). strongly curved (2). 40 Spermathecal duct shorter than abdomen. twice as long as abdomen (a). as long as spermatheca (b). 41 Spermathecal duct fine (narrower than nod- ulus at apex). 42 Spermathecal duct membranous. 3 Apex of aedeagus acute. moderately to very wide (wider than nodulus at apex) (1). sclerotized (1). truncate (a). arrow-pointed (b). 44 Aedeagal apodemes shorter than aedeagus. as long as aedeagus (1). longer than aedeagus (2). 45 Sclerites of internal sac inconspicuous. conspicuous (1). RESULTS AND DISCUSSION A single most parsimonious cladogram was obtained (Fig. 6), with 65 steps, a con- sistency index of 0.72, and a retention index of 0.74. The relationship between the outgroup Thoracocyphus and the group under study is based on the synapomorphies “‘disc of pronotum convex, with distinct groove” (16), “coxites of ovipositor strongly scler- otized”’ (33), and “‘style concealed by cox- Itesin(3 5): The monophyly of Priocyphus and related genera is justified by the synapomorphies “‘antennae slender” (8), “scape reaching or slightly exceeding hind margin of eyes”’ (9.1), “maxillary mala subrectangular, with long Table 3. Data matrix. lacinial teeth” (12), ‘“‘article 3 of maxillary palpus truncate-conical”’ (13), ““prementum lacking setae”’ (15.2), ““dorsal comb of hind tibiae as long as to longer than apical comb” (29), and “‘sclerites of internal sac conspic- uous” (45). Lamprocyphopsis is the sister genus to the remaining taxa, which form a clade based on the synapomorphies “ros- trum slightly truncate-conical’’ (1.1), ““scrobes evanescent” (4), “‘preocular de- pression indistinct” (5), ““humeri moderate- ly to slightly prominent” (21), ““spermathe- ca subcylindrical”’ (36), ““spermathecal duct moderately to very wide” (41), ““sperma- thecal duct sclerotized”’ (42), and ‘“‘aedeagal apodemes as long as aedeagus” (44.1). Prio- cyphopsis is the sister genus to Mendozella, Lamprocyphus Thoracocyphus Cyrtomon gibber species group Cyrtomon glaucus Cyrtomon ovalipennis Lamprocyphopsis Mendozella Priocyphopsis hirsutus Priocyphopsis humeridens Priocyphus bosgi species group Priocyphus inops species group 000000000000000000000000000000000000000000000 0000000000 10000 101000a00010000a1 1010000000000 000110112011101000000011000010000001121a11b11 110110112011101110001010000010b00001121a11b11 100110112011101100001000000010000001111lal1b11 00000001201 110210000001000001 1a01110000000a01 100111112001111100001c00100010001011010a11b11 20b1 10011001 102100101b00000110001011000011021 10a110211101101100011000000010001011121011b11 VOLUME 93, NUMBER 2 283 a 2 S = = — “ Ce = = = Ss = = cc = eal 2s 23 S = 3 es es —e 3 = = 5 a =" aay a) os oo. o 2 o ® o = Paites E 3s Ss ar =a ice ca peal 2 = Ss iw = oz Sos oS = o> a 5 < 24 4 43a 31b 21 4Ot 34 642 17 16 39.2 Ha 28 25 384 2 1 27 vn 22b 22¢ 23 23 19 14 17 "1 3b 92 25 40 16 92 1.2 6 3a B 3a 12 "1 20 92 10 72 2 Ss 35 = oS > —) S o a 2 33 E 2 391 38.2 37 43b 40a 381 151 7A 44) 42 4 36 21 5 32 4 Hla W 26 22a 45 18 29 n 192 13 12 91 8 35 3 16 Fig. 6. Most parsimonious cladogram. —: apomorphies; =: parallelisms; *: reversals. Cyrtomon, and Priocyphus; these three gen- ‘‘spermathecal duct twice as long as abdo- era share the apomorphies “‘eyes slightly to men’’ (40.a), and “‘apex of aedeagus arrow- moderately convex” (7.1), “prementum with pointed” (43.b). The taxa included in Cyr- less than 11 coarse setae” (15.1), ‘“‘nodulss tomonand Priocyphus form a monophyletic of spermatheca medium-length”’ (38.1), group having three synapomorphies of the 284 spermatheca, “nodulus tubular” (37), “long” (38.2), and “‘slightly curved”’ (39.1). Cyrtomon is a monophyletic group that is justified by the reversals “‘coxites of ovi- positor slightly sclerotized”’ (33) and “‘styli not concealed by coxites’’ (35), and com- prises C. hirsutus, C. ovalipennis, C. glau- cus, and the C. gibber species group. Cyr- tomon ovalipennis is related to the clade C. glaucus—C. gibber species group by the syn- apomorphies “scape largely exceeding hind margin of eyes” (9.2) and ‘“‘mandibular pha- ryngeal process long and narrow” (11), with parallel evolution in Lamprocyphopsis. The sister taxa C. glaucus and C. gibber species group share the synapomorphy “‘elytral de- clivity abrupt” (23), with parallel evolution in Lamprocyphopsis and the P. inops species group. Priocyphus includes the inops and bosqi species groups. The monophyly of the genus is supported by the synapomorphies “eyes flat” (7.2), “‘funicular article 2 about as long as 1” (10), and “‘base of pronotum and elytra straight”’ (20). All the terminal units are characterized by autapomorphies. Character analysis.—External morphol- ogy: Characters of the rostrum do not con- tribute substantially to the elucidation of relationships among the taxa under study. The shape of the rostrum evolves from trun- cate-conical to moderately truncate-coni- cal, a change that occurs in other genera of the tribe Naupactini, e.g. the genus Afri- chonotus Buchanan (Lanteri and O’Brien 1990). The eyes are slightly to moderately con- vex 1n Mendozella and Cyrtomon. The transformation has a further change in Prio- cyphus where the eyes become flat. The most apomorphous condition has evolved coin- cidentally in single species of several genera of Naupactini, e.g. Eurymetopus Schoen- herr (Lanteri 1984), and Phacepholis Horn (Lanteri 1990c). The evanescent scrobes and indistinct preocular depression are correlated char- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON acters that justify the monophyly of the in- group except Lamprocyphopsis. These char- acters are rather uniform in the tribe Naupactini and vary among genera or groups of genera. The antennae do not vary significantly within the group under study, but help dis- tinguish it from the outgroups. Within the ingroup, the most conspicuous change is the shortening of funicular article 2 in Priocy- phus. Most Naupactini have funicular ar- ticle 2 longer than 1, but in some genera, such as Atrichonotus (Lanteri and O’Brien 1990) and Eurymetopus (Lanteri 1984) both articles have the same length or article 1 is slightly shorter than article 2. The shape of the mala and palpal article 3 of the maxillae and the reduction of setae on the labial prementum are correlated characters that establish the monophyly of the ingroup. The remaining characters of mouthparts are autapomorphies of different taxa. Characters of the pronotum and elytra are not very helpful to recognize clades within the group under study, but are important to characterize taxa. The humeri evolve from strongly prominent to slightly prominent and sometimes (Priocyphopsis and Men- dozella) there are also tubercles. The re- duction of humeri and the occurrence of tubercles are apomorphic characters that occur independently in other Naupactini, e.g. Enoplopactus Heller (Lanteri 1990d). The presence of denticles on the tibiae is the most primitive condition in the tribe Naupactini. Within the ingroup these den- ticles are lost in the Priocyphus inops species group. In other genera, such as Eurymetopus (Lanteri 1984), all species lack the tibial denticles. Genitalia: Sternum 8 of most Naupactini is elongate and subrhomboidal, with the apodeme about 2x as long as the plate. Within the group under study, the shape of sternum 8 of each taxon is slightly different, but in Lamprocyphopsis the shape is trans- versal and completely different. The length VOLUME 93, NUMBER 2 of the apodeme evolves in two opposite di- rections, it becomes much shorter in Lam- procyphopsis and longer in C. glaucus. In the ovipositor, the presence of strongly sclerotized coxites concealing styli is an apomorphic condition that is reversed in Cyrtomon. This condition appears indepen- dently in other non-related genera of Nau- pactini, e.g. Atrichonotus (Lanteri and O’Brien 1990), Eurymetopus (Lanteri 1984), and Teratopactus (Lanteri 1981). The spermathecal body evolves from subglobose to subcylindrical, and the nod- ules from subconical and short to tubular, long, and curved. The second condition is a clear synapomorphy at Priocyphus and Cyrtomon. Other non-related genera of Naupactini such as Asynonychus Crotch or Aramigus Horn (Lanteri et al. in press) also have tubular and long nodulus, but the sper- mathecal shape is different. The length and width of the spermathecal duct evolve in two different directions, very long and wide, and very short and narrow. In some genera of Naupactini, e.g. Enoplopactus (Lanteri 1990d), the spermathecal duct is very uni- form, whereas in others, e.g. Aramigus (Lanteri et al. in press) it has a great vari- ation. The apex of the aedeagus arrow-pointed is a clear synapomorphy of Mendozella, Priocyphus, and Cyrtomon, and seems to be correlated to apomorphic conditions of the spermatheca. In most Naupactini, the apex of the aedeagus is acute. CONCLUSIONS According to the most parsimonious cladogram Priocyphus sensu Kuschel (1950) is not a natural group because it does not include either the genus Mendozella or the Cyrtomon gibber species group. This result corroborates the hypothesis of non-mono- phyly that led Lanteri (1990a) to split Prio- cyphus into three different genera. Lamprocyphopsis was named in reference to its similarity to Lamprocyphus (Lanteri 1990a). The cladogram obtained shows that 285 this similarity is due to plesiomorphic char- acters. Priocyphopsis sensu Lanteri (1990a) is not a natural group because P. humeridens and P. hirsutus do not share any synapomorphy. Therefore, this generic name 1s herein re- stricted to the type species, P. humeridens. The similarity of Priocyphopsis and Prio- cyphus is due to plesiomorphies. The critical species hirsutus was previ- ously assigned to Priocyphus (Hustache 1939) and Priocyphopsis (Lanteri 1990a), and according to the cladogram, it belongs to Cyrtomon. The external morphology of this species is a mosaic of characters of the three genera, but the female genitalia de- termine its inclusion in Cyrtomon. On the same basis, C. ovalipennis and C. glaucus are placed in Cyrtomon. The results of the cladistic analysis sup- port the hypothesis of Lanteri (1989) that Mendozella is closely related to Cyrtomon and Priocyphus, and that the latter two are sister genera. Moreover, they confirm that the male and female genitalia are essential to elucidate the relationships among these genera. Since cladistics does not provide any rule to rank the natural groups in a cladogram, we may treat the group under study (Prio- cyphus sensu Kuschel + Mendozella + Cyr- tomon gibber species group) as a single ge- nus, or keep the five generic names pro- posed until present (Lamprocyphopsis, Priocyphopsis, Mendozella, Cyrtomon, and Priocyphus), with the condition that C. hir- sutus, C. ovalipennis, and C. glaucus belong to Cyrtomon. The first alternative would imply the recognition of a very heteroge- neous genus, difficult to diagnose. On the contrary, the second allows definition of small, homogeneous, and easily identifiable genera, with greater chance of representing natural units. We chose the second option because it agrees better with the criteria applied to rec- ognize other genera of the tribe Naupactini (Lanteri 1989, 1990c, d; Lanteri and O’Brien 286 1990). This option disrupts the present clas- sification as little as possible and is partic- ularly convenient for the tribe Naupactini, where the construction of a phylogenetic system is still developing. The cladistic classification proposed, ap- plying the method of phylogenetic sequenc- ing (Nelson, 1974), is as follows: Genus Lamprocyphopsis Lanteri 1990a L. viridinitens (Kuschel 1950) L. paraguayensis Lanteri 1990a Genus Priocyphopsis Lanteri 1990a P. humeridens (Hustache 1926) Genus Mendozella Hustache 1939 M. curvispinis (Hustache 1926) Genus Cyrtomon Schoenherr 1823 C. hirsutus (Hustache 1939) nov. comb. C. ovalipennis (Hustache 1938) C. glaucus (Bovie 1907) C. gibber species group C. inhalatus (Germar 1824) C. luridus (Boheman 1840) C. pistor (Boheman 1833) C. gibber (Pallas 1781) Genus Priocyphus Hustache P. bosqi species group P. bosqi Hustache 1939 P. hustachei Kuschel 1950 P. inops species group P. inops Kuschel 1950 P. kuscheli Lanteri 1990a ACKNOWLEDGMENTS We are grateful to the following people who loaned us material (institutions abbre- viated as in Materials and Methods): G. Ro- sado-Neto (DZUP), A. Teran (FIML), G. Kuschel (NZAC), A. Bachmann (MACN), H. Perrin (MNHN), R. Ronderos (MLPC), and C. Costa (MZSP), and to Jorge Crisci and Maria M. Cigliano for critically reading the manuscript. Preliminary versions of this paper were presented at the XV Congresso Brasileiro de Zoologia, Curitiba, Brazil (February 1988) and at the XVIII Interna- tional Congress of Entomology, Vancouver, Canada (July 1988). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Boheman, C. H. 1833. Jn Schoenherr, C. J., ed.,Gen- era et species curculionidum, cum synonymia hu- jus familiae. Roret, Paris, 1(1): 1-381; 1(2): 383- 681. 1840. Jn Schoenherr, C. J., ed., Genera et species curculionidum, cum synonymia hujus familiae. Roret, Paris; Fleischer, Lipsiae, 6(1): 1- 474. Bovie, A. 1907. Notes sur les Curculionides. Annales de la Société Entomologique de Belgique 51: 67— 7k Cabrera, A. L. and A. Willink. 1973. Biogeografia de America Latina. Monografias de la OEA, Serie Biologia, monografia 13. Diaz, N. B., M. S. Loiacono, M. del C. Coscar6on, and A. A. Lanteri. 1990a. Importancia taxonomica de las piezas bucales en la tribu Naupactini. I. Género Cyrtomon Schoenherr y taxa afines (Co- leoptera: Curculionidae). Revista Brasileira de En- tomologia (In press). Diaz, N. B., M.S. Loiacono, A. A. Lanteri, and M. del C.Coscaron. 1990b. Importancia taxonomica de las piezas bucales en la tribu Naupactini. II. Las especies del género Cyrtomon Schoenherr (Cole- optera: Curculionidae). Neotropica (In press). Emden, F. I. van. 1936. Die Anordnung der Brach- yderinae-Gattungen im Coleopterorum Catalo- gus. Stettiner Entomologische Zeitung 97(1): 66— 99, (2): 211-239. Farris, J.S. 1988. HENNIG86 reference. Version 1.5. Published by the author. 1989. The retention index and the rescaled consistency index. Cladistics 5: 417-419. Germar, E. F. 1824. Insectorum species novae aut minus cognitae, descriptionibus illustratae. Vol- ume I, Coleoptera. Hendel and Sons, Halde. Hustache, A. 1926. Contribution a l’étude des Cur- culionides de la République Argentine (premiére note). Anales del Museo Nacional de Historia Na- tural “Bernardino Rivadavia”’ 34: 155-261. 1938. Nouveaux Naupactini de l’Amérique meridionale (Coleoptera: Curculionidae). Bulletin de la Société Entomologique de France 43: 93-96. . 1939. Curculionides nouveaux de l’Argentine et autres régions Sudamericaines. Anales de la So- ciedad Cientifica Argentina 128: 38-64, 99-124. Kluge, A. G. and J. S. Farris. 1969. Quantitative phyletics and the evolution of anurans. Systematic Zoology 18: 1-32. Kuschel, G. 1950. Die Gattung Priocyphus Hustache 1939. (10. Beitrag zu Coleoptera: Curculionidae). Revista de Entomologia, Rio de Janeiro 21(3): 545- 550. 1958. Neotropische Riisselkaéfer aus dem Museum G. Frey (Coleoptera: Curculionidae). En- VOLUME 93, NUMBER 2 tomologische Arbeiten aus dem Museum G. Frey 9(3): 750-798. Lanteri, A. A. 1981. Estudio comparativo de las es- tructuras genitales en la tribu Naupactini. I. Los caracteres a nivel genérico de Naupactus Schén- herr, Teratopactus Heller y Trichonaupactus Hus- tache (Coleoptera: Curculionidae). Revista de la Sociedad Entomologica Argentina 40(1-4): 273- 278. 1984. Revision sistematica del género Eu- rymetopus Schénherr (Coleoptera: Curculionidae) mediante la aplicacion de técnicas numéricas. Re- vista de la Sociedad Entomologica Argentina 43(1- 4): 247-281. Lanteri, A. A., N. B. Diaz, M. S. Loiacono, and M. del C. Coscar6n. 1987. Aplicacion de técnicas nu- méricas al estudio sistematico del grupo de Asy- nonychus durius (Germar) (Coleoptera: Curcu- lionidae). Entomologische Arbeiten Museum G. Frey 35/36: 171-198. 1989. Estudio sistematico de los géneros Trichocyphus Heller y Mendozella Hustache (Co- leoptera: Curculionidae). Boletin de la Sociedad de Biologia de Concepcion 60: 139-147. . 1990a. Revision sistematica del género Prio- cyphus Hustache 1939 y creacion de los géneros Priocyphopsis y Lamprocyphopsis (Coleoptera: Curculionidae). Revista Brasileira de Entomologia 34(2): 403-422. 1990b. Revision sistematica del género Cyr- tomon Schonherr (Coleoptera: Curculionidae). Revista Brasileira de Entomologia 34(2):387-402. 1990c. Systematic revision and cladistic analysis of Phacepholis Horn (Coleoptera: Cur- culionidae). Southwestern Entomologist 15(2): 179-204. 1990d. Revision sistematica y analisis filo- genético de las especies del género Enoplopactus 287 Heller 1921 (Coleoptera: Curculionidae). Boletin de la Sociedad de Biologia de Concepcion 61: 71- 92. Lanteri, A. A., N. B. Diaz, and E. Ortiz Jaureguizar. Analisis multivariado y congruencia taxonomica en la delimitacion de los géneros Asynonychus Crotch y Aramigus Horn (Coleoptera: Curculion- idae). Revista de la Asociacion de Ciencias Na- turales del Litoral (In press). Lanteri, A. A. and C. W. O’Brien. 1990. Taxonomic revision and cladistic analysis of Atrichonotus Bu- chanan (Coleoptera: Curculionidae). Transactions of the Entomological Society of America 116(3): 697-725. Marshall, G. A. K. 1922. On new genera and species of Neotropical Curculionidae. Transactions of the Entomological Society of London, 1922, 1-2: 181- 224. Nelson, G. 1974. Classification as an expression of phylogenetic relationships. Systematic Zoology 22: 344-359. Nelson, G. and N.E. Platnick. 1981. Systematics and Biogeography. Cladistics and Vicariance. Colum- bia University Press, New York. Pallas, P.S. 1781. Icones insectorum praesertim Ros- siae Sibiriaeque peculiarium quae collegit et des- criptionibus illustravit. Walther, Erlangae. Part 1, 6 (2 pp.; ““Praemonenda,” 4 pp.), + 56 pp. Schénherr, C. J. 1833. Genera et species curculio- nidum, cum synonymia hujus familiae. Roret, Paris, 1(1): 1-381; 1(2): 383-681. Watrous, L. E. and Q. D. Wheeler. 1981. The out- group comparison method of character analysis. Systematic Zoology 30: 1-16. Wiley, E. O. 1981. Phylogenetics: The Theory and Practice of Phylogenetic Systematics. John Wiley and Sons, New York. PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 288-297 LARVA AND PUPA OF CLEONIDIUS ERYSIMI (FALL) WITH A DISCUSSION OF THE PHYLOGENETIC POSITION OF LIXINI (SENSU KUSCHEL) (COLEOPTERA: CURCULIONIDAE) RosertT S. ANDERSON Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4, Canada. I first met Don Whitehead in December of 1980 when I visited him in Washington to discuss a topic in weevil systematics suitable for a Ph.D. dissertation. With his help and advice I eventually selected North American Cleoninae as the group I would study. Not only was Don instrumental in the selection of my dissertation topic but he was also the reviewer of the published version which appeared in 1988; his insightful and in-depth commentary greatly improved the final product. All those who have ever discussed phylo- genetics with Don know of his keen interest in the subject and his unique means of dem- onstrating relationships (he brought new meaning to the term “hand cladistics’”’ and I suspect was often frustrated that he only had ten fingers). I feel that it is appropriate that the present paper, which continues to deal with the Cleoninae and their phylogenetic re- lationships, be dedicated to his memory. Abstract.—The last instar larva and pupa of Cleonidius erysimi (Fall) are described. Larvae mine the roots and crowns of Lobularia maritima (L.) Dev., an introduced species of Cruciferae, in southern California. Pupation takes place in the larval mines. The pres- ence of mandibular setae in the pupa of traditional Adelognatha, and Lixini, Thecesternini, Pachyrhychina (sensu Kuschel), and Ithycerus noveboracensis (Forster) suggests this char- acter state is symplesiotypic. A review of the phylogenetic placement of Lixini (sensu Kuschel) 1s confused by conflicting character state distributions. Key Words: Curculionidae, immature stages, phylogeny Recent systematic revision of adults of the New World weevils placed in the tribe Cleonina of the tribe Lixini (sensu Kuschel) (= tribe Cleonini of the subfamily Cleoninae in Anderson [1988]) questioned the tradi- tional placement ofall New World Cleonina in the genus Cleonis Dejean (O’Brien and Wibmer 1982), and instead showed there to be four rather distantly related genera rep- resented: Cleonis Dejean (1 introduced spe- cies), Stephanocleonus Motschulsky (6 na- tive species), Ap/eurus Chevrolat (8 native species), and Cleonidius Casey (19 native species) (Anderson 1988). Whereas phylo- genetic relationships of Cleonis, Stephano- cleonus and Apleurus were considered to be with various other traditional Cleonina, re- lationships of Cleonidius were more difficult to firmly establish, but appeared to be with one or more subgenera of the extremely spe- ciose, geographically widespread and struc- turally diverse genus Lixus Fabricius of the VOLUME 93, NUMBER 2 subtribe Lixina as well as with a genus of traditional Cleonina, Cylindropterus Chev- rolat (Anderson 1988). These relationships were established solely on the basis of the distribution of adult character states largely because of the unavailability of other life stages representative of both Old, but es- pecially, New World species. In a review of the immature stages of American Curculionoidea, Burke and An- derson (1976) noted that larvae and/or pu- pae have been sufficiently well-described for only the introduced Cleonis piger (Scorpoli) and Rhinocyllus conicus (Froelich), and for but two native species of Lixus, L. scrobi- collis (Boheman) (Boving and Craighead 1930) and L. concavus Say (Peterson 1951). Immature stages of species of Ap/eurus, Cleonidius, and North American Stepha- nocleonus (an unidentified Palearctic spe- cies was very briefly described by Scherf [1978]) have not previously been described. Immature stages of a number of Old World Lixini (sensu Kuschel) have been very brief- ly described and diagrammatically illus- trated by Scherf (1964). This paper describes the pupal and last larval instars for Cleonidius erysimi (Fall) and thus represents the first description of the immature stages of that genus. Unfor- tunately, few meaningful phylogenetic in- terpretations can be made of characters of immature Cleonidius because of the general lack of broad taxonomic representation within the tribe, particularly those taxa ap- parently closely related to Cleonidius as de- termined on the basis of adult characters. Although nothing can be said of the phy- logenetic relationships of Cleonidius and other Lixini (sensu Kuschel) based on char- acter states of the immature stages, the phy- logenetic relationships of Lixini (sensu Kus- chel) and other Curculionidae can be discussed in this light. Terms for the pupa follow those of May (1978); for the larva, those of Anderson (1947) and Ahmad and Burke (1972), and May (1977) for the alimentary canal. Pupae 289 were examined in alcohol with a binocular dissecting microscope. Larvae were pre- pared for examination as in Ahmad and Burke (1972). They were examined in gly- cerine in depression slides with a compound microscope with interference contrast. DESCRIPTION OF LAST INSTAR LARVA Figs. 1-6 Body (Fig. 1).—Robust, slightly curved. Asperites minute, various in degree of de- velopment, apices acutely pointed, evenly distributed over body except in areas sur- rounding bases of setae and in primary folds of body cuticle. Color creamy white except for head (medium to dark brown), prono- tum (pale yellowish-brown), prothoracic pleural area and meso- and metanotal pedal areas (very pale golden yellow). Head (Figs. 2—5).—Free. Mouthparts di- rected ventrally. Width of head capsule 1.44— 1.59 mm (n = 4). Frontal sutures distin- guishable throughout length, incomplete anteriorly. Epicranial suture distinct throughout length, length approximately one-half length of head capsule. Anterior ocellus small, indistinct; posterior ocellus absent. Basal article of antenna bearing tall, circular, cone-like accessory projection and three or four minute setae. Endocarina dis- tinct, approximately one-half length of frons. Hypopharyngeal bracon present. Frons with five pairs fine, apically tapered, frontal setae (fs 1-5); fs 1 slightly shorter than fs 2, 4-5 which are subequal in length; fs 3 very short. Two pairs frontal sensillae (fsl 1-2); fsl 1 situated near midline between fs | and fs 2; fs] 2 located equidistant between fs 3 and fs 4, immediately posteriad of line joining fs 3 and fs 4. Five pairs dorsal epicranial setae (des 1-5); des 1, 3, 5 very long, tapered api- cally, subequal in length; des 2, 4 slightly shorter, subequal in length; des | situated slightly posterolaterad of confluence of epi- cranial and frontal sutures; des 2 situated distinctly laterad, slightly posteriad of des 1: des 4 situated slightly posterolaterad of des 3, des 3 situated very close to frontal 290 Figs. 1-6. suture. Four pairs very small, peg-like, post- epicranial setae (pes 1—4) arranged in an arc directed from vertex to des 2. Three pairs dorsoepicranial sensillae (desl 1-3); desl 1 situated posteriad of des 1, slightly anteriad of pes 1; desl 2 situated slightly laterad of des 1, mediad of des 2, slightly closer to des 1 than to des 2; desl 3 situated posteriad of des 5. Two pairs lateral epicranial setae (les 1-2); les 1 shorter than les 2; les 2 subequal in length to des 1, 4-5. Two pairs ventral PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Last instar larva of Cleonidius erysimi (Fall). 1, lateral view of larva (scale line = 1.75 mm). 2, frontal view of head (scale line = 0.5 mm). 3, ventral view of epipharynx (scale line = 0.1 mm). 4, dorsal view of left mandible (scale line 0.1 mm). 5, ventral view of maxillae and labium (scale line = 0.1 mm). 6, lateral view of alimentary canal (scale line = 1.5 mm). epicranial setae (ves 1—2); moderately long, subequal in length. Clypeus with two pairs short, stout, clypeal setae (cls 1-2); cls 1 longer than very short cls 2; also with single pair clypeal sensillae (clsl) close to anterior margin of frons. Labrum with two pairs short, stout, labral setae (Ims 1-2); Ims 1 longer than very short Ims 2; Ims 2 situated laterad and slightly anteriad of Ims 1; also with one pair lateral labral sensillae (Imsl) situated between Ims | and Ims 2; and single VOLUME 93, NUMBER 2 median labral sensilla (mlmsl) situated on midline at anterior clypeal margin. Epi- pharynx with six or seven very short, peg- like, anteromedian setae (ams); and four pairs short, stout anterolateral setae (als). Labral rods well-developed, convergent posteriorly; with two pairs short, stout, epi- pharyngeal median spines (msp 1-2); msp 1 longer and stouter than msp 2; single pair epipharyngeal sensory pores situated be- tween msp 1 and msp 2. Mandibles with single apical tooth and with two mandibular setae (mds 1-2); mds 1 longer than very short, stout mds 2. Maxillary palpus with two subequal in length articles; apical article with number of small, cone-like projections at apex. Stipes with four setae (sts 1-4); sts 1, 3-4 moderately long, subequal in length; sts 2 very short. Mala with six stout dorsal (dmsa) and five linearly arranged stout ven- tral setae (vsma). Labial palpus of two ar- ticles. Premental sclerite with long anterior and posterior processes. One pair long, slen- der, premental setae (prms); three pairs short, stout, glossal setae (gls) and numerous minute asperites along apical margin. Post- mentum with three pairs setae (pms 1-3); pms | very long, tapered apically; pms 2 shorter than pms 1; pms 3 shorter than pms 2; pms 2 and pms 3 situated close together in anterolateral angle of postmentum. Prothorax (Fig. 1).—(Setae described for one side of body). Pronotum with six mod- erately long setae arranged around perim- eter of darkly sclerotized area. Spiracular area with two setae, anterior seta shorter than posterior. Spiracular bicameral, air- tubes short, each with four or five annula- tions, peritreme elongate-oval in shape. Pleural region with two moderately long se- tae. Pedal area with seven setae arranged around perimeter of lightly sclerotized area. Meso- and metathorax (Fig. 1).—(Setae described for one side of body). Prodorsum with single short seta (prs). Postdorsum with four setae (pds 1-4) arranged in a transverse line; pds 3-4 slightly longer than pds 1-2. Spiracular area with four short setae ar- 291 ranged in a longitudinal line. Epipleural re- gion with single short seta (eps). Pleural re- gion with single moderately long seta. Pedal area with six short setae arranged around perimeter of lightly sclerotized area. Ster- nellum with single short setae. Most setae (excluding pronotal setae and pds 3-4) sub- equal in length. Abdomen (Fig. 1).—(Setae described for one side of body). Abdominal segments I- VII each with three folds; segment VIII with two folds. Prodorsum of each segment with single short seta (prs). Postdorsum with five setae (pds 1-5) arranged in transverse line on abdominal segments I-VI; pds 1-5 short, subequal in length on segments I-IV; pds 1-2, 4 longer than pds 3, 5 on segments V— VI. Abdominal segment VII with seven postdorsal setae (pds 1-7); pds 4, 6 shortest of series, pds 1-2, 4 subequal in length. Spi- racles bicameral, present on abdominal seg- ments I-VIII; peritreme circular, smaller than on prothorax; airtubes short, as long as diameter of peritreme, each with four or five annulations. Spiracular area with two setae (ss 1-2); ss | shorter than ss 2. Epi- pleural region with two short setae (ss 1-2); ss 1 shorter than ss 2. Epipleural region with two short setae (eps 1-2); eps 2 slightly lon- ger than eps |. Pleural region with two short setae (ps 1-2); ps 2 slightly longer than ps 1. Pedal area with single short seta. Euster- num with two short setae (eus 1-2). Ab- dominal segment IX with five short dorsal setae; one short epipleural seta; and, one short eusternal seta. Anus terminal. Alimentary canal (Fig. 6).—Proventric- ulus short, numerous small mycetomes present around cardiac valve. Anterior ven- triculus large, deeply folded with numerous variously sized and shaped protruding lobes, narrowed gradually to posterior ventriculus. Gastric caecae small, not numerous. Six malphighian tubules, grouped 4 + 2 on thickened portion of posterior ventriculus, rejoined to alimentary tract in uniform manner distant from rectal bracon and ap- pressed for some length to well-developed, 292 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ap ay —\ fet > " TF j§ ff tats 9 * 1 t 7 > T9-y—7_t__y—y—I-7—¥ 5 t ’ Y--y-"- Rema aan Tt a (ery tr—vyKi] , ? ? 7 % a Pn Pol} ca on of & = AP 799 Tag NeIN ( \ ve PMT itt rs ~ peal Ve i \ aX Spe \ ea en”, An i NA k= at 10 Figs. 7-10. Pupa of Cleonidius erysimi (Fall). 7, frontal view of head. 8, dorsal view of pronotum and mesonotum. 9, dorsal view of metanotum and abdomen. 10, ventral view of abdomen. Scale line = 1.1 mm. asymmetrically expanded cryptonephrid- ium. Rectal bracon membranous, distinctly delimiting posterior end of cryptonephrid- ium. Cryptonephridrium much longer than very short anal tube. Specimens examined.—USA. CA. San Bernardino Co., Guadalupe, Los Alamos, 20.vil.1987, ex. crowns Lobularia mariti- ma (L.) Dev. (Cruciferae), T. Seeno & J. Davidson (15 specimens). USA. CA. San Bernardino Co., Harris Ranch, Los Alamos, 25.vii.1986, ex. crowns Lobularia maritima (L.) Dev. (Cruciferae), J. Davidson (2 spec- imens). USA. CA. San Bernardino Co., Harris Ranch, Los Alamos, 14—21.vii.1987, ex. crowns and roots Lobularia maritima (L.) Dev. (Cruciferae) (1 specimen). Species identity was established based on co-occur- rence of adults with larvae and pupae in mines in crowns and roots of the host plants. Specimens are deposited in CMNC, TAMU, USNM. DESCRIPTION OF PUPA Figs. 7-10 Body.— Length 9.6-13.0 mm (n = 9). Head (Fig. 7).—Creamy white except for compound eye and mandibles which are light to dark brown in color in some spec- imens. All setae sessile, fine, tapered api- VOLUME 93, NUMBER 2 cally. One pair moderately long vertical se- tae (vs), subequal in length to each of two pairs supraorbital setae (sos 1-2) and pair orbital setae (os 1). One pair “‘ocular” setae situated on posterior portion of ocular area subequal in length to, or shorter than, each of vs, sos 1-2 and os 1. Five pairs postan- tennal setae (pas 1-5); pas 1-4 subequal but various in length, successively more distal in position; pas 5 situated laterad of, and slightly shorter than, pas 4. Rostrum short, extended only to base of prothoracic tarsi. One pair short to moderately long rostral setae (rs 1) situated immediately anterior to site of antennal attachment, subequal in length to pair lateral rostral setae (rls 1); rls 1 borne on distinct lateral tubercle at an- terolateral angle of rostrum. Mandible with one pair very short, fine, mandibular setae (ms) at extreme anterolateral angle of man- dible. Prothorax (Fig. 8).—Pronotal setae ses- sile, fine, tapered apically; straight to slight- ly medially or anteriorly apically curved. One pair very long apical setae (aps), sub- equal in length to each of two pairs antero- lateral setae (als 1-2), pair of lateral setae (Is 1), and lateralmost two of four pairs pos- terolateral setae (pls 3-4). Two pairs short to moderately long discal setae (ds 1-2) sit- uated approximately equidistant from aps and pair moderately long basal setae (bs); ds | slightly shorter than ds 2 on most spec- imens, situated slightly posteromediad of ds 2. Aps, ds 2 and bs situated in longitudinal linear manner. Two pairs very long antero- lateral setae (als 1-2); als 1 situated mediad and slightly anteriad of als 2. One pair long lateral setae (Is 1) situated posterolaterad of als 2 and anterolaterad of pls 4. Four pairs posterolateral setae (pls 1-4); pls 3-4 very long; pls 4 situated moderately anterolat- erad of pls 3; pls 2 shorter than pls 1, 3-4, situated slightly mediad of pls 3; pls 1 mod- erately long, situated approximately equi- distant from pls 2 and bs, laterad and var- ious from slightly anteriad to slightly posteriad of bs. 293 Mesonotum (Fig. 8).—Six pairs short to moderately long, fine, straight, sessile meso- notal setae (msns 1-6); msns 4, 6 subequal in length, longest of series. Metanotum (Fig. 9).— Five pairs short to moderately long, fine, straight, sessile meta- notal setae (mtns 1—5); mtns 1-2 short, sit- uated very close together in anterior one- half of metanotum; mtns 3-5 longer than mtns 1-2, situated equidistant from each other in slightly anteriorly directed trans- verse linear manner. Abdomen (Figs. 9—10).—Abdominal seg- ment I with four pairs very short, very fine, sessile postdorsal setae (pods 1-4); two pairs short, fine laterodorsal setae (Ids 1-2), the latter each borne on summit of very small, rounded, laterally directed tubercle. Abdom- inal segment II with five pairs very short, fine postdorsal setae (pods 1-5), each seta borne on summit of very small, rounded, posteriorly directed tubercle; two pairs short, fine laterodorsal setae (Ids 1—2); one pair very short, fine, prodorsal setae (prds 1); and two pairs moderately long spiracular setae (ss 1- 2), each borne on summit of small rounded tubercle, the bases of tubercles narrowly con- tiguous. Abdominal segments HJ—VII each with six pairs short postdorsal setae (pods I- 6); pods 1-2, 4, 6 short, very stout, darkly sclerotized, each borne on summit of apically darkly sclerotized, posteriorly directed, rounded tubercle; pods 3, 5 short to mod- erately long, fine, sessile or situated on sum- mit of small, unsclerotized tubercle; two pairs short to moderately long laterodorsal setae (Ids 1-2), each borne on summit of small rounded tubercle, Ids | approximately one- half length Ids 2, Ids 1 and Ids 2 more closely situated on successively more posterior ab- dominal segments; one pair small, fine, ses- sile prodorsal setae (prds 1); and two pairs long to very long, fine spiracular setae (ss 1- 2) each borne on summit of laterally directed rounded tubercles, tubercles slightly larger and bases increasingly contiguous on suc- cessively more posterior abdominal seg- ments. Abdominal segment VIII with five 294 pairs postdorsal setae (pods 1-5); pods 1, 3, 5 short, very stout, darkly sclerotized, each situated on summit of moderately large, pos- teriorly directed, darkly sclerotized tubercle; pods 2, 4 long, fine, each situated on summit of small, rounded tubercle; one pair prodor- sal setae (prds); two pairs laterodorsal setae (Ids 1-2) situated very close to each other; and two pairs long, fine, spiracular setae, slightly shorter in length than those on ab- dominal segment VII. All setae very slightly increased 1n length on successively more pos- terior abdominal segments unless otherwise noted. Ventrally with two pairs short, fine, sessile lateroventral setae (lvs 1-2) on each of abdominal segments IV—VII; and three pairs transversely linearly arranged, very short, fine, sessile midventral setae (mvs 1|- 3). Abdominal segment VIII ventrally with single pair short, fine, sessile lateroventral setae (Ivs) and single pair short, fine, sessile midventral setae (mvs). Abdominal segment IX with pair of large, apically acuminate, darkly sclerotized, posteriorly directed pro- cesses (pp), each process with three or four moderately long, fine setae around and an- terior to base, each seta borne on summit of variously developed, small, rounded tuber- cle. Basal region of processes to apical mar- gin abdominal segment VIII slightly ven- trally swollen. Specimens examined.—USA. CA. Santa Barbara Co., Guadalupe, Los Alamos, 20.vil. 1987, ex. crowns Lobularia maritima (L.) Dev. (Cruciferae), T. Seeno & J. David- son (4 specimens). USA. CA. Santa Barbara Co., Harris Ranch, Los Alamos, 14— 21.vui.1987, ex. crowns Lobularia maritima (L.) Dev. (Cruciferae) (8 specimens). Speci- mens are deposited in CMNC, TAMU, USNM. NATURAL HISTORY This species is widely distributed throughout western North America from British Columbia east to Manitoba, south to southern California, Arizona and ex- treme west Texas (Anderson 1988). Adults PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of Cleonidius erysimi have been collected from a variety of plants, most commonly Cruciferae, in a variety of habitats through- out the species range (Anderson 1988). Al- though Cruciferae were suspected as the lar- val host due to the abundance of associations of adults with plants of this family, the as- sociation of the immature stages with Lob- ularia maritima (L.) Desv. is the first con- firmed larval host record. Lobularia maritima is an introduced, low, woody shrub of Mediterranean origin which is a common escapee from gardens; it generally grows along roadsides or in wasteplaces in Cali- fornia (Munz 1974). Lobularia maritima is cultivated for its seeds which are used in flower beds. Although seeds develop on in- fested plants, the plants are forced into early maturation and therefore, produce fewer seeds. Cleonidius erysimi adults and immature stages were found associated with commer- cially planted L. maritima in Santa Barbara County, California. Natural vegetation in the areas surrounding the infestations was checked but no C. erysimi adults or im- mature stages were found. Infested plant- ings were in one-half to ten acre lots. Most infestations involved 10-15% of the plants but in some lots up to an estimated 70% of the plants were involved. Most infestations were dense nearest the service roads with evidence of activity getting lighter toward the center of the planting. One larva was present per plant and occupied the entire area inside the crown. Pupation takes place in the larval mine. Plants with purple flow- ers appeared to have the heaviest infesta- tions. Larvae mine the roots and crowns of the plants; pupation takes place in the larval mine. THE PHYLOGENETIC POSITION OF LIXINI Anderson (1988) summarized evidence, albeit symplesiotypy, to suggest Lixini (sen- su Kuschel) as a primitive lineage within Curculionidae but otherwise could not re- VOLUME 93, NUMBER 2 solve their precise relationships further. Such a proposal was made solely on the basis of character state distribution in adults. Whereas precise relationships are still left as unresolved, based on the present descrip- tions of the immature stages of C. erysimi, examination of pupae of L. scrobicollis (per- sonal observation), and descriptions and figures of pupae of Lixini (sensu Kuschel) in Scherf (1964), some additional evidence bearing on this topic can now be considered. The traditional division of Curculionidae into Adelognatha (““broad-nosed weevils’’) and Phanaerognatha (“‘long-nosed wee- vils’’) carries with it the implication that the two groups are sister-taxa. This is clearly not so, and as a result, the resolution of phylogenetic relationships in Curculionidae iS proving to be a very complex problem, one brought about largely by conflicting character state distributions in a number of presumed primitive curculionid groups. Adelognatha appear to represent a mono- phyletic group based on a number of apo- typic features; in adults, the deciduous man- dibular cusp and resulting mandibular scar, and in larvae, the accessory sensory ap- pendage of the antenna distinctly wider than tall and the frons lacking an endocarina. May (1970, 1978) suggested that the occurrence of mandibular setae in pupae of only Ade- lognatha was thus associated with the pres- ence of the deciduous cusp in the adult. Ab- sence of the deciduous mandibular cusp in adults of species of Pantorhytes (Pachy- rhynchina), but the presence of mandibular setae in pupae, suggested the secondary loss of the cusp in this taxon (May 1978). Man- dibular setae, however, are now known in pupae of Jthycerus noveboracensis (Forster) (Sanborne 1981), Thecesternus hirsutus Pierce (McClay and Anderson 1985), Cleonidius erysimi (Fall) and Lixus scrobi- collis (Boheman) (personal observation), and Cleonis pigra (Scopoli) (Scherf 1964). Adults of each of these species do not possess a deciduous mandibular cusp. If absence of the cusp is truly a unique secondary loss (as 295 suggested by May 1970, 1978) and presence of mandibular setae is apotypic, their pres- ence in the pupa suggests inclusion of these taxa in a monophyletic group, sister to the paraphyletic traditional Adelognatha. If, however, it is not assumed that the man- dibular setae of pupae are associated with the presence of the deciduous cusp in the adult, then the alternative proposal, that presence of mandibular setae in the pupa is an independent apotypic feature, can be considered. Such would result in Adelo- gnatha, Ithycerus noveboracensis, Pachy- rhynchina, Thecesternini and Lixini (sensu Kuschel) as a monophyletic group but with relationships further unresolved. Either of these arrangements is fraught with prob- lems and thus it appears best if a third al- ternative is considered: that the presence of mandibular setae in the pupa is a plesiotypic character state, not related to the presence of deciduous cusps in the adults. Kuschel (in press) has recently proposed the unification of traditional Adelognatha with Ithycerus noveboracensis (Forster) and a number of other presumed primitive Cur- culionidae (including Pachyrhynchina and Thecesternini among others, but not Lixi- ni), as the Brachycerinae. Inclusion of Jthy- cerus noveboracensis is at least in part based on the presence of mandibular setae in pu- pae of this species as well as a number of other features of the adult and immature stages (Kuschel in press). That pupae of Lix- ini (sensu Kuschel) are now known to pos- sess mandibular setae suggests, if the pres- ence of the setae were considered apotypic, inclusion of this taxon in Brachycerinae (sensu Kuschel in press); however, among other characters used by Kuschel (in press) to define Brachycerinae, some support this placement whereas others dispute it. Adult Lixini agree with the Brachycerinae (sensu Kuschel in press) in having the epistoma raised in relation to the surrounding area (although this is not so in some Lixus spe- cies and may be correlated with increased overall rostrum length), mandibles pluri- 296 setose, underside of elytra lacking stridu- latory files, sclerolepidia absent, and bladal part of sternite 9 of male extensively pig- mented. Adult Lixini disagree in not having the proximal and distal hemisternites (gono- coxites I and II) of the female distinct (proximal hemisternite [gonocoxite I] is lacking), fore coxae placed posteriorly on the prosternum, and in the rostrum being sexually dimorphic (although this is not true for many traditional Cleonina) and where natural history information is available, in being used (in females of at least some Lixus species) to excavate the oviposition site. As far as larval characters are concerned, Lixini disagree in that des | is not situated in the dorsoepicranial suture or on the frons, and, where natural history information is avail- able, they are known to be endophytic in roots and stems. Anderson (1988) was unable to suggest a sister-group relationship between Lixini (sen- su Kuschel) and any other Curculionidae and suggested a primitive phylogenetic position in Curculionidae, apparently near Molytini, but cautioned that this could not be further resolved as it was based solely on symplesi- otypy. The now apparent phylogenetic posi- tion near traditional Adelognatha is not in- consistent with this previous suggestion of a primitive phylogenetic position for Lixini, but disputes the claim of a close relationship to Molytini. At present, Lixini as a Curculionini (sensu Kuschel in press) is supported by only one apotypic character; that of not hav- ing the proximal and distal hemisternites (gonocoxites I and II) of the female distinct (proximal hemisternite [gonocoxite I] is lack- ing). On the other hand, apotypic characters suggesting Lixini as Brachycerinae are: bladal part of sternite 9 of male extensively pig- mented, and, for some of the species, having the epistoma raised in relation to the sur- rounding area and the rostrum not sexually dimorphic. At present Lixini are best left as Curcu- lioninae in the new scheme of Kuschel (in press), but special note should be taken of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON their heretofore unrecognized close and per- haps transitional relationship with Brachy- cerinae. ACKNOWLEDGMENTS I thank Terry N. Seeno of the California State Department of Food and Agriculture and Jerry W. Davidson, Santa Barbara County Department of Agriculture, for the larvae and pupae of C. erysimi and for in- formation on the host plant association and natural history. I also acknowledge the use of Dr. Kuschel’s soon to be published new classification scheme for Curculionoidea (see Literature Cited). LITERATURE CITED Ahmad, M. and H. R. Burke. 1972. Larvae of the weevil tribe Anthonomini (Coleoptera: Curculion- idae). Miscellaneous Publications of the Ento- mological Society of America 8(2): 31-81. Anderson, R. S. 1988. Systematics, phylogeny and bio- geography of New World weevils traditionally of the tribe Cleonini (Coleoptera: Curculionidae; Cleoni- nae). Quaestione Entomologicae 23: 431-709. Anderson, W. H. 1947. A terminology for the ana- tomical characters useful in the taxonomy of wee- vil larvae. Proceedings of the Entomological So- ciety of Washington 49: 123-132. Boving, A. G. and F. C. Craighead. 1930. An illustrated synopsis of the principal larval forms of the order Coleoptera. Entomologica Americana 11: 1-351. Burke, H. R.and D. M. Anderson. 1976. Systematics of larvae and pupae of American Curculionoidea: Status report, historical review and bibliography. Southwestern Entomologist 1: 56-73. Kuschel, G. (In press). A phylogenetic classification of Curculionoidea to family and subfamily level. Memoirs of the Entomological Society of Wash- ington. May, B. M. 1970. Immature stages of Curculionidae of Heard Island. Pacific Insects Monographs 23: 261-270. 1977. Immature stages of Curculionidae: Larvae of the soil-dwelling weevils of New Zea- land. Journal of the Royal Society of New Zealand 7: 189-228. . 1978. Immature stages of Curculionidae (Co- leoptera): Some species in the genus Pantorhytes Faust (Pachyrhynchini) from the Papuan Region and phylogenetic implications of certain charac- ters. Journal of the Australian Entomological So- ciety 17: 351-360. McClay, A. S. and D. M. Anderson. 1985. Biology VOLUME 93, NUMBER 2 and immature stages of Thecesternus hirsutus Pierce (Coleoptera, Curculionidae) in north-eastern Mex- ico. Proceedings of the Entomological Society of Washington 87: 207-215. Munz, P. A. 1974. A Flora of Southern California. University of California Press, Berkeley, Califor- nia. 1086 pp. O’Brien, C. W. and G. J. Wibmer. 1982. Annotated checklist of the weevils (Curculionidae sensu lato) of North America, Central America, and the West In- dies (Coleoptera: Curculionoidea). Memoirs of the American Entomological Society 34: x + 382 pp. Peterson, A. 1951. Larvae of insects. Part II. Cole- optera, Diptera, Neuroptera, Siphonaptera, Me- 297 coptera, Trichoptera. Edward Bros. Inc., Ann Ar- bor, Michigan. 146 pp. — Sanborne, P. M. 1981. Biology of Ithycerus novebo- racensis (Forster) (Coleoptera) and weevil phylog- eny. Evolutionary Monographs 4: 1-80. Scherf, H. 1964. Die Entwicklundsstadien der mitteleu- ropaischen Curculioniden (Morphologie, Bionomie, Okologie). Abhandlungen der senckenbergischen na- turforschenden Gesellschaft 506: 1-335. . 1978. Bestimmungstabelle fur die Larven der Curculionidae (partim), pp. 344-367. Jn Klaus- nitzer, B. ed., Ordung Coleoptera (Larven). Dr. W. Junk bv. Publishers, The Hague, Netherlands. PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 298-316 WEST INDIAN COCCINELLIDAE IV (COLEOPTERA): NEW GENERA AND SPECIES OF STICHOLOTIDINI ROBERT D. GORDON Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, % U:S. National Museum of Natural History Washington, D.C. 20560. Abstract. — Five new genera of West Indian Sticholotidini are described. Nelasa, n. gen., includes the new species N. erugonota, N. beckeri, N. iricolor, N. howdeni, N. duncansi, N. schwarzi, N. cubensis, N. haitiensis, and N. dominicensis. Paranelasa, n. gen., includes the new species P. jamaicensis and P. polita. Semiviride, n. gen., includes the new species S. loisobrienae and S. portoricensis. Neaptera, n. gen., includes the new species N. purpurea, N. viridissima, and N. viola. Nexophallus korschefskyi Duverger is transferred to Neaptera. Parinesa, n. gen., is established for P. whiteheadi, n. sp. Key Words: The classification of the Western Hemi- sphere Sticholotidinae was revised by Gor- don (1977). At that time the subfamily was composed of four tribes with one additional tribe described later (Gordon et al. 1989). Four genera were recognized in the tribe Sticholotidini, only one of these from the West Indies. Five West Indian genera dis- covered subsequently are described here and the key to genera modified accordingly. A major change in the definition of the tribe (Gordon 1977) is necessitated by the pres- ence of six abdominal sterna in three of the genera described herein. All previously known members of Sticholotidini had five visible sterna. These minute to moderate sized cocci- nellids occur worldwide but only recently has the extent of Western Hemisphere spe- ciation been appreciated. This contribution is another in the ongoing process of making known the entire New World fauna of Sti- cholotidinae. Type material is deposited in the Cana- dian National Collection, Ottawa (CNC), the U.S. National Museum, Washington Sticholotidinae, Sticholotidini, West Indies (USNM), the Natural History Museum, De- partment of Zoology, Vienna, Austria (NHMA), the Carnegie Museum, Pitts- burgh (CM), M. Ivie, Montana State Uni- versity, Bozeman (MI), and the University of California, Berkeley (UCB). KEY TO GENERA OF New WORLD STICHOLOTIDINI 1. | Epipleuron foveate for reception of femur; eye divided by genal extension; anterior tibia broadly expanded, externally dentate or not ~ Epipleuron not foveate for reception of fe- mur; eye not divided; anterior tibia simple, unmodified) 425 cee ere 3 2(1). Pronotum with sparse, fine pubescence; ex- ternal margin of anterior tibia not angulate SN Rn oe oe ERR LR Glomerella Gordon - Pronotum without pubescence; external margin of anterior tibia sharply angulate. (Fig. 14) ee see nk ee cee Parinesa, n. gen. . Terminal segment of maxillary palpus long, slender, apically acuminate et, WL cary ane cr ase Nexophallus Gordon - Terminal segment of maxillary palpus elon- gate or not, not apically acuminate ....... 4 4(3). Postcoxal line on Ist abdominal sternum complete (Rigs8)) saeco 5 VOLUME 93, NUMBER 2 - Postcoxal line on 1st abdominal sternum in- complete (Fig. 15) Eye large, finely faceted; abdomen 5-seg- mented) sc, soe we ete eee Nesina Gordon - Eye small, coarsely faceted; abdomen 6-seg- MENTEGEN Aree artes Neaptera, n. gen. Postcoxal line on 1st abdominal sternum ex- tended parallel to posterior margin of ster- num in apical 2; abdomen 5-segmented ... 7 - Postcoxal line on Ist abdominal sternum short, not parallel to posterior margin of Ist abdominal sternum; abdomen 6-segmented 8 Dorsal surface smooth, polished; lateral margin of prosternal process not ridged SO Oa Bee ae Oar one aoe one Neotina Gordon - Dorsal surface roughly sculptured, feebly shiny; lateral margin of prosternal process heavily ridged! 7.04 .%,.6.l4c Semiviride, n. gen. Terminal segment of maxillary palpus elon- gate (Fig. 2); antennal club elongate (Fig. 6) Sp Bo On eee ere er Paranelasa, n. gen. - Terminal segment of maxillary palpus short (Fig. 1); antennal club short, broad (Fig. 5) eA RL Se eh = tt ee ae SE Nelasa, n. gen. 7(6). 8(6). Nelasa, NEw GENUS Sticholotidini with form generally oval; without pubescence except on clypeus, lat- eral margin of head near eye; dorsal surface with metallic sheen throughout, or at least on elytron; punctation on elytron much coarser than on head, pronotum. Head broad; clypeus short, truncate apically, an- terior angle rounded. Eye coarsely faceted, small; eyes separated by 4 times width of eye; gena slightly extended onto eye. An- tenna 10-segmented; club 3-segmented, short, broad (Fig. 5); insertion mostly ex- posed. Terminal segment of maxillary pal- pus short, barrel-shaped, slightly narrowed apically (Fig. 1). Epipleuron narrow, de- scending externally, not foveate for recep- tion of femoral apices. Prosternum with coxae narrowly separated by protuberant, rectangular process, process carinate on each side, carinae joined at apex (Fig. 10). Leg with femur robust, shallowly grooved for reception of tibia; tibia slender, shorter than femur; tarsus cryptotetramerous, tarsal claw simple, lacking tooth. Abdomen with 6 vis- ible sterna; 1st sternum with postcoxal line short, incomplete (Fig. 15); apex of male 5th 299 sternum broadly rounded, apex of female 5th sternum strongly, abruptly rounded. Male genitalia symmetrical; basal lobe lon- ger than paramere. Female genitalia with unmodified spermathecal capsule (Fig. 191); without infundibulum. Membranous wing present. Type species: Nelasa beckeri, new species. Nelasa is most similar to Neotina Gordon but Neotina differs by having the prosternal process short, broadly triangular, without lateral carinae, and the postcoxal line on the first abdominal sternum not joining hind margin of sternum. Ne/asa also resembles Paranelasa, n. gen., see remarks under that genus. Nine species of Nelasa are known thus far: five from Jamaica, two from Cuba, and two from Hispaniola. Surprisingly, no specimens have been seen from Puerto Rico or the Virgin Islands, but the genus is prob- ably represented there also. Ne/asa species are difficult to differentiate externally but the male genitalia provide excellent diag- nostic characters. The female genitalia, where known, are essentially identical in all species, therefore only those of N. beckeri are illustrated. Food and habitat preferences for this genus are unknown; most of the available specimens were collected by beat- ing vegetation. The generic name is an arbitrary com- bination of letters and the gender is femi- nine. KEY TO SPECIES OF NELASA 1. | Species known only from Jamaica Species not known from Jamaica Head, pronotal surface polished, shiny ... ea seers yada: Wyk ebetles kanes Se poke: erugonota, Nn. sp. - Head, pronotal surface dull or feebly shiny, with alutaceous sculpture Dorsal surface with strong, metallic greenish sheen; head, pronotum feebly shiny, surfaces feeblyalutaceous)) 2.442. 4: beckeri, n. sp. - Dorsal surface with purple or copper sheen, or if sheen greenish then base color brown; head, pronotum dull, surfaces strongly alu- taceous 4(3). Elytron brown with feeble greenish sheen .. . duncansensis, Ni. sp. 300 - Elytron black with purple or coppery sheen Rina RE PENT ee Tho Sue cabils borat 5 5(4). Elytron with purple tint not mixed with cop- pery Sheen. 80: s-.5ece5 eee howdeni, n. sp. - Elytron with intermixed purple, coppery Sheen? 2152 Manet ee one eee iricolor, n. sp. 6(1). Species known only from Cuba ........... 7 Species known only from Hispaniola ...... 8 7(6). Head, pronotal surfaces as smooth, polished as suriace ol elytronis a4 ae cubensis, n. sp. - Head, pronotal surfaces dull, alutaceous; surface of elytron shiny, with only trace of alutaceous sculpture .......... schwarzi, n. sp. . Elytron brown with metallic greenish sheen; surface of head, pronotum shiny, feebly alu- (ACCOUSMe rer rises. cas dominicensis, Nn. sp. - Elytron black with metallic greenish sheen; surface of head, pronotum dull, strongly alu- TACCOUS BORN EA: rhe acs aes haitiensis, n. sp. Nelasa erugonota, NEw SPECIES Description: Holotype male, length 1.5 mm, width 1.2 mm. Form oval. Color black with strong, blue metallic sheen on dorsal surface, equally strong throughout; mouth- parts, antenna, tibia, tarsus yellow; femur, ventral surface dark brown. Head smooth, polished, finely punctured, punctures sep- arated by about a diameter. Pronotum smooth, polished, punctures equal in size to head punctures, separated by 2 to 4 times a diameter. Elytron smooth, polished, punc- tures coarser than on head, separated by about twice a diameter. Type material: Holotype; Jamaica, Port- land, Silver Hill Gap, 22.VIII.1980, 1000- 1100 m, A. Norrbom (CM). Remarks: The smooth, polished head and pronotum of N. erugonota distinguish it from the other known Jamaican species. The male genitalia were not dissected to avoid damaging the only available specimen. The specific name is from the Latin erugo, mean- ing smooth, and refers to the smooth pro- notal surface. Nelasa beckeri, NEw SPECIES Description: Holotype male, length 1.6 mm, width 1.3 mm. Form oval. Color black with strong, green, metallic sheen on dorsal surface, particularly strong on elytron; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mouthparts, antenna, tibia, tarsus brownish yellow; ventral surface black; femur brown. Head feebly shiny, surface alutaceous, finely punctured, punctures separated by about a diameter. Pronotum feebly shiny, surface finely alutaceous, punctures slightly coarser than on head, separated by less than to twice a diameter. Elytron smooth, polished, mod- erately coarsely punctured, punctures much larger than on pronotum, separated by less than to twice a diameter. Genitalia with sides of basal lobe somewhat sinuate in ventral view; sipho short, unmodified (Figs. 19a- Cc) Allotype: Length 1.8 mm, width 1.4 mm. Similar to male except genitalia as in Figure 191. Variation: Length 1.5 mm to 1.8 mm, width 1.3 to 1.4 mm. Type material: Holotype; Jamaica, Hard- war Gap, 4000’, VII.10.1966, Howden & Becker (CNC). Allotype; same data as ho- lotype except date VII.4.1966 (CNC). Para- types, total 16; 5, same data as holotype; 7, same data as holotype except additional dates VII.5.1966, VII.11.1966, VII.16.1966. VII.18.1966, VII.25.1966; 3, Jamaica, Try. Barbecue Bottom, VII.6.1966, VII.13.1966, H. Howden; 1, Jamaica, Shirley Castle, VII.1971, on Pinus caribae. (CNC) (USNM). Remarks: The strong, green metallic sheen on the dorsal surface, feebly shiny prono- tum and head distinguish N. beckeri from the other Jamaican species. The elytral punctures are also less coarse than in the other species. The species is named for Ed Becker, one of the collectors of the type series and an eminent coleopterist. Nelasa iricolor, NEw SPECIES Description: Holotype male, length 1.5 mm, width 1.3 mm. Form oval, slightly rounded. Color black, elytron with purple, coppery metallic sheen, pronotum with green, metallic copper sheen, head with pur- ple metallic sheen; mouthparts, antenna, tibia, tarsus brownish yellow; ventral sur- VOLUME 93, NUMBER 2 301 Figs. 1-12. 1-4, maxillary palpus: 1, Nelasa iricolor, 2, Paranelasa jamaicensis: 3, Semiviride loisobrienae; 4, Neaptera purpurea: 5-8, antenna: 5, Nelasa iricolor; 6, Paranelasa Jamaicensis; 7, Semiviride loisobrienae: 8. Neaptera purpurea: 9-12, prosternum: 9, Nelasa duncansensis; 10, Paranelasa Jamaicensis; 11, Semiviride loisobrienae; 12, Parinesa whiteheadi. 302 eS at Figs. 13-18. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 18 13, 14, Parinesa whiteheadi: 13, facial view; 14, protibia: 15-18, lst abdominal sternum: 15, Nelasa iricolor; 16, Paranelasa jamaicensis; 17, Semiviride loisobrienae; 18, Neaptera purpurea. face mostly yellowish brown; femur, ab- domen brown. Head dull, surface strongly alutaceous, finely punctured, punctures sep- arated by about a diameter. Pronotum dull, surface strongly alutaceous, punctures equal in size to head punctures, separated by | to 2 times a diameter. Elytron smooth, pol- ished, coarsely punctured, punctures much larger than on pronotum, separated by about a diameter. Genitalia with sides of basal lobe not sinuate, broad in lateral view; sipho broken (Figs. 19d—f). Type material: Holotype; Jamaica, St. And. St. Peters, VII-7-1966, Howden & Becker collectors (CNC). Other specimens: 1, same data as holo- type; 3, Jamaica, Port., Port. Antonio, VII. 1- 7-1966, E. C. Becker (CNC). Remarks: All specimens except the ho- lotype are females tentatively considered conspecific with the holotype. They differ slightly in color and punctation from N. iri- color, more so from the other Jamaican spe- cies, and they may represent another un- described species. However, males are needed to decide this. Nelasa howdeni, NEw SPECIES Description: Holotype male, length 1.6 mm, width 1.25 mm. Form oval. Color black, elytron with metallic purple sheen; mouthparts, antenna, coxa, trochanter, tar- sus brownish yellow; femur, tibia dark brown. Head, pronotum extremely dull, punctures very fine, barely perceptible. El- ytron smooth, polished, coarsely punctured, VOLUME 93, NUMBER 2 punctures separated by a diameter or less. Genitalia with basal lobe slender in lateral view, paramere slender; sipho broken (Figs. 19g, h). Type material: Holotype; Jamaica, Hard- war Gap, 4000’, VII.21.1966, Howden & Becker (CNC). Remarks: The holotype is the only spec- imen examined. It was collected at Hardwar Gap where most of the type series of N. beckeri was taken. In addition to very dif- ferent male genitalia, Nelasa howdeni differs from N. beckeri by having the head and pro- notum very dull and black, without a me- tallic sheen. The species is named for Henry Howden, one of the collectors and a noted scara- baeidologist. Nelasa duncansensis, NEw SPECIES Description: Holotype male, length 1.5 mm, width 1.2 mm. Form oval, slightly rounded. Color dark brown, head, prono- tum darker than elytron, with strong green metallic sheen, elytron with faint green me- tallic sheen; antenna, mouthparts, trochan- ter, tarsus brownish yellow. Head dull, sur- face alutaceous, finely punctured, punctures separated by less than to 3 times a diameter. Pronotum dull, surface alutaceous, punc- tures equal in size to head punctures, sep- arated by | to 2 times a diameter. Elytron smooth, polished, coarsely punctured, punctures separated by less than to twice a diameter. Genitalia with basal lobe % lon- ger than paramere, apex rounded, paramere slender; sipho short, unmodified (Figs. 20a— c). Allotype: Length 1.5 mm, width 1.2 mm. Similar to male except for genitalia. Variation: Length 1.4 to 1.6 mm, width 1.0 to 1.3 mm. Type material: Holotype; Jamaica, Try., Duncans, VIII.19.1966, Howden & Becker (CNC). Allotype; same data as holotype. Paratypes, total 14; 4, same data as holo- type; 8, same data as holotype except dates VII.13.1966, VIII.14.1966, VIII.15.1966, 303 VIII.25.1966; 1, Jamaica, Mandeville, Manchester Parish, VIII.16.1966, Howden & Becker collectors; 1, Jamaica, St. Eliz., Hermitage 7) Vil 1721966, 5E).C. Becker. (CNC) (USNM). Remarks: The brown elytra with faint green metallic sheen combined with the dark, strongly tinted head and pronotum distinguish N. duncansensis from other known Jamaican species. The species is named for the type locality. Nelasa schwarzi, NEw SPECIES Description: Holotype male, length 1.4 mm, width 1.1 mm. Form broadly oval, slightly rounded. Color black, dorsal surface with distinct purple sheen; antenna, mouth- parts, coxa, tibia, tarsus brownish yellow; venter, femur dark brown. Head dull, sur- face strongly alutaceous, punctures very fine, barely visible, separated by 2 to 3 times a diameter. Pronotum dull, surface strongly alutaceous, punctures slightly coarser than on head, separated by less than to twice a diameter. Elytron shiny, surface with trace of alutaceous sculpture, punctures coarse, separated by about a diameter. Genitalia with basal lobe straight in lateral view, very slightly longer than paramere (Figs. 20d, e); sipho lost. Type material: Holotype; Cuba, Caya- mas, 25.5, EA Schwarz Collector (USNM). Remarks: The holotype is the only spec- imen seen. Nelasa schwarzi resembles the Jamaican N. howdeni in external appear- ance but the male genitalia of each species are different. The species is named for E. A. Schwarz, collector of the holotype. Nelasa cubensis, NEw SPECIES Description: Holotype female, length 1.4 mm, width 1.1 mm. Form oval. Color black with strong, metallic green sheen through- out dorsal surface; antenna, mouthparts, trochanter, tarsus brownish yellow; venter, femur, tibia brown. Head with surface smooth, polished, finely punctured, punc- 304 tures separated by less than to 3 times a diameter. Pronotum with surface smooth, polished, punctures equal in size to head punctures, separated by less than to 3 times a diameter. Elytron shiny, surface with faint trace of alutaceous sculpture, punctures coarse, separated by less than to about a diameter. Type material: Holotype; Cuba, Bah. Honda, June 1-3, Wickham (USNM). Remarks: The holotype female is the only specimen examined. This species is thus far unique within the genus in having the head and pronotal surfaces as smooth and pol- ished as the elytra. The species is named for the country of origin. Nelasa haitiensis, NEw SPECIES Description: Holotype male, length 1.6 mm, width 1.3 mm. Form oval. Color black with faint coppery sheen on head, prono- tum; elytron with distinct metallic green sheen; antenna, mouthparts, trochanter, tarsus brownish yellow; tibia, femur dark brown. Head dull, surface strongly aluta- ceous, punctures very fine, barely visible, separated by less than to twice a diameter. Pronotum dull, surface strongly alutaceous, punctures equal in size to head punctures, separated by | to 3 times a diameter. Ely- tron smooth, surface with faint trace of alu- taceous sculpture, punctures coarse, sepa- rated by a diameter or less. Genitalia with short, robust projections in apical 4 of basal lobe (Figs. 20f-g); sipho lost. Type material: Haiti, Kenscoff, VIII-11- 35, Sta 23, Blackwelder (USNM). Remarks: The holotype is the only spec- imen examined. The species is named for the country of origin. Nelasa dominicensis, NEw SPECIES Description: Holotype male, length 1.5 mm, width 1.2 mm. Color dark brown with metallic greenish sheen on dorsal surface except lateral margin of elytron narrowly light brown; antenna, mouthparts yellow; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON venter, leg yellowish brown. Head shiny, surface finely alutaceous, punctures fine, separated by a diameter or less. Pronotum shiny, surface finely alutaceous, punctures equal in size to head punctures, separated by about a diameter. Elytron smooth, pol- ished, coarsely punctured, punctures sepa- rated by a diameter or less. Genitalia with basal lobe slender, strongly curved in lateral view, apex with apical projection; sipho short, unmodified (Figs. 20h-). Allotype: Length 1.5 mm, width 1.2 mm. Similar to holotype except for genitalia. Variation: Length 1.5 to 1.6 mm. Type material: Holotype; Dominican Re- public, La Matica, Boca-Chica, III-6-1955, A.M. Nadler (USNM). Allotype; same data as holotype (USNM). Paratypes, 3; same data as holotype. (USNM). Remarks: In addition to male genitalia, the relatively shiny head and pronotum dis- tinguish N. dominicensis from the other known Hispaniola species, N. haitiensis. Paranelasa, NEw GENUS Sticholotidini with form oval; without pubescence except on clypeal apex, lateral margin of head near eye; dorsal surface with slight metallic sheen; punctation on elytron not greatly coarser than on pronotum. Head broad; clypeus short, slightly, broadly emar- ginate apically, anterior angle abruptly rounded. Eye coarsely faceted, small, eyes separated by 5 times width of eye; gena ex- tended onto eye. Antenna 10-segmented; club 3-segmented, elongate (Fig. 6); inser- tion exposed. Terminal segment of maxil- lary palpus long, distinctly narrowed api- cally (Fig. 2). Epipleuron broad, descending externally, not foveate for reception of fem- oral apices. Prosternum with coxae narrow- ly separated by protuberant rectangular pro- cess, process carinate on each side, joined at apex (Fig. 10). Leg with femur robust, shallowly grooved for reception of tibia; tib- ia slender, shorter than femur; tarsus cryp- totetramerous, tarsal claw simple, lacking tooth. Abdomen with 6 visible sterna; Ist VOLUME 93, NUMBER 2 305 Fig. 19. a-i, genitalia. ac, male genitalia of Nelasa beckeri; d—f, male genitalia of Nelasa iricolor; g, h, male genitalia of Nelasa howdeni; i, female genitalia of Nelasa beckeri. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 306 Boe SNe Sa IS Awe. Sa E> SEE aa iy ——— ee aD ad = a-j, male genitalia. ac, Nelasa duncasensis; d, e, Nelasa schwarzi; f, g, Nelasa haitiensis; h-j, Nelasa Fig. 20. dominicensis. VOLUME 93, NUMBER 2 sternum with postcoxal line short, widely incomplete (Fig. 16); apex of male 5th ster- num briefly truncate, apex of female 6th sternum broadly rounded. Male genitalia symmetrical; basal lobe longer than para- mere. Female genitalia without spermathe- cal capsule or infundibulum. Membranous wing completely absent. Type species: Paranelasa jamaicensis, new species. Paranelasa is most similar to Nelasa, but Paranelasa differs by having an elongate an- tennal club, an elongate terminal maxillary palpal segment, no membranous wings, and no spermathecal capsule or infundibulum. Food and habitat preferences are unknown. The generic name is composed of the pre- fix Para- added to the genus name Nelasa and the gender is feminine. KEY TO SPECIES OF PARANELASA 1. Punctures on elytron distinctly visible; elytron with coppery or greenish metallic sheen; pro- sternal carinae angled toward each other api- Calvert Seti certreecaten eee jJamaicensis, 0. sp. — Punctures on elytron faint, distinctly visible only under high magnification; elytron with faint bluish metallic sheen; prosternal carinae Marallelaw nce wionce ne Ss ce eee: polita, n. sp. Paranelasa jamaicensis, NEw SPECIES Description: Holotype male, length 1.8 mm, width 1.4 mm. Color black, dorsal sur- face with faint metallic copper sheen; an- tenna yellow except club brown; mouth- parts, leg brown. Head smooth, polished, with faint trace of alutaceous sculpture, punctures fine, separated by less than to 3 times a diameter. Pronotum smooth, pol- ished, with faint trace of alutaceous sculp- ture, punctures slightly finer than on head, separated by | to 3 times a diameter. Ely- tron smooth, polished, with faint trace of alutaceous sculpture, punctures fine, indis- tinct, slightly coarser than pronotal punc- tures, separated by | to 3 times a diameter. Prosternal carinae angled toward each other apically. Genitalia with sipho gradually nar- 307 rowed before constriction at apical ’s, apex with small dorsal crest (Figs. 21a—c). Allotype: Length 1.7 mm, width 1.3 mm. Similar to holotype except for genitalia. Variation: Length 1.6 to 2.0 mm, width 3.10.7 mm: Type material: Holotype; Jamaica, Blue Mt. Peak, 7400’, VII.27—28.1966, Howden & Becker (CNC). Allotype; same data as holotype (CNC). Paratypes, total 99; 97, same data as holotype; 1, Jamaica, Port- land, N. side of Mossman’s Peak, 16- VIII.1980, 1400 m, A. Norrbom; 1, Ja- maica, St. Thomas, Portland Gap, 1600 m, 24-VIII.1980, A. Norrbom. (CNC) (CM) (USNM). Remarks: This species is named for the country of origin. Paranelasa polita, NEw SPECIES Description: Holotype male, length 1.8 mm, width 1.5 mm. Color black, dorsal sur- face with metallic sheen; antenna yellow ex- cept segments 5-8 brown; mouthparts, tar- sus yellow; leg, venter, dark brown to black. Head smooth, polished, finely punctured, punctures separated by a diameter or slight- ly more. Pronotum smooth, polished, punc- tures slightly finer than on head, separated by less than to twice a diameter. Elytron smooth, polished, punctures equal in size to head punctures, separated by 1 to 2 times a diameter. Prosternal carinae parallel, not angled toward each other apically. Genitalia with sipho broad before constriction at api- cal %, abruptly narrowed at constriction, apex without crest (Figs. 21d-f). Type material: Holotype; Jamaica, Hard- war Gap, Portland Par., 4000’, 03 Aug 1956, B&B Valentine, Hardwood cloud forest, beating (USNM). Paratypes, total 3; 1, Ja- maica, N side of Mossman’s Peak, 20- VIII.80, 1450 m, A. Norrbom; 1, Jamaica, Portland, N side of Mossman’s Peak, 26- VIII.1980, 1500 m, A. Norrbom; 1, Ja- maica, St. Thomas, Portland Gap, 1600 m, VIII.1980, A. Norrbom. (CM) (USNM). Remarks: The two known species of 308 Paranelasa are quite similar in external ap- pearance but the sipho of the male genitalia differs. The specific name refers to the smooth, polished appearance of the dorsal surface. Semiviride, NEw GENUS Sticholotidini with form oval, not strong- ly convex; without pubescence except on clypeal apex, lateral margin of head near eye; dorsal surface heavily sculptured, with metallic sheen; punctation on elytron not coarser than on head, pronotum. Head broad, clypeus short, apically truncate, an- terior angle abruptly rounded. Eye coarsely faceted, small, eyes separated by 4 times width of eye; gena extended onto eye. An- tenna 10-segmented; club 3-segmented, very broad (Fig. 8); insertion exposed. Terminal segment of maxillary palpus elongate, slen- der, distinctly narrowed apically (Fig. 3). Epipleuron broad, slightly descending ex- ternally, not foveate for reception of fem- oral apices. Prosternum with coxae narrow- ly separated by protuberant, triangular process, process heavily ridged on each side, carinae joined at apex (Fig. 11). Leg with femur robust, shallowly grooved for recep- tion of tibia; tibia slender, equal in length to femur; tarsus cryptotetramerous; tarsal claw simple, lacking tooth. Abdomen with 5 visible sterna; Ist sternum with postcoxal line incomplete, parallel to hind margin of sternum in apical '2 (Fig. 17); apex of male 5th sternum briefly truncate medially, apex of female Sth sternum broadly rounded. Male genitalia symmetrical. Female geni- talia with spermathecal capsule, without in- fundibulum (Fig. 21j). Fully developed membranous wing absent. Type species: Semiviride loisobrienae, new species. Semiviride is most similar to Neotina but Neotina is strongly convex, dorsally smooth, polished, the prosternal process lacks ridg- es, the antennal club is not as strongly ex- panded, and the terminal segment of the maxillary palpus is shorter, less strongly PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON narrowed apically. Food and habitat pref- erences are completely unknown. The generic name is a combination of the Latin semi, meaning half, and viride, a neu- ter Latin noun meaning green, and refers to the semigreen dorsal surface of the known species. KEY TO SPECIES OF SEMIVIRIDE 1. Pronotal surface strongly wrinkled, impunc- tate; elytron densely alutaceous, punctures ob- scured by surface sculpture ... /oisobrienae, n. sp. -— Pronotal surface feebly alutaceous, distinctly, coarsely punctured; elytron alutaceous but punctures not obscured by surface sculpture . . 3 if sella ett ake Bia ne ined os Bled enas portoricensis, n. sp. Semiviride loisobrienae, NEw SPECIES Description: Holotype male, length 1.8 mm, width 1.5 mm. Color black, dorsal sur- face with metallic green sheen; antenna yel- low except apical 4 segments pale, straw yel- low; mouthparts, coxa, trochanter, apex of femur, tibia, tarsus brownish yellow; ven- tral surface, basal % of femur dark brown. Head with surface alutaceous, slightly wrin- kled, feebly shiny, punctures fine, separated by about a diameter, difficult to see in sur- face sculpture. Pronotum with surface coarsely alutaceous, strongly wrinkled, dis- tinctly shiny, impunctate. Elytron densely alutaceous, not wrinkled, punctures larger than on head, separated by less than to 4 times a diameter, difficult to see because of surface sculpture. Genitalia with basal lobe slightly longer than paramere; paramere very slender; sipho elongate (Figs. 21g-1). Allotype: Length 1.7 mm, width 1.6 mm. Similar to holotype except genitalia as in Fig. 21). Variation: Length 1.7 to 1.8 mm, width 125)to*l.6°mim: Type material: Holotype; Puerto Rico, Carib. N.F. El Yunque Hwy. (191)K11H2, July 1979, G. B. Marshall (USNM). Allo- type; same data as for holotype except (191)K10H9, Lois O’Brien (USNM). Para- types, total 7; 1, same data as holotype; 1, Puert. Rico, Carib. Nat. For. base Mt. Brit- VOLUME 93, NUMBER 2 ton Tr., 17 March 1983, R. S. Miller colr.; 2, Puerto Rico, El Yunque, 16-17 July 1954, M. W. Sanderson; 1, Puerto Rico, Carib- bean Nat. Forest, El Yunque Trail, 610- 1050 m. 23 Sep 1987, M. A. Ivie, beating; 1, Puerto Rico, kCarib. N.F., El Yunque Hwy, (191)K11H4, July 29, 1979, C. W. O’Brien; 1, PR: Sierra Luquillo, Caribbean Nat. For. Rd 191, 2500’ (12 KmS. Palmer), XII-22-86, J. Doyen & J. Santiago. (MI) (UCB) (USNM). Additional specimen: 1, Puerto Rico, El Yunque, USFS aviary, July-Aug 1985, E. LaRue, at light. Remarks: The species is named for Lois O’Brien, collector of the allotype. The single specimen listed above collected at the USFS aviary is a female that possibly represents an undescribed species. It is slightly larger than S. /oisobrienae and the dorsal surface has a coppery sheen; however, because it is a unique female it is tentatively considered a variant of S. loisobrienae. Semiviride portoricensis, NEW SPECIES Description: Holotype male, length 1.75 mm, width 1.5 mm. Color black; dorsal sur- face with blue green sheen; antenna yellow except apical 4 segments slightly paler than remaining segments; mouthparts, coxa, tro- chanter, tibia, tarsus brownish yellow; ven- tral surface, femur dark brown. Head with surface alutaceous, not wrinkled, feebly shiny, punctures fine, separated by about a diameter. Pronotum with surface feebly alu- taceous, shiny, coarsely punctured, punc- tures separated by | to 3 times a diameter. Elytron distinctly alutaceous, punctures as coarse as pronotal punctures, not obscured by surface sculpture, separated by less than to 3 times a diameter. Genitalia with phal- lobase as illustrated for S. /oisobrienae ex- cept trabes slender with unmodified apex; sipho with basal capsule reduced, feebly de- veloped (Figs. 22a—g). Type material: Holotype; Puerto Rico, Carib. N.F., El Yunque Hwy., (191)K12H7, July 29, 1979, L. B. O’Brien (USNM). 309 Remarks: In addition to the key charac- ters, S. portoricensis is distinguished from S. loisobrienae by the feebly developed si- phonal capsule and slender, unmodified trabes; however, the phallobases are virtu- ally identical. Neaptera, NEw GENUS Sticholotidini with form rounded, strong- ly convex; without pubescence except for row of long setae on clypeal apex; dorsal surface with metallic sheen, at least on el- ytron; elytron with coarse, widely spaced punctures much larger than head, pronotal punctures. Head broad; clypeus short, trun- cate apically; anterior angle abruptly round- ed. Eye very coarsely faceted, small, eyes separated by 4 times width of eye; gena ex- tended onto eye. Antenna 10-segmented; club 3-segmented, broad (Fig. 8); insertion exposed. Terminal segment of maxillary palpus elongate, slender, slightly tapered apically (Fig. 4). Epipleuron narrow, flat, not foveate for reception of femoral apices. Prosternum with coxae narrowly separated by protuberant, rectangular process, process carinate on each side, carinae not joined at apex. Leg with femur robust, shallowly grooved for reception of tibia; tibia slender, shorter than femur; tarsus cryptotetramer- ous, tarsal claw simple, lacking tooth. Ab- domen with 6 visible sterna (6th sternum barely visible); 1st sternum with complete, slightly angulate, postcoxal line (Fig. 18); apex of male 5th sternum broadly rounded; apex of female 5th sternum strongly round- ed. Male genitalia symmetrical. Female gen- italia with unmodified spermathecal cap- sule; without infundibulum. Membranous wing lacking. Type species: Neaptera purpurea, new species. Neaptera is similar to Nexophallus but the latter genus differs by having the head and pronotum finely pubescent, the terminal segment of the maxillary palpus strongly ta- pered apically, the prosternum produced anteriorly, the epipleuron broad, slightly de- 310 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 21. a-j, genitalia. a-c, male genitalia of Paranelasa jamaicensis, d—f, male genitalia of Paranelasa polita; g-i, male genitalia of Semiviride loisobrienae; j, female genitalia of Semiviride loisobrienae. VOLUME 93, NUMBER 2 311 Fig. 22. a-g, genitalia. ac, male genitalia of Semiviride portoricensis; d-f, male genitalia of Neaptera purpurea; g, female genitalia of Neaptera purpurea. 312 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 23. a-h, male genitalia. a-c, Neaptera viridissima; d-f, Neaptera korschefskyi; g, h, Neaptera viola. scending externally, no membranous wings, and the postcoxal line not angulate. Four species of this flightless genus are thus far known from Guadeloupe, Puerto Rico, and the Virgin Islands. Food preferences are un- known but label data for N. viridissima in- dicate that the habitat is quite different from that of most coccinellids. The data are “‘at VOLUME 93, NUMBER 2 base of Kapok,” “litter in clump of bam- boo,” “‘base of large trees,” “litter along walls” and “‘leaf axils of coconut palm.” This information and the wingless condition of all known species explain why members of Neaptera are infrequently collected and probably indicate the presence of still more species on various West Indian islands. The generic name refers to the wingless condition and the gender is feminine. KEY TO SPECIES OF NEAPTERA 1. Elytron metallic green; Puerto Rico a EET OPEL oe Cp ee eae viridissima, N. sp. - Elytron metallic blue, purple, or violet; not knownitrom Puerto Rico: {sa5.6. 4-60. 2 2(1). Species occurring on Guadeloupe or Mont- SEIT a Caria ete ae ete a gate et eee 3 - Species known only from the Virgin Islands PARENT SOS oes eek Ce purpurea, N. sp. Elytron mostly metallic blue; punctures dense, coarse punctures separated by a di- AMELEIZOTMIESS ere tetehs ee ean ae korschefskyi (Duverger), n. comb. - Elytron metallic violet or purple; coarse punctures separated by a diameter or more » fe hoe kOe eee Eee ear ae ae viola, n. sp. Neaptera purpurea, NEW SPECIES Description: Holotype male, length 1.25 mm, width 1.0 mm. Color light yellowish brown; elytron mostly dark metallic purple with greenish iridescence; lateral, basal bor- ders of elytron, entire head, pronotum, clear, dark reddish brown; antenna, mouthparts, tibia, tarsus pale yellow. Head smooth, pol- ished, punctures distinct, separated by less than to twice a diameter. Pronotum smooth, polished, punctures finer than on head, sep- arated by less than to twice a diameter. El- ytron smooth, polished, surface with faint trace of alutaceous sculpture, with sparse, intermixed fine, coarse punctures, coarse punctures separated by | to 3 times a di- ameter. Genitalia as in Figs. 22d-f. Allotype: Length 1.3 mm, width 1.1 mm. Similar to male except genitalia as in Fig. 22g: Variation: Length 1.2 to 1.3 mm, width 1.0 to 1.1 mm. Some specimens have no 313 trace of greenish iridescence on the elytron, others have distinct iridescence. Type material: Holotype; Virgin Islands, St. John, Lameshur Bay, VIERS, 23 FEB1984, base of Kapok, W.B. Muchmore (USNM). Allotype; Virgin Islands, St. John, Little Lameshur Bay, 24JAN1986, litter in clump of bamboo, W.B. Muchmore colr. (USNM). Paratypes, total 15; 2, same data as holotype; 1, same data as holotype except date O4MAY 1984; 4, same data as allotype; 2, Virgin Islands, Trunk Bay, 08 JUN 1980, leafaxils of coconut palms, colr W.B. Much- more; |, Virgin Islands, St. John, Annaberg ruins, 13 JUN 1980, litter along wall, W.B. Muchmore; 2, Virgin Islands, St. John, Johnny Horn Trail summit over Emmaus, 13 MAY 1984, base of lg. trees, WBMuch- more; 1, Virgin Islands, St. John, Estate Carolina, King Hill, south O8SMAY1984, ground litter, WBMuchmore; 1, Virgin Is- lands, St. John, King Hill, 21 MAY 1982, W.B. Muchmore; |, Virgin Islands, St. John, Est. Maho Bay, Windberg Ruins, 20 & 31 MAY 1979, litter at base of walls, colr. W.B. Muchmore; 1, Virgin Islands, St. Thomas, Jan 20, 1963, Paul J. Spangler, Pond, | mi. E. Charlotte Amalie. (MI) (USNM). Remarks: This is the only species of Neaptera represented by more than a few specimens. Thus far known only from the Virgin Islands, it is most similar to the Puer- to Rican N. viridissima in form of the male genitalia. The elytral punctation and color are different however, and they are here treated as two valid species. The specific name is Latin and refers to the metallic purple dorsal color. Neaptera viridissima, NEw SPECIES Description: Holotype male, length 1.3 mm, width 1.0 mm. Color light yellowish brown; elytron metallic green; head, pro- notum clear, dark reddish brown; antenna, mouthparts, tibia, tarsus pale yellow. Head smooth, polished, punctures distinct, sep- arated by less than to a diameter. Pronotum smooth, polished, punctures equal in size 314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to head punctures, separated by less than to twice a diameter. Elytron smooth, polished, with fine punctures sparse, very small, bare- ly visible, coarse punctures sparse, separat- ed by 2 to 4 times a diameter. Genitalia as in Figs. 23a-c. Allotype: Length 1.35 mm, width 1.1 mm. Similar to male except for genitalia. Variation: Length 1.3 to 1.4 mm. Type material: Holotype; Puerto Rico, nr Fajardo, rtl94 km45.7, VIII-20-61, Flint&Spangler (USNM). Allotype; Puerto Rico, Mayaguez, 10.9.‘37, P.R. 2035, H.K. Plank (USNM). Paratypes, total 3; 2, same data as holotype; 1, same data as allotype. Remarks: This species is most similar to N. purpurea but can be recognized by the distinctly metallic green elytra. See remarks under N. purpurea. The specific name is Latin and refers to the metallic green dorsal coloration. Neaptera korschefskyi (Duverger), New ComMBiNATION Nexophallus korschefskyi Duverger, 1986: 223: Description: Length 1.6 mm, width 1.2 mm. Color dark brown; dorsal surface mostly dark metallic blue, with some cop- per, violet iridescence; antenna, mouth- parts, tibia, tarsus brownish yellow. Head smooth, polished, finely punctured, punc- tures separated by less than to twice a di- ameter. Pronotum smooth, polished, punc- tures equal in size to head punctures, separated by less than to twice a diameter. Elytron shiny, densely punctured, coarse punctures separated by a diameter or less, with numerous intermixed fine punctures. Male genitalia as in Figs. 23d-f; sipho bro- ken. Type locality: Guadeloupe, Vitrac, Trois rivieres. Type depository: Museum National d’Histoire Naturelle, Paris. Specimens examined: The female holo- type from the Paris Museum and 4 addi- tional specimens, all of which are labeled ‘““Guadeloupe”’ with no further data. Remarks: Neaptera korschefskyi has the most densely punctured elytra thus far known within the genus. This character, the mostly metallic blue dorsal color, and form of the male genitalia distinguish N. kor- schefskyi from N. viola which also occurs on Guadeloupe. Duverger (1986) placed N. korschefskyi in the genus Nexophallus which is understandable because the two genera are superficially very similar and he did not have a specimen of Nexophallus with which to make a direct comparison. In addition to the holotype, Duverger (1986) described an allotype and 18 paratypes from various lo- calities on Guadeloupe. The holotype has been examined courtesy of N. Berti of the Paris Museum. Neaptera viola, NEw SPECIES Description: Holotype male, length 1.4 mm, width 1.0 mm. Color brown; elytron dark metallic purple with violet iridescence; pronotum dark purple with anterior, lateral margins dark reddish brown; head dark red- dish brown basally, becoming paler reddish brown toward clypeal apex; antenna, mouthparts, tibia, tarsus yellow. Head smooth, polished; punctures fine, separated by less than to twice a diameter. Pronotum smooth, polished, with basomedian area of punctures coarser than on head; remainder of pronotal surface with very fine, barely visible punctures. Elytron smooth, pol- ished, with intermixed fine, coarse punc- tures, coarse punctures separated by | to 3 times a diameter, fine punctures indistinct, widely separated. Genitalia as in Figs. 23g, h; sipho lost. Allotype: Similar to holotype except for female genitalia. Variation: The elytral color ranges from that described above to having a mixture of purple, green, and violet reflections. Type material: Holotype; Insel Guade- loupe, Coll. Mus. Vindob., Pentilia egena Muls., Scymnillodes (NHMA). Allotype; VOLUME 93, NUMBER 2 Insel Guadeloupe (NHMA). Paratype, 1; same data as allotype (USNM). Other specimen: 1, Montserrat, BWI, VII- 24-36, Blackwelder (USNM). Remarks: This species is most similar to N. korschefskyi, see remarks under that spe- cies. The single female specimen from Montserrat is apparently conspecific with the type specimens of N. viola from Gua- deloupe but is not designated a paratype. The specific name is Latin and refers to the predominantly metallic violet tint of the elytron. Parinesa, NEw GENUS Sticholotidini with form rounded, very convex; without pubescence except head with fine, sparse, short, decumbent hairs, row of long setae on clypeal apex; dorsal surface with metallic sheen, at least on el- ytron; elytron with punctures not larger than head, pronotal punctures. Head broad; clypeus short, truncate apically; anterior an- gle broadly rounded. Eye very coarsely fac- eted, small; eyes separated by 4 times width of eye; gena broadly extended onto eye forming shelf dividing eye nearly in half (Fig. 13). Antenna 10-segmented; club 3-seg- mented; insertion concealed. Terminal seg- ment of maxillary palpus elongate, slender, distinctly tapered apically. Epipleuron nar- row, abruptly descending externally, with shallow depression for reception of femoral apices. Prosternum with coxae very narrow- ly separated by prosternal process, apex of prosternum scoop shaped, partially con- cealing mouthparts (Fig. 12). Leg with fe- mur robust, grooved for reception of tibia; tibia modified, anterior, middle tibiae broadly expanded with sharp angulation on outer margin (Fig. 14), posterior tibia not expanded, with slight external angulation; tarsus cryptotetramerous; tarsal claw sim- ple, lacking tooth. Abdomen with 5 visible sterna; Ist sternum with incomplete post- coxal line joining hind margin of sternum. Genitalia not examined. Presence or ab- 315 sence of membranous wings not deter- mined. Type species: Parinesa whiteheadi, new species. Parinesa is similar to Glomerella in hav- ing the eye broadly divided, the prosternum apically expanded, the epipleuron foveate, the anterior pair of tibiae modified, and ab- domen with 5 visible sterna. Parinesa dif- fers from Glomerella in having the body less convex, the pronotum without pubescence, the clypeus anteriorly truncate with broadly rounded angles, and the anterior and middle tibiae externally dentate. Only a single spec- imen has thus far been seen and it was not dissected, therefore the genitalia, antenna, and mouthparts are not illustrated and pres- ence or absence of membranous wings not determined. Habitat and food preferences are unknown. The generic name is an arbitrary com- bination of letters and the gender is femi- nine. Parinesa whiteheadi, NEw SPECIES Description: Holotype female, length 1.25 mm, width 0.90 mm. Elytron black with faint metallic blue sheen; head, pronotum dark brown except clypeus light reddish brown; antenna, mouthparts, leg, 5th ab- dominal sternum yellow, remainder of ven- ter dark reddish brown. Head finely aluta- ceous, somewhat shiny, punctures fine, indistinct, separated by about a diameter. Pronotum finely alutaceous, shiny, punc- tures barely visible. Elytron smooth, pol- ished, punctures extremely fine, shallow, barely visible. Genitalia not examined. Type material: Holotype; Dominican Re- public, 5.22.36, S. Francisco - 8073, E.C. Decker (USNM). Remarks: Named for my old friend and companion, Donald R. Whitehead. ACKNOWLEDGMENTS I thank N. Berti, Museum National d’Histoire Naturelle, for allowing me to ex- amine the holotype of Nexophallus kor- 316 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON schefskyi Duverger. I also thank R. D. Pope, British Museum, London, N. Vandenberg, Berkeley, California, and F. C. Thompson, Systematic Entomology Laboratory, Wash- ington, D.C., for reviewing the manuscript. Kelly Marsh prepared the illustrations. LITERATURE CITED Duverger, C. 1986. Revision des Coccinellidae de Guadeloupe (Antilles francaises) I—Subfamilies des Sukunahikonini et Sticholotidinae (Coleoptera Coccinellidae). L’Entomologiste 42: 219-225. Gordon, R. D. 1977. Classification and phylogeny of the New World Sticholotidinae. The Coleopterists Bulletin 31: 185-228. Gordon, R. D., J. Pakaluk, and S. A. Slipinski. 1989. Carinodulini, a new tribe of Sticholotidinae based upon a new genus and species from Mexico (Co- leoptera: Coccinellidae). The Coleopterists Bulle- tin 43: 359-364. PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 317-321 PHYLOGENETIC PLACEMENT OF PHAENOCEPHALUS WOLLASTON (COLEOPTERA: PHAENOCEPHALIDAE & PHALACRIDAE) JAMES PAKALUK Snow Entomological Museum, University of Kansas, Lawrence, Kansas 66045. Abstract.—The taxonomic history of the family Phaenocephalidae is reviewed. This family currently consists of two species, Phaenocephalus castaneus Wollaston and P. coomani Paulian, described from Japan and Viet Nam respectively. The genus is rede- scribed and selected structures are illustrated. The relationship of this enigmatic family to other beetles has been disputed for years. The uncertainty of its phylogenetic position has been due, in part, to the fact that type material has been unavailable for study for forty years or more. After examining types in collections in London and Paris it seems that these two species, previously attributed to a separate family, should be placed in the Phalacridae. Key Words: The Phaenocephalidae is one of the smallest families of Coleoptera, with only two species described, as well as one of the most enigmatic. Ever since this family was described nearly 100 years ago there have been few published reports for these beetles. While studying types and other material of Cucujoidea recently in London and Paris, I was able to locate specimens of Phaenoce- phalidae that have likely been unavailable for study for forty years or more. Phaeno- cephalus was not listed in the generic list of Coleoptera at The Natural History Muse- um, London, and the material at the Mu- séum National d’Histoire Naturelle, Paris was among Paulian’s dissections of Cory- lophidae which were until recently consid- ered lost or destroyed. It seems appropriate to now review the taxonomic history of these beetles, redescribe the genus, illustrate im- portant structural features, and discuss the phylogenetic position of this family. Wollaston (1873) described Phaenoceph- alus castaneus as a corylophid based upon a single specimen from Japan. He thought Coleoptera, Cucuioidea, Phaenocephalidae, Phalacridae it was most similar to Corylophus Stephens but grouped Phaenocephalus with Sacium LeConte and Microstagetus Wollaston based upon their 11-segmented antennae. Mat- thews (1899) established a new family, Phaenocephalidae, for Wollaston’s species and provided a more detailed description and some illustrations. Both authors em- phasized the large, deflexed head and an- teriorly emarginate edge of the pronotum in Phaenocephalus. Matthews, however, thought these features, as well as others, were sufficient to exclude this species from Cor- ylophidae. He speculated that Phaenoce- phalidae was a link between Corylophidae and Silphidae. Paulian (1950) described another species of Phaenocephalus, P. coomani, from Viet Nam and illustrated a few structures. In a footnote he indicated that he had seen ad- ditional material of an undetermined spe- cies of Phaenocephalus from Sumatra; I was unable to locate this material. I did find, however, a specimen labelled as Phaeno- cephalus sp. from Viet Nam, but this is re- 318 ally a species of Orthoperus Stephens (Cor- ylophidae). There are few subsequent references to this family, and these were presumably made without referring to type material. Crowson (1955) treated Phaenocephalidae as Cole- optera incertae sedis, Lawrence (1982) con- sidered this family synonymous with Phal- acridae, and Sasaji (1985) included a photograph of Phaenocephalus castaneus and treated it as a distinct family of Cucu- joidea. The quality of preserved material for this family is extremely poor. Wollaston’s type of Phaenocephalus castaneus is virtually lost; the only parts that I could find were the labium and a maxilla mounted in balsam on an acetate card. These structures agree with Matthews’s (1899) illustrations. All of Paulian’s material, including the type of P. coomani, were poorly mounted on slides and still in glycerin jelly. These specimens have been removed from their slides and transferred to glycerin-filled microvials. There are few features that can be compared to Wollaston’s or Matthews’s descriptions of P. castaneus. I have examined the following material of Phaenocephalus: P. castaneus.—The labium and a maxilla of the holotype. The number and date **759/ Aug 5 1885,” written by Matthews, is at- tached to the pin. Two slide-mounted spec- imens from Japan were used for Paulian’s (1950) monograph. One specimen is rep- resented by the head only. Paulian (in Iitt., 1990) never examined Wollaston’s type, so these may not be conspecific. The specimen that Paulian listed from Formosa in the Grouvelle collection was not found. I have an additional specimen from Japan identi- fied by H. Sasaji. Most of the illustrations included in this paper were made from this specimen and compared to previously pub- lished descriptions and illustrations. This specimen is in my collection. P. coomani.—The slide-mounted type described by Paulian was the only material PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON available. This type consists of the head and prothorax; the rest of the specimen appears to be lost. This is likely a syntype since Pau- lian lists the size of this species as 1.15—1.20 mm long, suggesting more than one speci- men was used for his description. I am not designating a lectotype with the hope that the remaining type material for this species will eventually be located. REDESCRIPTION OF PHAENOCEPHALUS WOLLASTON The following description is based upon the material listed above. Certain features, such as total body length, were taken from descriptions. Most details are described from the specimen of the putative P. castaneus that is in my collection. Every effort was made to compare these features to previ- ously published descriptions and illustra- tions, as well as the preserved material listed above. It is possible that this description is inaccurate since the type of the type species of this genus is almost entirely lost. In gen- eral, this agrees with Matthews’s (1899) de- scription, except that the mesocoxal cavities are clearly closed and the ventrites are sub- equal in length. Description.—Length 1.15-1.70 mm. Body ovate, about 1.3 x longer than wide, light brown, glabrous, shiny; dorsum con- vex, venter flat. Head (Fig. 3) transverse, widest at eyes. Eye on antero-lateral margin of cranium. Fronto-clypeal suture absent, clypeus strongly declivious anteriorly. Antenna 1 1- segmented with 3-segmented club, apical club segment elongate, subequal in length to basal two club segments combined; an- tennal groove indestinct. Labrum (Fig. 4) transverse, sclerotized, on different plane from clypeus; tormae (Fig. 4) elongate, slightly longer than labrum, fused apically with labial rods, with mesal arms directed anteriorly; epipharyngeal rods (Fig. 4) sub- equal in length to mesal arms of tormae, fringed with setae mesally. Mandible (Fig. 5) bifid apically; mola well-developed; pros- VOLUME 93, NUMBER 2 theca broad, fringed with hairs, with a small basal tuft of setae. Maxilla (Fig. 6) with 4-segmented palp, segment 4 longest; galea broad, distigalea with dorsal membranous area basally, with broad brush of elongate setae apically; lacinia narrow, bifid apically. Labium (Fig. 8) with 3-segmented palp, seg- ments 2 and 3 subequal in length; mentum transverse, anterolateral angles produced; prementum mostly membranous, basal sclerite small, ligula large, divergent apically with distinct lobes, labial rods elongate, fused apically with tormae. Gular sutures indistinct. Tentorium reduced, represented by vestigial anterior arms only. Pronotum (Fig. 1) transverse, about 0.4 x longer than wide, anterior edge with deep U-shaped emargination, posterior edge with large medial lobe, lateral edges convergent anteriorly, with smooth margins. Proster- num reduced anteriorly; intercoxal process narrow, elevated, deflexed apically. Procoxa globose, with long internal extension, its cavity internally closed, externally widely open. Mesosternum between coxae sub- equal in width to mesocoxal cavity, without femoral lines, junction of meso- and meta- sternum straight-line type, without internal knobs. Mesocoxa globose, trochantin con- cealed, its cavity laterally closed. Metaster- num twice as wide as long, without femoral lines, with medial line extending to middle, with posterior edge deeply notched medi- ally. Metacoxa large, transverse, almost contiguous. Metendosternite with furcal arms divergent at about 90 degree angle, anterior tendons short, curved laterally at apex. Leg with trochanter small, subtrian- gular; femur widest at middle, progressively longer posteriorly, fore and midfemur slightly longer than tibia, hind femur sub- equal in length to hind tibia; tibia elongate, subcylindrical, with numerous stout spines, especially along posterior edge, tibial spur formula 1-2-2, fore spur short, stout, barely longer than tibial spines, midspurs distinct, unequal in length, hindspurs distinct, sub- equal in length; tarsi (Fig. 7) 4-4-4, all tarsi 319 subequal in length, segments 1-3 distinctly lobed, 4 elongate, subequal in length to 1- 3 combined; claws with large, broad tooth at base, empodium absent. Scutellum small, triangular. Elytra about 3.2 longer than pronotum, punctation minute, distinctly se- riate, with 9 interneurs, intervals smooth except for extremely fine, seriate punctures; epipleuron large, strongly angled, complete. Wing (Fig. 2) with venation reduced, cells and a subcubital fleck absent, with 2 anal veins, jugal lobe present. Abdomen with 7 spiracles; 5 ventrites subequal in length, first with large, narrow intercoxal process; femoral lines absent; all ventrites free; 2—5 with small, anterolateral internal apodemes. Male genitalia with teg- men (Fig. 9) elongate, with apical subtrian- gular sclerite deeply emarginate medially, median lobe (Fig. 10) weakly sclerotized. I agree with Lawrence’s (1982) decision to synonymize Phaenocephalidae with Phalacridae, although he did this based upon published information only (J. F. Lawrence, pers. comm.). In general habitus, Phaeno- cephalus is certainly a typical phalacrid. Its lack of a fronto-clypeal suture, maxilla with distinct lacinia and galea, broadly open pro- coxal cavities, laterally closed mesocoxal cavities, 4-4-4 tarsi distinctly lobed with dentate claws, and subequal ventrites are sufficient to exclude Phaenocephalus from all other families of Cucujoidea except Phal- acridae. With the shape of the tegmen and the almost completely reduced tentorium it seems that this genus can easily be included in this family. Moreover, I have observed elongate tormae fused apically with elongate labial rods only in other phalacrids. This condition has not, to my knowledge, been reported previously for cucujoids. It may be related to the reduced tentorium or it may be a useful phylogenetic character at another level. The only other phalacrid with similar tar- si and tarsal claws is the Australian Phal- acrinus Blackburn. Species that I examined, however, are larger that Phaenocephalus, the 320 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-10. Phaenocephalus castaneus. 1, Pronotum. 2, Wing. 3, Cranium, ventral. 4, Labrum, left side ventral, right side dorsal. 5, Mandible. 6, Maxilla. 7, Tarsus. 8, Labium. 9, Tegmen. 10, Median lobe. VOLUME 93, NUMBER 2 elytral punctures are more distinct, without punctation on the intervals, and the lateral margins of the elytra are distinctly and sometimes broadly explanate. It is likely that these two genera are sister groups or that they should eventually by synonymyzed. Although the phylogenetic importance of some of these features is still uncertain, it is reasonable to include Phaenocephalus in Phalacridae until a comprehensive phylo- genetic arrangement of this family is pro- posed. Even with such a system, the precise position of this genus will be tentative until the remainder of the types are discovered. Then we may more confidently discuss the placement of Phaenocephalus by utilizing specimens that have been compared with this material. ACKNOWLEDGMENTS I am deeply indebted to M. Kerley, The Natural History Museum, London, En- gland, and N. Berti, Muséum National d’Histoire Naturelle, Paris, France, for pro- viding material for this study. Both indi- viduals spent long hours searching for these specimens in their respective institutions. I am grateful to H. Sasaji for generously pro- viding me with a specimen of Phaenoceph- alus castaneus from his personal collec- tion and to J. F. Lawrence for a gift of Phalacrinus. 1 thank S. A. Slipinski for a spirited discussion of Phaenocephalus while visiting his laboratory and for his comments on this manuscript. R. A. Crowson, J. F. Lawrence, and W. E. Steiner also reviewed 321 this piece and contributed useful comments and suggestions. My visits to London and Paris were fund- ed by an Ernst Mayr Grant from the Mu- seum of Comparative Zoology, Harvard University, Cambridge, USA. These visits would have been impossible without an ad- ditional grant that transported me across the Atlantic from the Office of International Re- lations, Smithsonian Institution, Washing- ton, U.S.A. to study the phylogeny of the cerylonid series of Cucujoidea with S. A. Slipinski in Warsaw, Poland. This paper is dedicated to the late Don Whitehead in recognition of his numerous contributions to systematic entomology. LITERATURE CITED Crowson, R. A. 1955. The Natural Classification of the Families of Coleoptera. Lloyd, London. 187 pp. Lawrence, J. F. 1982. Coleoptera, pp. 482-553. In Parker, S. P., ed., Synopsis and Classification of Living Organisms. Volume 2. McGraw-Hill, New York. Matthews, A. 1899. A Monograph of the Coleopter- ous Families Corylophidae and Sphaeriidae. Jan- son & Son, London. 220 pp. Paulian, R. 1950. Les Corylophidae d’Afrique. Mé- moires de l’Institute Francais d’Afrique Noire. Number 12. 126 pp. Sasaji, H. 1985. Phaenocephalidae, p. 230, pl. 37, fig. 21. In Kurosawa, Y., S. Hisamatsu & H. Sasaji, eds., The Coleoptera of Japan in Color. Volume III. Hoikusha Publishing Co., Osaka. 500 pp. [in Japanese] Wollaston, T. V. 1873. On new Coleoptera from Ja- pan (part). Entomologists’ Monthly Magazine 10: 167-168 (part). PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 322-332 TAXONOMIC NOTES, NEW RECORDS, AND A KEY TO THE ADULTS OF NORTH AMERICAN BYRRHIDAE (COLEOPTERA) PAUL J. JOHNSON Department of Entomology, University of Wisconsin, Madison, Wisconsin 53706. Abstract. —Synonyms, lectotype designations, holotype recognitions and taxonomic notes are provided for North American Byrrhidae not treated elsewhere. Twenty-one names are reduced to junior synonym status, three junior synonyms are recombined, and one name is moved from subspecies to species status. Lectotypes are designated for 11 species described by T. L. Casey, G. H. Horn, J. L. LeConte, and W. F. Erichson. Holotype data is given for all species not previously reviewed. Byrrhus pilula L. is newly recorded from Canada, and Sierraclava cooperi Johnson is newly recorded from Mexico. A key is pre- sented for the identification of the subfamilies, genera, and species of Byrrhidae known to occur in North America. Key Words: The taxonomy of the North American Byrrhidae has not been fully treated since Casey (1912) monographed the family. Ca- sey described 46 species and subspecies, of which I consider 3 to be valid. In contrast, he described 6 genera and all are considered valid. Since Casey’s monograph, Sierracla- va (Johnson 1982) has been the only genus added, 5 new species have been described (Johnson 1985, 1986, 1991), 2 have been species recognized and reported as immi- grants from Europe (Johnson 1990), and one additional species is herein newly reported as a member of the North American byrrhid fauna. During the course of continuing taxo- nomic and ecological studies on the Byr- rhidae, a number of synonymies, lectotype designations, and name corrections were found to be necessary, and which could not be incorporated into revisionary studies in a timely manner. In addition, Byrrhus pilula L. is now recorded from North America for the first time, Sierraclava cooperi Johnson is newly reported from Mexico, and a key Coleoptera, Byrrhidae, North America, taxonomy to the known and recognized taxa in North America is provided. These data are pro- vided here due to delays in preparing an adequate monograph on the North Amer- ican byrrhids, and the need to complete no- menclatorial clarifications for final prepa- ration of the Byrrhidae fascicle for the United States Department of Agriculture Coleoptera Catalog (R. D. Gordon, editor), as well as clarifying names for use by other workers. Generally, only new synonymical data is presented here unless inclusion of previous synonymies is felt pertinent for clarity. Fur- ther, only the more important taxonomic and faunistic references are cited under each synonymy. Synonymical data and com- ments in Simplocaria are presented else- where (Johnson, submitted). Types mentioned are reposited in the fol- lowing institutions: Museum of Compara- tive Zoology, Cambridge (MCZ); U.S. Na- tional Museum of Natural History, Washington, D.C. (USNM); Zoologisches Museum Humboldt-Universitat, Berlin, VOLUME 93, NUMBER 2 DDR (ZMHU); The Natural History Mu- seum, London (BMNH); Canadian Nation- al Collection, Ottawa (CNC); and the Car- negie Museum of Natural History, Pittsburgh (CMNH). All specimens person- ally examined have my own handwritten designation or determination labels at- tached. Types of all species have been ex- amined except where noted. Amphicyrta chrysomelina Erichson Amphicyrta chrysomelina Erichson 1843: 40, of Casey 1912: 64, Hatch 1961: 301 (pars). Holotype, female: “9438; TYPUS; chrysomelina, Koching Mont., Oregon, Willcox; Zool. Mus. Berlin” (ZMHU). Amphicyrta chrysomelina oblonga Casey 1912: 64 New Synonym. Holotype, fe- male: “Cal; Casey bequest 1925; Type USNM 48378; oblonga Csy” (USNM). Amphicyrta chrysomelina parvuliceps Casey 1912: 65 New Synonym. Holotype, male: “Cal; Casey bequest 1925; Type USNM 48379; parvuliceps Csy” (USNM). Casey’s specimens of A. oblonga and A. parvuliceps are typical examples of A. chrys- omelina. Several hundred specimens of A. chrysomelina have been examined from throughout its range, and I can find no mor- phological or bionomical evidence for seg- regating local populations into several spe- cies or subspecies. Coloration differences noted by Casey (1912) are the same seen by specimens discolored by decomposing fatty tissues or specimens subjected to prolonged exposure to killing agents such as sodium cyanide. Amphicyrta dentipes Erichson Amphicyrta dentipes Erichson 1843: 40, Casey 1912: 65. Lectotype, sex not con- firmed: “9437; TYPUS; Amphicyrta den- tipes Esch., Californ. Esch.; Amphicyrta Esch. Er.; Zool. Mus. Berlin 1984” (ZMHU). Paralectotype: ‘‘California Eschsch. Nr. 9437; TYPUS; Zool. Mus. Berlin” (ZMHU). 323 Amphicyrta chrysomelina (pars) of Hatch 1961: 301. Amphicyrta elongata Casey 1912: 65 NEw Synonym. Holotype, female: “Cal; Casey bequest 1925; Type USNM 48380; elon- gata Csy” (USNM). Amphicyrta gentilis Casey 1912: 66 New SyNonyM. Lectotype here designated, sex not confirmed: ““Cal; Casey bequest 1925; Type USNM 48381; gentilis Csy” (USNM). Paralectotypes: 4, same data (USNM). Amphicyrta gentilis ventricosa Casey 1912: 66 New Synonym. Holotype, female: “Cal; Casey bequest 1925; Type USNM 48382; ventricosa Csy” (USNM). Amphicyrta nevadensis Casey 1912: 66 NEw SynonyM. Holotype, female: ““Nev.; Cas- ey bequest 1925; Type USNM 48383; nevadensis Csy” (USNM). Amphicyrta dentipes is a highly variable species in size and coloration, and these were the primary traits considered by Casey. None of the character states involving size or color are useful for unequivocal delimitation of species or subspecies. Variations examined do suggest some geographic regionalization, but there is extensive intergradation. Lioligus nitidus (Motschulsky) Simplocaria nitida Motschulsky 1845: 362, of Mannerheim 1852: 341, LeConte 1854: 116. Type not seen; probably lost (Kelen- ikova, 1n litt.) Lioligus nitidus (Motschulsky), of Casey 1912: 62, Hatch 1961: 300. Lioligus keeni Casey 1912: 61, of Hatch 1961: 301 New Synonym. Lectotype here designated, male: “Metlakatla, B. Col., Keen; Casey bequest 1925; Type USNM 48385: keeni Csy.’’ Paralectotypes: 3, same data (USNM). Lioligus striolatus Casey 1912: 61, of Hatch 1961: 300 New SynNonyoM. Lectotype here designated, female: ““Metlakatla, B. Col., Keen; Casey bequest 1925; Type USNM 48384; striolatus Csy.” Paralectotypes: 11, same data (USNM). 324 Lioligus aequabilis Casey 1912: 62, syn- onymy by Hatch 1961: 301. Holotype, female: “Br. C.; Casey bequest 1925; Type USNM 48386; aequabilis Csy.”” (USNM). Specimens attributable to L. nitidus have been examined from throughout the com- posite range of its synonyms and I can find no morphological or ecological justification for recognizing more than one species. Characters given by Casey in separating his ““species”’ are highly variable, and are minor variations in sculpture, body dimensions, and coloration. Although the type has apparently been lost, a neotype is not designated due to this species being readily identified on morpho- logical and distributional factors in con- junction with Motschulsky’s description and denoted locality of provenance. There is no available evidence suggesting nomenclatur- al confusion. Lioligus pallidus Casey Lioligus pallidus Casey 1912: 62. Lioligus nitidus, of Hatch 1961: 301. Lec- totype here designated, sex not con- firmed: “ID.; Casey bequest 1925; Type USNM 48387; pallidus Csy.” Paralecto- type: “Coeur d’Alene, Idaho” (USNM). The pallid coloration of the type is due to its teneral condition. Typical L. pallidus specimens are piceous with an olivaceous sheen dorsally, whereas L. nitidus is bril- liantly viridescent or aeneo-viridescent. Other traits for separating these two species are in the key below. Exomella pleuralis Casey Exoma pleuralis Casey 1908: 282, of Casey [912=337: Exomella pleuralis (Casey), of Casey 1914: 378, Hatch 1961: 299, Johnson & Russell 1978: 159, Johnson 1985: 155. Lectotype here designated, sex not confirmed: *“Metlakatla, B. Co., Keen; Casey bequest 1925; Type USNM 48357; pleuralis Csy.” PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (USNM). Paralectotypes: 10, same data (USNM). Two series of specimens with identical labelling, less the bequest and cataloging la- bels, are reposited in the CNC and the BMNH. These may be syntypical with the lectotype series, but are not so treated here due to a lack of confirming evidence. Morychus oblongus (LeConte) Pedilophorus oblongus LeConte 1857: 39, of Wickham 1903: 181. Morychus oblongus (LeConte), of Casey 1912: 8, Hatch 1961: 295. Lectotype here designated, male: ““medium blue disc in- dicating the Oregon Territory; Type 2293; Pedilophorus oblongus LEC., acuminatus # LEC.” (MCZ). Paralectotype: same data (MCZ). Morychus insulsus Casey 1912: 8, synony- my by Hatch 1961: 295. Holotype, sex not confirmed: ‘“‘Vernon, B.C., VI, Ven- ables; Casey bequest 1925; Type USNM 48324; insulsus Csy” (USNM). Pedilophorus subcupreous Fall 1907: 225. Morychus subcupreous (Fall), of Casey 1912: 9, synonymy by Hatch 1961: 295. Ho- lotype, male: ‘““Aberdeen, Wash.; subcu- preous TYPE; MCZ Type 24474; H.C. Fall Collection; Pedilophorus subcu- preous Fall’? (MCZ). Morychus insulsus represents a blue-green color variant of the typical M. oblongus which is rarely seen in living material, but is commonly induced by prolonged expo- sure to sodium cyanide. The type of M. sub- cupreous is only a slightly larger and slightly less brilliantly aeneous representative of M. oblongus. Morychus aeneolus (LeConte) Pedilophorus aeneolus LeConte 1863: 74, of Wickham 1903: 181. Morychus aeneolus (LeConte), of Casey 1912: 8. Holotype, female: ‘““Neb.; Pedi- lophorus aeneolus LEC.; Henry Ulke Coll. CMNH Acc. No. 1645” (CMNH). VOLUME 93, NUMBER 2 Pedilophorus subsetosus Fall 1907: 225 NEw SYNONYM. Morychus subsetosus (Fall), of Casey 1912: 14, Hatch 1961: 295. Holotype, female: ‘Kalispell, Mont., June 13, Wickham; subsetosus T Y PE; Type MCZ 24475; H.C. Fall Collection; Pedilophorus subsetosus Fall’? (MCZ). Pedilophorus lateralis Fall 1907: 225 New SYNONYM. Morychus lateralis (Fall), of Casey 1912: 9. Holotype, female: “N.M., Las Vegas, head of Daily Can.; 6.26.01; T.D.A.C.; lateralis TYPE; Type MCZ 24473; Pedilophorus lateralis Fall’? (MCZ). Pedilophorus hesperus Wickham 1903: 182 New SYNONYM. Morychus hesperus (Wickham), of Casey 1912: 9. Holotype, female: “Leadville, Col., H.F. Wickham, July 7-14 96, 10,000-11,000 ft.; Wickham Collection 1933; TYPE; Pedilophorus hesperus Wickham” (USNM). Morychus albertanus Brown 1932: 8 NEw SyNonyM. Holotype, sex not confirmed: ““Crow’s Nest Pass, Alberta, June 7, 1930, J.H. Pepper; No. 3246” (CNC). The various synonyms for this species re- flect slight variations of integument color and patterns of pubescence. Minor sculp- tural variations occur, but none correlate with discreet populations. Slight variations and distributional patterns of genitalic char- acters and pubescence support the synon- ymies, but also suggest the presence of clines over large geographic areas. Byrrhus cyclophorus Kirby Byrrhus cyclophorus Kirby 1837: 117, of El Moursy 1970: 329. Holotype, female: “Type; N.Amer.; 5828a; Byrrhus cy- clophorus Kirby, N. Amer., 5828, Rev. Wm. Kirby (BMNH). Byrrhus fasciatus, of El Moursy 1970: 330 NEw CoMBINATION. The application of the name B. fasciatus is here restricted in its application only to 325 North American Byrrhus determined as such by El Moursy (1970) and previous catalog- ers (Hamilton 1894a, b; Dalla Torre 1911; Leng 1920). Interestingly, neither of the mo- nographers of North American Byrrhidae, LeConte (1854) or Casey (1912), applied the name B. fasciatus to any native species. El- Moursy had incorrectly recognized B. fas- ciatus as occurring in North America, for reasons unknown to me. My examination of the type of B. cyclophorus and conspecific specimens from throughout North Ameri- ca, typical B. fasciatus from Europe, and reference to taxonomic treatments by John- son (1966), Paulus (1979), and earlier au- thors, clearly shows the regional misappli- cation of this name. Byrrhus geminatus LeConte Byrrhus geminatus LeConte 1854: 114, of El Moursy 1970: 330, Fiori 1982: 437. Holotype, female: “Light blue disc with two converging tangential cuts indicating the northern shore of Lake Superior; Type 2297; B. geminatus LEC., L. Sup.” (MCZ). Byrrhus pettiti Horn 1870: 76, synonymy by El Moursy 1970: 330, Fiori 1982: 438. Lectotype here designated: Female; ““Can.; B. pettiti Horn; Lectotype 3260” (MCZ). Paralectotype: same data (MCZ). Neither El Moursy (1970) or Fiori (1982) examined the type material of B. pettiti, and did not designate a type; however, they did correctly synonymize it with B. geminatus. The label reading ““Lectotype 3260” is ap- parently a cataloging label and has no tax- onomic standing, and was apparently placed on the specimen while the Horn Collection was held by the Academy of Natural Sci- ences, Philadelphia (A. Newton, pers. comm). Byrrhus kirbyi LeConte Byrrhus kirbyi LeConte 1854: 114, of El Moursy 1970: 331. Holotype, female: “light blue disc with two converging tan- gential cuts indicating the northern shore of Lake Superior” (MCZ). 326 Byrrhus fulvovestitus Casey 1912: 27, of El Moursy 1970: 329. NEw COMBINATION. Neither LeConte nor any subsequent worker labelled the above designated spec- imen of B. kirbyi in a manner indicating its identity or status. The entire byrrhid series in the LeConte collection was examined and only one specimen was found which fit LeConte’s description and possessed the correct label for the type locality. The spec- imen had been placed next to specimens of B. concolor Kirby and Porcinolus undatus (Melsheimer) in an unlabelled series, pos- sibly for comparative purposes. This spec- imen is herein considered LeConte’s unique type of B. kirbyi, and has been so labelled. El Moursy (1970) provided a lectotype designation, but incorrectly syonymized B. fulvovestitus with B. cyclophorus, apparently under the impression that Byrrhus females are not determinable to species. Casey’s specimens are badly rubbed and slightly narrow in form, but otherwise are typical B. kirbyi. Byrrhus pilula Linnaeus Recently, a series of Byrrhidae from northern Canada was gifted, with a request for determination by J. Pilny, Waterloo University, Ontario. Included were 78 spec- imens of a Byrrhus which could not be ac- curately assigned to any described or re- corded North American species; however, its close affinity with B. americanus Le- Conte was evident. Further investigation has shown that these specimens are conspecific with examples of B. pilula, of British and Austrian provenance. North American specimens ascribed to B. pilula have been seen from the following locality: Canada, Northwest Territories, Keewatin, Lat + 62.41 Long +97.03, What- ever Lake, June 1989. All specimens were collected by pitfall traps in tundra and ri- parian habitats. Specimens are in my col- lection and representatives will be depos- ited in the USNM and CNC. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Until now, B. pilula has been considered only a Eurasian species (Dalla Torre 1911, Winkler 1926, Fiori 1951, Horion 1955, Bonadona 1975). The lack of previous rec- ognition of this species in North America is ascribed to inadequate series of material, especially males, and to the failure of the most previous byrrhid workers to appreci- ate the potential of Holarctic distributions in the family (see also Johnson, submitted). Further complication of determination can be ascribed to a close relationship of B. pi- lula and B. americanus, as indicated by ae- deagal morphology, and a general difficulty of correlating unassociated Byrrhus females with males; this latter difficulty may also help explain occasional confounding of B. pilula with B. cyclophorus by previous workers. Porcinolus undatus (Melsheimer) Byrrhus undatus Melsheimer 1844: 117. Byrrhus murinus Fabricius 1794: 437, of LeConte 1854: 115. Porcinolus undatus (Melsheimer), of Casey 1912: 33. Holotype, female: ““Melsh.; un- datus M.; murinus; [red torn paper]” (MCZ). Byrrhus glabellus Melsheimer 1844: 117, of Casey 1912: 33. Holotype, sex not con- firmed: ‘“Melsh.; glabellus *Melsh.; [red torn paper] (MCZ). Porcinolus crescentifer Casey 1912: 32 NEw SyNonyM. Holotype, sex not confirmed: “Baldwin, Kansas; Casey bequest 1925; Type USNM 48354; crescentifer Csy” (USNM). Porcinolus hystrix Casey 1912: 33 NEw SyNonyM. Lectotype here designated, sex not confirmed: ‘““Aweme, Manitoba, S. Criddle, 23.VI.10; Casey bequest 1925; Type USNM 48355; hystrix Csy.” Para- lectotype: same data (USNM). All of the Melsheimer and Casey names represent simple color pattern variations of the pubescence. Porcinolus undatus is dis- tributed from the Atlantic coast to the east- VOLUME 93, NUMBER 2 ern slopes of the Rocky Mountains, with the western specimens being slightly small- er. The transelytral crescentiform pattern is highly variable from well colored and con- tiguous to indistinctly colored and broken in outline. Cytilus alternatus (Say) Byrrhus alternatus Say 1825: 186, of Le- Conte 1854: 115. Cytilus alternatus (Say, of LeConte 1870: 398, Casey 1912: 18. Type not found, pre- sumed lost (LeConte 1859, Lindroth & Freitag 1969). Byrrhus trivittatus Melsheimer 1844: 117, original synonymy by LeConte 1854: 115, LeConte” 1877; 108, Casey 192: 18, Hatch 1961: 297. Holotype, sex not con- firmed: “‘Melsh.; varius; red torn paper; trivittatus ‘Melsh.’’’ (MCZ). Cytilus nigrans Casey 1912: 19 NEw Synonym. Holotype, female: “Little Riv- er, Colroy, NFLD, July 10-18 ‘07; Casey bequest 1925; Type USNM 48331; ni- grans Csy” (USNM). My interpretation of C. alternatus is based on LeConte’s understanding of this species as represented by his specimens at the MCZ. The type of Cytilus nigrans is badly rubbed of pubescence, as noted by Casey (1912), while that of C. trivittatus lacks aeneous or viridescent elytral patches. Otherwise these specimens are typical examples of C. alter- natus. Cytilus mimicus Casey Cytilus mimicus Casey 1912: 18 NEw STATUS. Cytilus alternatus mimicus Casey 1912: 18. Lectotype here designated, sex not con- firmed: “‘Cal.; Casey bequest 1925; Type USNM 48329; mimicus Csy’” (USNM). Paralectotypes: 2, same data (USNM). Cytilus alternatus longulus Casey 1912: 18, Hatch 1961: 297 New ComBINATION. Lectotype here designated, sex not con- firmed: ““W.T.; Casey bequest 1925; Type 327, USNM 48330; longulus Csy” (USNM). Paralectotypes: 10, same data (USNM). Cytilus mimicus differs from C. alternatus by its larger size and more elongate form, slight differences in aedeagal structure, and habitat. This proposed new status for C. mimicus 1s an attempt at better recognition of two ecologically segregated, but geo- graphically sympatric forms of Cytilus in North America, which are difficult to di- agnose morphologically. Considerable ef- fort is still required for a full evaluation of both species and their relationship to Eur- asian species. Curimopsis echinata (LeConte) Syncalypta echinata LeConte 1850: 224, of LeConte 1854: 114. Curimopsis echinata (LeConte), of Casey 1912: 34, Johnson 1986: 42. Holotype, female: light blue disc indicating the Lake Superior region (MCZ). Curimopsis brevicollis Casey 1912: 35, of Hatch 1961: 299, synonymy noted by Johnson 1986: 42. Holotype, male: “W.T.; Casey bequest 1925; Type USNM 48356; brevicollis Csy.”” (USNM). Morphological differences between C. brevicollis and C. echinata used by Casey (1912) seem to be allometric and sexual. The identical aedeagi and female gonocox- ites of the types and other specimens sup- port this synonymy. Curimopsis albonotata (LeConte) Syncalypta albonotata LeConte 1861: 344. Curimopsis albonotata (LeConte), of Casey 1912: 34, Hatch 1961: 299, Johnson 1986: 42. Holotype, female: ““W.T.; S. albono- tata LEC.; Type 2302” (MCZ). Syncalypta grisea LeConte 1879: 514. Curimopsis grisea (LeConte), of Casey 1912: 35, synonymy noted by Johnson 1986: 42. Curimopsis brevicollis, Hatch 1961: 299. Holotype, female: “Garland, Col., 19.6; 3762S.erlsca LEG salype.23007 ((MCZ): 328 It is difficult to understand why LeConte did not observe the quite obvious similarity of these two species. His own descriptions indicate only minor color shading and size differences between the specimens, and di- rect comparison reveals these same differ- ences. Conspecificity is supported by genital morphology. Sierraclava cooperi Johnson This species was originally recorded (Johnson 1982) from the vicinity of Sequoia National Park, in the southern Sierra Ne- vada, California, U.S.A. Since then, addi- tional material has accumulated from 8 new disparate locations through central and southern California, as well as a new coun- try record from Baja California Norte, Mex- ico. These new records [abbreviated data] are: U.S.A., California, Amador Co., Elec- tra, Mokelumne River; Calaveras Co., 3 mi south of Mokelumne Hill; Fresno Co., 3 mi northeast of Auberry, 8 mi southwest of Au- berry, and 9 mi east of Coalinga; Riverside Co., Lamb Canyon, 2 mi northwest of Gil- man Hot Springs; San Benito Co., 18.4 mi northwest of New Idria, and 1.8 mi south- west of [New] Idria; Mexico, Baja California Norte, 10.7 km east of El Rosario, Lat + 30.04.35 Long +115.38.25, 11.7 km east of El Rosario, Lat +30.04.30 Long +115.37.55, and 14.7 km east of El Rosa- rio, Lat +30.04.10 Long + 115.36.00. Most specimens were collected in pitfall traps during late winter and spring months. Cal- ifornia specimens were collected largely by the staff of the California Department of Food and Agriculture, Sacramento, and the Mexico specimens by William H. Clark, College of Idaho, Caldwell. The apical portion of the penis of the Mexican specimens is much less broadened and spatulate than of northern specimens from Sierran localities, and specimens from Coast Range sites are intermediate in penial form. Coast Range and Mexican specimens have been found in desert or semidesert communities dominated by Juniperus-Ar- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON temisia scrub, Adenostoma chaparral, or mixed sarcophyllous scrub, while Sierran specimens were found in Quercus-Pinus woodlands. Whether the morphological and habitat differences can be refined to indicate two allopatric species is still being investi- gated. KEY TO THE ADULTS OF NorTH AMERICAN ByYRRHIDAE la. Antennae filiform, compressed; palps with ultimate segment securiform; appendages not retractile; tarsomere 3 with large fleshy lobe; integument appearing glabrous, rufocasta- neus to piceous ..... Amphicyrtinae ..... 2 1b. Antennae short, clavate or capitate; palps with ultimate segment fusiform or pyri- form: 222 aes ee eee 3 2a. Usually larger, 8-12 mm, elongate; integu- ment usually rufocastaneus with metallic cu- preous sheen, occasionally with viridescent reflections; coastal coniferous forests of Or- egon and northern California ............ ee oe Amphicyrta chrysomelina Erichson 2b. Usually smaller, 5-10 mm, ovoid, occasion- ally inflated posteriorly; integument rufo- piceous to piceous, without metallic sheen; prairies and oak-pine woodlands of central California and southwestern Oregon Ree OA READ Ge Amphicyrta dentipes Erichson 3a. Antennae capitate; body small (0.9-2.6 mm), shallowly to moderately convex; appendages closely retractile; dorsum with clavate or truncate bristles ..... Syncalyptinae 3b. Antennae clavate; body various (1.9-8.7 mm), moderately to strongly convex; ovate to elongate; appendages not or partially re- ceived into fossae; dorsum with decumbent to erect fine setae Byrrhinae 4a. Length 2.4—3.2 mm; ovoid, dorsum with ap- pressed scale-like setae and erect bristles . 5 4b. Length 0.9-2.1 mm; ovate, strongly convex; elytral striae punctate; dorsum with bristles only; northern U.S. and southern Canada, Europe eee: Chaetophora spinosa (Rossi) 5a. Ovate to oval, shallowly to moderately con- vex, lateral margins arcuate; striae on disc shallowly impressed, or serially punctate . 6 5b. Elongate, strongly convex, parallel-sided; el- ytra with striae deeply punctate and im- pressed, sulciform; central and southern California, Baja California Norte BS Hake Ae aes Sierraclava cooperi Johnson 6a. Length 2.1-2.8 mm, subparallel laterally; appressed squamae linear; median lobe of aedeagus slender, narrowly acuminate api- VOLUME 93, NUMBER 2 6b. as Tb. 8a. 8b. 9a. 9b. 10a. 10b. lla. 11b. 12a. 12b. cally -Alaskagierrrr..ceretme bere etnias Bek cere Curimopsis setulosa (Mannerheim) Length 2.8—3.2 mm; oval in lateral outline; aedeagus not as above Dorsal squamae round, set into round punc- tures; 3.0-3.2 mm; subparallel laterally; western and northern U.S., Canada Rie eae Curimopsis albonotata (LeConte) Dorsal squamae more-or-less linear to nar- rowly subtriangular, emergent from small usually barely evident punctures; usually smaller Body oval, rounded laterally; striae not im- pressed as lines or shallow sulci, usually rep- resented by rows of punctures; south-central Wiss ta cee hee Curimopsis strigosa (Melsheimer) Body elongate, subparallel laterally; striae narrowly, shallowly impressed, not evident- ly punctate Length 2.9-3.2 mm; appressed squamae short, subtriangular; elytral apical declivity long, gradually sloping; sutural stria at de- clivity shallowly impressed; northeastern WEeSenCanadawAlaskay-en sosen aoe eenee ae: Curimopsis moosilauke Johnson Length 2.8-2.9 mm; appressed squamae longer, linear; elytral apical declivity short- er, abruptly sloping; sutural stria at declivity distinctly and deeply impressed, sulcate; northern U.S., Canada, Alaska 5 ee oe Curimopsis echinata (LeConte) Frontal margin beaded, occasionally re- flexed and carinate; mesosternum strongly reduced except for median fossa; palps with ultimate segment fusiform to narrowly pyr- iform; metacoxae small, distant from elytral margin; elytra often connate Frontal margin rounded or truncate, never reflexed, beaded or carinate; mesosternum distinct laterad of median fossa; palps with ultimate segment compressed or cylindri- cally pyriform; metacoxae flattened, nearly reaching elytral margin; elytra not connate 19 Elytra separate and metathoracic wings pres- ent; integument rufopiceous to piceous, oc- casionally with submetallic reflection; me- socoxae moderately separated ........... 12 Elytra connate, metathorax apterous; me- socoxae widely separated ............... 14 Form short, ovate, slightly depressed dor- sally; elytral pubescence long, moderately dense, evenly distributed; elytral stria be- coming evanescent at midlength; northeast- ern U.S., southeastern and southwestern Canadawann ae Simplocaria semistriata (F.) Form elongate, ovoid; elytral pubescence lisa‘ 13b. 14a. 14b. 1Sa. 15b. l6a. 16b. 17a. 17b. 18a. 18b. 19a. 19b. 329 tessellate; elytral striae complete or becom- ing evanescent preapically Length 3.54.5 mm; pronotal punctures fine and sparse on disc; elytral striae becoming evanescent towards apex; pubescence de- cumbent; northern Canada, Greenland, Eur- asiaeneneece Simplocaria elongata J. Sahlberg Length 2.8-3.5 mm; pronotal punctures moderate on disc; elytral striae complete to apex; pubescence recumbent to suberect; northern U.S., Canada, Greenland, Eurasia Simplocaria metallica (Sturm) Pubescence simple, slender, fine; epipleura flat; integument shining to metallic ...... 16 Pubescence stout, recurved; epipleura deep- ly emarginate to receive femoral knee; in- LesuIMeENntiDrUNnCOUSE ee Fae eee 15 Length 1.8-2.3 mm; lateral elytral margin crenate anteriorly; coastal Oregon, Wash- ington, British Columbia EER AR AY at Ae ae Exomella pleuralis (Casey) Length 2.3-2.4 mm; lateral elytral margin smooth throughout; northern Idaho ities meh eee oY Exomella merickeli Johnson Epipleura broad, extending length of ely- tron; dorsal integument piceous to oliva- CEQUSHSHININ Sure eee ee eee Ia Epipleura narrow, attenuating and termi- nating before second visible abdominal ster- nite; dorsal integument viridescent ....... 18 Larger, 3.2-4.1 mm, lateral margins inflated; integument piceous, shining, occasionally with submetallic reflections; pubescent sparse to moderate, short to long; north-coastal California to southeastern Alaska, and west of Cascade Range crest in Oregon, Wash- ington, British Columbia Lioon simplicipes (Mannerheim) Smaller, 2.9-3.6 mm, lateral margins weakly rounded; integument piceous, olivaceous, without submetallic reflections; pubescence moderately dense, long; northern Idaho .. Ro) Nae Rn nih oe ee Lioon nezperce Johnson Length 2.4-2.6 mm; dorsal integument bright viridescent; west of Cascade Range crest in Oregon, Washington, British Columbia . syne eat Bin Me. Lioligus nitidus (Motschulsky) Length 2.2-2.4 mm; dorsal integument ol- ivaceous with evanescent viridescent reflec- tions; northern Idaho .. Lioligus pallidus Casey Frontal margin truncate and thickened; ab- dominal sternite 1 without crural modifi- CatlOns6 ey} Sak Pe By ee Pe sea eon 20 Frontal margin obtusely rounded; abdomi- nal sternite | with distinct crural depressions OMTOSSASMR ASR ae eee eS 22 330 20a. Diltae 21b. D2ar 22b. 23a. 230: 24a. 24b. 250% PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Form subparallel, moderately convex dor- sally; piceous, without metallic patches or reflections; pubescence long; decumbent, with erect black setae; northern U.S., Can- ada, Greenland ... Tylicus subcanus (LeConte) . Form ovoid to subparallel laterally, strongly convex dorsally; piceous, frequently with cu- preous or viridescent patches or intervals; pubescence short, appressed, without erect SCAG ag ye PR rn OEE Love niais or Seowebioye 21 Form ovoid, short; integument rarely with cupreous or viridescent patches; pubescence black and cinereous, tessellate on alternate elytral intervals; transcontinental Se os oh ooo aoe Cytilus alternatus (Say) Form elongate to subparallel; elytra usually with viridescent patches or intervals; pu- bescence pale cinereous, rarely tessellate; montane western U.S. and Canada Cytilus mimicus Casey Tarsomere 4 with membranous ventral lobe; dorsal integument shining, usually virides- Cent and/OMCcupreousysas eee 23 Tarsi simple; dorsal integument opaque or shining. piceous to) black =. .....45. 420-6. Dai Form ovate; aeneous to viridescent; punc- tures obsolescent or coarse and forming ru- gose patches on elytra; pubescence minute or moderately long and patchy, usually mixed cinerous and rufobrunneous ............. 24 Form subparallel; piceous with viridescent reflections to cupreous; punctures fine to moderate; pubescence moderately long, evenlvadistnibutedeer aa na oe. 26 Length 6.5—10.0 mm; dorsal punctures fine; subglabrous, pubescence minute; aeneous to viridescent, with elytra bearing distinct ae- neous and viridiaeneous vittae; northern Idaho, northeastern Washington, western Montana, southeastern British Columbia ade see. See Eusomalia lecontei (Wickham) Length 3.2-4.0 mm; punctures coarse, pu- bescence usually evident, moderately long 25 Dorsal punctures small to moderately coarse, simple, forming rugose patches with little suggestion of strial formation, or smooth with finely punctate striae on elytra; aeneous to viridiaeneous, often vittate on elytra; north- coastal California to Sitka, Alaska, west of Cascade Range crest in Oregon, Washing- ton) BritishyGolumbial@er- see ee Sid bn Sm Le Listemus acuminatus (Mannerheim) . Dorsal punctures large, coarse, umbilicate, not condensing into rugose patches on ely- tra; piceous, with only faint aeneous reflec- tions; northern Idaho, northeastern Wash- INSTOHe eee Listemus kootenai Johnson 26a. 26b. 27a. 27b. 28a. 28b. 29a. 29b. 30a. 30b. 31a. 31b. 32a. 32D. 33a. 33b. Integument piceous, frequently with viri- descent reflection; punctures moderate to dense; pubescence cinereous with dark brun- neous patches; montane western U.S., Can- ada eter suns ae Morychus aeneolus (LeConte) Integument aeneous to cupreous, occasion- ally with viridescent reflections; punctures fine to moderately dense, moderately to sparsely distributed; pubescence mixed ru- fobrunneous and cinereous; western U.S. and Canadamareeee ee Morychus oblongus (LeConte) Form oval to subparallel, strongly convex dorsally; pubescence simple, appressed ... 28 Form ovate, shallowly convex dorsally; pu- bescence including erect, bristle-like setae; central and eastern U.S., southern Canada ph Aire te eA Porcinolus undatus (Melsheimer) Pronotal and elytral integument shining be- tween moderate sized punctures; sparsely and finelysmicroreticulateiss..-= ee eee 29 Pronotal and elytral integument dulled due to dense, fine to coarse microreticulation or micorugosities; punctures small, often oblit- eratedh aac eat Oo era 32 Pronotal punctures smaller, well separated on disc by interspaces greater than own di- AMECLEL 5. enero Ea! Ere ee 30 Pronotal punctures larger, separated by less than own diameter; northern U.S., Canada ee en tt Byrrhus geminatus LeConte Punctures of head coarse, confluent, rugose; similar but shallower on elytra Punctures of head large and shallow on frons; deep and moderately dense on elytra; north- ern U.S., Canada .. Byrrhus cyclophorus Kirby Punctures on pronotal disc small, subequal or only slightly larger in diameter than base of seta; northeastern U.S., southeastern Can- Aa a nasa. eer Byrrhus americanus LeConte Punctures on pronotal disc larger, 2-3 x wider than seta; northern Canada, Eurasia RAE ee aren eT ure LAREN dy. Byrrhus pilula L. Punctures on elytra larger, distinct; sculp- turing: weak: 950s aaecerencr rice 33 Punctures on elytra minute, often obliter- ated; sculpturing transverse, reticulate, mi- crorugose, or undulating; northeastern U.S., Canadas. axe eee Byrrhus concolor Kirby Length >6.5 mm; elytral punctures smaller, dense but separated, rarely rugulose; north- ern and western U.S., Canada syns hay cel epee Ars See Byrrhus kirbyi LeConte Length <6.0 mm; elytral punctures dense, usually coalesced, rugulose; western U.S. and Canadaefee en eee Byrrhus eximius LeConte VOLUME 93, NUMBER 2 ACKNOWLEDGMENTS My thanks are extended to J. M. King- solver, U.S. National Museum of Natural History, Washington, D.C.; A. F. Newton, Jr. (formerly) and D. Furth, Museum of Comparative Zoology, Cambridge; H. Uh- lig, Humboldt-Universitat, Berlin; J. M. Campell, Agriculture Canada, Ottawa; R. L. Davidson, Carnegie Museum of Natural History, Pittsburgh; F. G. Andrews, Cali- fornia Department of Food and Agriculture, Sacramento; and William H. Clark, College of Idaho, Caldwell, for the loan of material and/or permission to examine material un- der their care. Thanks are also extended to the entomologists of the CNC and Carleton University, Ottawa, for their hospitality and visitation grants from the CanaColl Foun- dation. Thanks are extended to A. F. New- ton, Jr., S. E. Miller, for their comments on early drafts, J. M. Kingsolver for consid- erable assistance with specimens, informa- tion, and reviewing of manuscripts, and to D. S. Chandler for comments on the draft key. LITERATURE CITED Bonadona, P. 1975. Les Byrrhus (sensu lato) de France (Col., Byrrhidae). L’Entomologiste 31(6): 193-209. Brown, W. J. 1932. New species of Coleoptera, II. Canadian Entomologist 64: 8-9. Casey, T. L. 1908. A new genus of Byrrhidae. Ca- nadian Entomologist 40: 281-282. . 1912. Descriptive catalogue of the American Byrrhidae. Memoirs on the Coleoptera 3: 1-69. 1914. Miscellaneous notes and new species. Memoirs on the Coleoptera 5: 378. Dalla Torre, K. W. von 1911. Fam. Byrrhidae, pp. 5-36. In Schenkling’s Coleopterorum Catalogus, pars 33. W. Junk, Berlin. El Moursy, A.A. 1970. The taxonomy of the Nearctic species of the genus Byrrhus Linnaeus (Coleoptera: Byrrhidae). Quaestiones Entomologica 6: 327-338. Erichson, W. F. 1843. [revisionary notes and mis- cellaneous species descriptions]. /nSteffhany, G., ed., Tentamen Monographiae Generis Byrrhi. Zeitschrift fiir die Entomologie 4: 1-42. Fall, H. C. 1907. The Coleoptera of New Mexico: Descriptions of new species. Transactions of the American Entomological Society 33: 145-272. Fiori, G. 1951. Alcuni Appunti sui Byrrhus L., s.str., 331 Europei: II Contributo alla conoscenza della fa- miglia Byrrhidae (Coleoptera). Bolletina de Insti- tut Entomologia Universite, Bologna 18: 293-304. . 1982. Byrrhus geminatus LeConte: Specie ad ampia geonemia olartica, XIII Contributo all co- noscenza della famiglia Byrrhidae (Coleoptera). Bolletina de Museo Divisio Storia Naturella, Ve- rona 9: 437-447. Forster, J. R. 1771. Novae Species Insectorum. Cen- turia I. T. Davies & B. White, London. Hamilton, J. 1894a. Catalogue of the Coleoptera of Alaska, with the synonymy and distribution. Transactions of the American Entomological So- ciety 21: 1-38. . 1894b. Catalogue of the Coleoptera common to North America, northern Asia, and Europe with distribution and bibliography. Transactions of the American Entomological Society 21: 345-416. Hatch, M.H. 1961. The Beetles of the Pacific North- west, part III: Pselaphidae and Diversicornia I. University of Washington Press, Seattle. Horion, A. 1955. Faunistik der Mitteleuropaischen Kafer: Sternoxia, Fossipedes, Macrodactyla, Bra- chymera. Entomologische Arbeiten 4: 222-249. Horn, G. H. 1870. Contribution to the coleopter- ology of the United States. Transactions of the American Entomological Society 3: 69-142. Johnson, C. 1966. The British species of the genus Byrrhus L., including B. arietinus Steffahny (Col., Byrrhidae) new to the British list. Entomological Monthly Magazine 101: 111-115. Johnson, P. J. 1985. Anew species of Exomella from Idaho, with notes on the biology of Exomella pleuralis (Casey) (Coleoptera: Byrrhidae). Cole- opterists Bulletin 39(2): 151-157. . 1986. Anew species and a key to the Nearctic species of Curimopsis Ganglbauer (Coleoptera: Byrrhidae). Coleopterists Bulletin 40(1): 37-43. . 1990. Notes on the naturalization of two spe- cies of European Byrrhidae (Coleoptera) in North America. Journal of the New York Entomological Society 98(4): 434-440. . 1991 (In press). Taxonomic reviews of Lioon Casey and Listemus Casey, with descriptions of two new species (Coleoptera: Byrrhidae). Proceed- ings of the Entomological Society of Washington. (submitted). Synonymical notes for some species of the metallica-group of Simplocaria Ste- phens (Coleoptera: Byrrhidae). Entomologica Fen- nica. Johnson, P. J. and L. K. Russell 1978. Notes on the rediscovery, habitat, and classification of Exo- mella pleuralis (Casey) (Coleoptera: Byrrhidae). Coleopterists Bulletin 32(2): 159-160. Kirby, W. 1837. Northern zoology, part IV, Insecta, pp. 11i-xxxix, 1-325. Jn Richardson, J., ed., Fauna 332 Boreali-Americana: Or the Zoology of the North- ern Parts of British America . . . Josiah Fletcher, Norwich. LeConte, J. L. 1850. General remarks on the Cole- optera of Lake Superior, pp. 224-225. In Agassiz, J. L. R. and J. E. Cabot, eds., Lake Superior: Its Physical Character, Vegetation, and Animals, Compared with Those of Other and Similar Regions. Gould, Kendall, Lincoln, Boston. . 1854. Synopsis of the Byrrhidae of the United States. Proceedings of the Academy of Natural Sciences, Philadelphia 7: 114-117. 1857. Report upon insects collected on the survey, Zoological Report, part I, pp. 1-72. In Explorations and Surveys for a Railroad Route from the Mississippi River to the Pacific Ocean. War Dept. Route near the 47th and 49th Parallels, Explored by I. I. Stevens, Governor. Washington, D.C. (1860 reprint) . 1859. The Complete Writings of Thomas Say on the Entomology of North America, Vol. 1. New York. 1861. New species of Coleoptera inhabiting the Pacific district of the United States. Proceed- ings of the Academy of Natural Sciences, Phila- delphia 13: 338-359. 1863. New species of North American Co- leoptera, part I. Smithsonian Miscellaneous Col- lections 6(167): 1-92. . 1870. Synonymical notes on North American Coleoptera. Annals and Magazine of Natural His- tory, ser. 4, 6: 394-409. 1877. List of Coleoptera. Reports on Prog- ress, Canadian Geological Survey, 1875-76, 5: 107- 109. 1879. The Coleoptera of the alpine Rocky PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mountain regions, part II. Bulletin of the United States Geological and Geographical Survey of the Territories 5(3): 499-520. Leng, C. W. 1920. Catalogue of the Coleoptera of America, North of Mexico. John D. Sherman, Mount Vernon, New York. Lindroth, C. H.and R. Freitag 1969. North American ground-beetles (Coleoptera: Carabidae, excluding Cicindelinae) described by Thomas Say: Desig- nation of lectotypes and neotypes. Psyche 76(3): 326-361. Mannerheim, C. G. 1852. Zweiter Nachtrag zur Kaefer-fauna der North-Amerikanischen Laender des Russischen Reiches. Bulletin de la Société Im- périale de Naturalistes, Moscou 25(4): 283-387. Melsheimer, F. W. 1844. Descriptions of new Co- leoptera of the United States. Proceedings of the Academy of Natural Sciences, Philadelphia 2: 98- 118. Motschulsky, V. 1845. Observations sur le Musee Entomologique de L’Université Impériale de Moscou. Bulletin de la Société Impériale de Na- turalistes, Moscou 18(4): 332-387. Paulus, H. F. 1979. Familie: Byrrhidae, pp. 328-351. In Freude, H., K. W. Harde, and G. A. Lohse, eds., Die Kafer Mitteleuropas, vol. 6. Goecke & Evers, Krefeld. Say, T. 1825. Descriptions of new species of cole- opterous insects. Journal of the Academy of Nat- ural Sciences of Philadelphia 5: 160-204. Wickham, W. F. 1903. The North American species of Pedilophorus. Canadian Entomologist 35: 179- 182. Winkler, A. 1926. Catalogus Coleopterorum Region- is Paléarcticae. Wien. PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 333-355 SYSTEMATIC ANALYSIS OF ACANTHOCOCCUS SPECIES (HOMOPTERA: COCCOIDEA; ERIOCOCCIDAE) INFESTING ATRIPLEX IN WESTERN NORTH AMERICA DouGLAss R. MILLER Systematic Entomology Laboratory, Agricultural Research Service, U.S. Department of Agriculture, Beltsville, MD, USA 2070S. Abstract. —Salt bush (Atriplex spp.) (Chenopodiaceae) is an important source of forage for livestock at critical times of the year. Salt bush species are declining and suffering dieoff over extensive areas in the Great Basin and on the Colorado Plateau of western North America (Nelson et al. in press). Surveys of potential insect pests of several salt bush species have been conducted and several members of the scale insect genus Acan- thococcus (= Eriococcus) have been discovered. The purpose of this paper is to provide systematic information on the Acanthococcus species that occur on Atriplex to assist ongoing research on dieoff problems. Included are seven species of which three are new and four are redescribed. Key Words: In the United States, the genus Acantho- coccus has been treated as a junior synonym of Eriococcus. However, the occurrence of distinctive enlarged tubular ducts on the type species of Eriococcus (Coccus buxi Fonsco- lombe) which are present on other species from Australia and Europe and are absent from the type species of Acanthococcus (Acanthococcus aceris Signoret) and most other species from around the world, have convinced me that Borchsenius (1948) was correct when he treated Acanthococcus and Eriococcus as valid and separate genera. Be- cause of this change, all of the U.S. species previously treated as members of Eriococ- cus, should now be considered as members of Acanthococcus. METHODS Terminology used in this paper follows Miller and McKenzie (1967), Miller and Gonzales (1975), and Miller (1984). When first reading the terminology for the en- Scale insect, eriococcid, saltbush larged setae there may be some confusion. There generally are two distinct sizes of en- larged setae, i.e. large-sized enlarged setae and small-sized enlarged setae. Unfortu- nately, these sizes are relative within a spe- cies; no consistent size criterion can be used to decide whether a seta is large sized or small sized among all species. However, in nearly all cases these relative sizes are dis- tinct and there should be no confusion when studying a particular species or specimen. In one or two cases a species may have setae that intergrade from very large to small, but this circumstance is unusual and is a useful character state. Leg measurements are taken on the outer surface of each segment. Counts of enlarged setae include all enlarged setae on the segment dorsal and ventral. All dor- sal setae are considered to be enlarged even though some may be quite small. Measure- ments and numbers are taken from 10 spec- imens when available and are given as a 334 range of numbers followed by the average in parentheses. Depositories of specimens are as follows: British Museum (Natural History), London (BM); California Department of Food and Agriculture, Sacramento (CDA); University of California, Davis (UCD); University of Hawaii, Honolulu (UH); National Museum of Natural History, Washington, D.C. (USNM); Virginia Polytechnic Institute and State University, Blacksburg (VPI); Zoolog- ical Institute, Academy of Sciences of USSR, Leningrad (ZAS). Other abbreviations are: specimen (spm.), slide (sl.), and ad. (adult). ACKNOWLEDGMENTS I am grateful to the following individuals for their criticisms and comments of this manuscript: C. Riley Nelson, Department of Zoology, University of Texas, Austin; John A. Davidson, Department of Ento- mology, University of Maryland, College Park; Robert L. Smiley and Sueo Nakahara, Systematic Entomology Laboratory, ARS, USDA, Beltsville, Maryland. Riley Nel- son’s suggestions were especially helpful as a potential user of the information provided in the publication. KEY TO ADULT FEMALES OF ACANTHOCOCCUS SPECIES THAT INFEST ATRIPLEX Ie Five setae on hind tibia Four'setae on hinditiblay = 4.5.2.4... 42) 2 Anal lobes each with 4 enlarged setae .... BT EA TEMAS Se ICE eee froebeae Miller, n. sp. Anal lobes each with 3 enlarged setae .... Loni aR: Se On As, i ayia tinsleyi (Cockerell) Enlarged setae with round or blunt apices .. 4 Enlarged setae with acute apices (Fig. 7) .. era eee on Ce Ca whiteheadi Miller, n. sp. Fewer than 65 enlarged setae on segment V including those on dorsum and venter ... 5 More than 65 enlarged setae on segment V including those on dorsum and venter ... Sol oS HOR ee eee ee barri Miller, n. sp. Enlarged setae not fusiform in shape Enlarged setae fusiform in shape (Fig. 5) .. Nae co ee tae ee, en ae salarius (Ferris) 6(5). Microtubular ducts without sclerotized ring 3(1). 4(3). 5(3). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON at dermal orifice; cruciform pores normally absent arenosus (Cockerell) Microtubular ducts usually with sclerotized ring at dermal orifice; cruciform pores pres- Cnto ach Se eee ee ae eriogoni (Ehrhorn) TREATMENT OF SPECIES Acanthococcus arenosus (Cockerell), New ComMBINATION Sand eriococcin Fig. 1 Eriococcus arenosus Cockerell, 1897, 1899, 1900; Fernald, 1903; Ferris 1955; Mc- Daniel, 1959; Hoy, 1963. Type material: From the syntypes I have chosen and marked as lectotype an adult female labeled “‘Eriococcus arenosus Ckll., On ?, N. Mex., Cockerell, April 16, 1897 Type” (USNM). The slide contains only 1 specimen. In addition, there are 4 paralec- totypes on 2 slides. Field features: Adult female elongate oval. Body varies from gray to light purple. A smooth, heavy, white ovisac may be inter- mixed with grains of sand. This species is found on the spines and branches of its host. Recognition characters: Adult female, mounted, 2.2-3.8 (3.0) mm long, 1.7-2.4 (2.1) mm wide. Anal lobes lightly sclero- tized ventrally; each lobe dorsally with 3 enlarged setae (size variable, either lateral seta equal to posteromedial seta, antero- medial seta shortest, or all setae equal in size), with from 1—5 (3) microtubular ducts; each lobe ventrally with 3-4 (3) body setae and 2-9 (4) sessile pores. DORSUM with enlarged setae of 2 pri- mary sizes: with 2 larger setae along margin of each abdominal segment, also present along thorax and head; remaining setae small. Largest large seta 29-44 (37) u long, largest small seta 19-44 (35) uw; on abdom- inal segments VIII through III longest large seta 1.2-1.7 (1.5) times longer than longest small seta. All enlarged setae slightly curved; those from northern areas slender, apices rounded; those from southern areas robust, VOLUME 93, NUMBER 2 335 Fig. 1. Acanthococcus arenosus (Cockerell). 14 miles W. Phoenix, Arizona, October 15, 1968, on Atriplex sp. 336 apices truncate; all with thin setal rings. En- larged setae ranging from infrequent to abundant—e.g. abdominal segment V with 22-55 (37)—large setae showing no longi- tudinal pattern. Macrotubular ducts vari- able in length (4-8 (6) uw long), with area farthest from dermal orifice sclerotized and divided into 2 parts, apical portion round- ed, equal or slightly shorter than remaining sclerotized portion; total sclerotized portion unusually short, varying from 0.5-2.0 (1.0) times length of unsclerotized portion; der- mal orifice with no sclerotized ring. Micro- tubular ducts scattered over surface. Anal ring either dorsal or ventral, with 4, rarely 5, pairs of setae. VENTER with lanceolate body setae long (longest seta on abdominal segment VIII from 32-64 (48) uw long, on segment III from 53-84 (67) u), medial setae rarely capitate. Enlarged setae same as on dorsum, except more slender, present along margins of ab- dominal segment VIII through head. Mac- rotubular ducts of 2 sizes: larger size on lat- eral areas only; smaller size on medial and sublateral areas of entire surface, most abundant on abdomen. Microtubular ducts restricted to lateral margins only, uncom- mon. Multilocular sessile pores of 3 kinds: septeloculars rarely present; quinquelocu- lars abundant on posterior margins of ab- dominal segment, rare on anterior margins of abdomen, thorax, and head; triloculars most abundant on thorax and head, also present on anterior margins of abdomen. Cruciform pores usually absent, rarely with a few near lateral margins of anterior ab- dominal segments, thorax, and head. Legs: hind coxae dorsally with 4-35 (15) pores, ventral surface with 0-25 (10); hind femora dorsally with 0-7 (3) pores, ventral surface with 0-2 (1); tibiae with 5 setae; inner, apical, tibial setae robust on hind 2 pairs of legs, lanceolate on front pair of legs; hind tarsi usually slightly longer than tibiae (hind tibia/tarsus ratio 0.89-1.06 (0.95)); claws with denticle near tip. Antennae 7-segmented, third or fourth segment lon- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gest. Segment 7 with 3 sensory setae; seg- ment 6 with | noticeably longer than single sensory seta on segment 5. Notes: There appear to be two extreme forms of this species, one that occurs in northern Oregon, northern Utah, and northern and central Nevada, and another that occurs in central and southern Nevada, Arizona, New Mexico, and Texas. The northern form is characterized by slender, enlarged setae with rounded apices; where- as, the southern form possesses robust, en- larged setae with truncate apices. Interme- diate forms occur in southern Nevada and northern Arizona. This species is similar to Acanthcoccus eriogoni (Ehrhorn), but differs in possessing: few or no cruciform pores, microtubular ducts without sclerotized ring orifice; A. erlogoni, on the other hand, possesses: many cruciform pores, microtubular ducts usually with heavily sclerotized ring orifice. Specimens examined: ARIZONA, MAR- ICOPA Co.: Phoenix, X-10-1899, on Atri- plex canescens (Chenopodiaceae), T. D. A. Cockerell (3 spm. on 2 sl.) USNM; 14 mi. W. Phoenix, X-15-68, on Atriplex sp., P. F. Min and Miller (2 ad. female on 2 sl.) CDA; Tempe, butte, VIII-(?)-18, on Atriplex sp., G. F. Ferris (1 ad. female) UCD. NEVADA, LYON Co: Weeks, VII-5-68, on Atriplex sp., D. R. Miller and R. F. Den- no (5 ad. female on 2 sl.) UCD. NYE Co.: Tonopah, VII-6-62, on (?), collector (?) (6 ad. female on 3 sl.) CDA. WASHOE Co.: Nixon, VI-24-64, on Bassia hyssopifolia (Chenopodiaceae), J. A. Froebe (1 ad. fe- male) UCD; near Reno, VII-2-47, on Atri- plex sp., G. F. Ferris (4 ad. female) UCD. WHITE PINE Co.: 3 mi. N. McGill on Atri- plex sp., D. R. Miller and R. F. Denno (1 ad. female) UCD; 6 mi. NE. McGill, on Atriplex sp., D. R. Miller and R. F. Denno (1 ad. female) UCD. NEW MEXICO, DONA ANA Co.: Las Cruces, date (?), on A. canescens, ““M. and F.”’ (1 spm.) USNM; VIII-4-66, on Atriplex sp., D. R. Miller (3 ad. female on 2 sl.) UCD; VOLUME 93, NUMBER 2 Mesilla Park, VIII-(?)-1898, on A. canes- cens, T. D. A. Cockerell (3 spm. on 2 sl.) USNM. RIO ARRIBA Co.: Embundo, IV- 26-1897, on Psoralea micrantha (Legumi- nosae), T. D. A. Cockerell (1 ad. female lectotype, 4 ad. female paralectotypes on 2 sl.) USNM. OREGON, LAKE Co.:: Alkali Lake, VIII- 4-68, on Gutierrezia sp. (Compositae), D. R. Miller and R. F. Denno (1 ad. female) UCD; 24 mi. E. Christmas Valley, VIII-4- 68, on Atriplex canescens, D. R. Miller and R. F. Denno (4 ad. female on 2 sl.) UCD; 9 mi. N. Valley Falls, VIII-4-68, on A. ca- nescens, D. R. Miller and R. F. Denno (5 ad. female on 3 sl.) UCD. TEXAS, PRESIDIO Co.: Presidio, XI- 19-43, on A. canescens, J. H. Russell (4 spm.) USNM; Presidio, IV-29-52, on A. canes- cens, J. H. Russell (5 spm.) USNM. UTAH, BOX ELDER Co.: 40 mi. SW. Rosette, VIII-2-67, on Atriplex sp., D. R. Miller and D. S. Horning (1 ad. female) UCD. KANE Co: Kanab, III-28-59, on Sar- cobatus vermiculatus (Chenopodiaceae), G. F. Knowlton (4 ad. female) USNM. Hosts and distribution: Found on Atri- plex, Gutierrezia, Psoralea, and Sarcobatus. Acanthococcus arenosus is the most com- monly collected on Atriplex. Distributed in arid areas of Arizona, Ne- vada, New Mexico, Oregon, Texas, and Utah. Acanthococcus barri Miller, New SPECIES Barr eriococcin Fig, 2; Type material: Adult female holotype (1 specimen on slide) with right label “‘Erio- coccus 2 mi. E. Tonopah, Nye Co., NE- VADA 7-VII-1968 Atriplex canenscens D. R. Miller and R. F. Denno 1230”’; left label “Eriococcus barri Miller Holotype TYPE” (deposited at UCD). In addition there are 116 paratypes. Field features: The body is white or light yellow; legs are yellowish-brown. The adult 337 female is heavily coated with many short, squat crystalline rods that give the body a white appearance. One rod on lateral mar- gin of each abdominal segment is longer and broader than the others; these rods are slightly curved posteriorly. The ovisac is noticeably tough and difficult to break open. This species occurs on the crown and roots of its host. Recognition characters: Adult female ho- lotype, mounted, 1.9 mm long, 1.2 mm wide (paratypes 1.5-3.3 (2.3) mm long, 1.0—2.5 (1.9) mm wide). Anal lobes slightly pro- truding, rounded, lightly sclerotized; each lobe dorsally with 3 enlarged setae (antero- medial seta longest and most slender, lateral seta shortest and most robust), with 4 mi- crotubular ducts; each lobe ventrally with 4 body setae and 4 sessile pores. DORSUM with enlarged setae of 3 sizes: 1 larger seta on margin of each abdominal segment, also present on lateral margin of thorax and head; | pair of smaller setae present on medial area and sublateral areas of each abdominal segment; remaining se- tae of intermediate size. Largest large seta 37 uw long (paratypes 30-38 (35) yw), largest intermediate seta 28 uw long (paratypes 25— 31 (28) u), largest small seta 16 u long (para- types 13-16 (15) yu); longest large seta 1.3 times longer than longest intermediate sized seta (paratypes 1.2-1.3 (1.2) times); longest large seta 2.3 times longer than longest small seta (paratypes 1.9-2.4 (2.1) times). Large setae extremely broad, with truncate or blunt apices; intermediate setae similar except apices more rounded; small setae relatively slender, with rounded apices; larger and in- termediate setae with setal rings so thin as to appear fused to main body of seta; small- er setae with thin, unfused setal rings. En- larged setae abundant—e.g. abdominal seg- ment V with 113 (paratypes with 69 to 111 (81))—small setae forming 2 pairs of lon- gitudinal lines (medial, sublateral), large se- tae forming | pair of longitudinal lines (lat- eral). Macrotubular ducts in small numbers over surface. Microtubular ducts elongate PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 338 9 dd XN 9 Acanthococcus barri Miller, n. sp. 2 miles E. Tonopah, Nevada, July 7, 1968, on Atriplex canescens. Fig) 2: VOLUME 93, NUMBER 2 (13 uw long) (paratypes 11-14 (13) wu), with area farthest from dermal orifice sclerotized and undivided; total sclerotized area ap- proximately 0.2 length of unsclerotized area; dermal orifice sclerotized. Microtubular ducts in small numbers over surface. Anal ring dorsal, with 4 pairs of setae. VENTER with lanceolate body setae elongate (longest seta on abdominal seg- ment VIII 47 uw long (paratypes 34—50 (42) a), on segment III 56 uw (paratypes 47-62 (57) «), medial setae with acute apices. En- larged setae present along lateral margin from abdominal segment VIII through tho- rax. Macrotubular ducts same as on dor- sum, scattered over surface, most abundant on lateral areas. Microtubular ducts most abundant on lateral areas, also present an- terior of each leg. Multilocular sessile pores in unusually small numbers on thorax, and of 3 kinds: septeloculars uncommon, pres- ent on posterior abdominal segments; quin- queloculars most abundant, present over abdomen and near spiracles; triloculars un- common, present on anterior abdominal segments and spiracles. Cruciform pores ab- sent. Legs: hind coxae dorsally with 45 and 43 pores (paratypes with 27-58 (43)), ventrally with 9 and 30 (paratypes with 18-48 (31)); hind femora dorsally with 6 and 5 (para- types with 1-7 (5)), ventrally with 0 and 2 (paratypes with 1-6 (4)); tibiae each with 5 setae; inner, apical, tibial setae unenlarged; tarsi slightly longer than tibiae (hind tibia/ tarsus ratio 0.88) (paratypes 0.82-0.98 (0.87)); claws with large denticle. Antennae 7-segmented, fourth segment longest. Seg- ment 7 with 3 sensory setae; segment 6 with 1 longer and more slender than single sen- sory seta on segment 5. Variation: Some of paratypes vary from holotype in possessing the following char- acteristics: cluster of 6 or 7 small sized setae on dorsum of abdominal segment VIII; an- teromedial seta on anal lobes may be unen- larged, appearing as a body seta; 1-4 mi- crotubular ducts on each anal lobe; sessile 339 pores normally absent on anal lobes; anal ring rarely ventral and with 3 pairs of setae; microtubular duct orifice ring often com- pletely unsclerotized; enlarged setae on ven- ter may be present from abdominal segment VIII through head; antennae rarely 6- or 8-segmented. Notes: I take great pleasure in naming this species A. barriin honor of W. F. Barr, Uni- versity of Idaho. He has provided speci- mens of this unusual species, and has also contributed many other Coccoidea collec- tions. His assistance has contributed greatly towards a more comprehensive understand- ing of the scale insects of the northwestern United States. This species is similar only to A. white- headi Miller. It differs in possessing: differ- ently shaped dorsal setae, and a different enlarged setal pattern. For detailed discus- sion see “notes” under A. whiteheadi. Specimens examined: IDAHO, CAN- YON Co.: 15 mi. S. Nampa, VI-27-53, on Atriplex confertifolia (Chenopodiaceae), W. F. Barr (3 ad. female paratypes on 3 sl.) UCD, CDA; IX-4-62, on Atriplex sp., W. F. Barr (6 ad. female paratypes on 6 sl.) UCD, VPI. ELMORE Co.: 15 mi. W. Mountain Home, VII-31-58, on Atriplex sp., W. F. Barr (4 ad. female paratypes on 4 sl.) BM, UCD, UH; IX-3-62, on Atriplex sp., W. F. Barr (4 ad. female paratypes on 3 sl.) UCD, ZAS. OWYHEE Co.: 5 mi. N. Mur- phy, VII-25-56, on Atriplex sp., W. F. Barr (2 ad. female paratypes on 2 sl.) UCD. NEVADA, ESMERALDA Co.: 6 mi. W. Tonopah, VII-6-68, on A. canescens, D. R. Miller and R. F. Denno (7 ad. female para- types on 5 sl.) UCD, USNM. NYE Co.: 2 mi. E. Tonopah, VII-7-68, on A. canescens, D. R. Miller and R. F. Denno (1 ad. female holotype, 32 ad. female paratypes, 26 first instar nymph paratypes, 19 ad. male para- types, 1 third instar male paratype, | fourth instar male paratype on 22 sl.) UCD; 28 mi. NE. Warm Springs, VII-7-68, on A. canes- cens, D. R. Miller and R. F. Denno (4 ad. female paratypes on 2 sl.) UCD. 340 Host and distribution: Found only on Atriplex. Probably occurring throughout much of the Great Basin. Acanthococcus eriogoni (Ehrhorn), NEw COMBINATION Eriogonum eriococcin Fig. 3 Eriococcus sidae Ferris, 1955; Hoy, 1963; McDaniel, 1964. Eriococcus sidae Ferris, 1955; Hoy, 1963; McDaniel, 1964. Type material: From the syntypes I have chosen and marked as lectotype an adult female labeled “‘Eriococcus eriogoni on Er- iogonum, Flagstaff Arizona Type” (USNM). There are 8 specimens on the slide; the spec- imen horizontally on the right and vertically in the middle is the lectotype. In addition, there are 7 paralectotypes. I have examined part of the type series of E. sidae. Field features: Adult female oval. Newly formed adult females vary from gray to green; becoming red with age. Crystalline rods numerous over entire dorsum giving eriococcin a wooly appearance. Present on roots and subterranean or ae- rial crown of hosts. Recognition characters: Adult female, mounted, 1.7-2.5 (2.2) mm long, 0.9-1.8 (1.2) mm wide. Anal lobes apically acute, moderately sclerotized; each lobe dorsally with 3 enlarged setae (lateral seta either equal to or larger than posteromedial seta, antero- medial seta shortest), with 0-3 (2) micro- tubular ducts; each lobe ventrally with 3, rarely 4, slender body setae and 2-9 (5) ses- sile pores. DORSUM with enlarged setae of 2 pri- mary sizes: | seta on margin of each ab- dominal segment and several on margins of thorax and abdomen large; remaining setae conspicuously smaller. Largest large seta 41- 67 (54) uw long, largest small seta 33-50 (40) u; on abdominal segments VIII through III PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON longest lateral setae, including large type, straight, with apices rounded, slightly more acute than apices of medial seta; medial sublateral setae conspicuously curve, with apices rounded; all with thin setal rings. En- larged setae abundant—e.g. abdominal seg- ment V with 27-55 (43)—these setae nor- mally showing no longitudinal pattern; although rarely there may be weak indica- tion of 3 pairs of longitudinal lines, these suggested by slightly larger setae, when pres- ent, these lines not visible anterior of ab- dominal segment V. Macrotubular ducts densely scattered over dorsum. Microtu- bular ducts moderate in length 6-8 (7) u long, with area farthest from dermal orifice sclerotized and weakly divided into 2 parts, apical portion rounded, from 0.5—1.0 (0.8) times length of remaining sclerotized por- tion; total sclerotized area varying from 1- 5 (2) times longer than unsclerotized area; dermal orifice varying from heavily scler- otized to totally unsclerotized. Microtubu- lar ducts abundant over surface. Anal ring ventral, rarely dorsal, with 3, normally 4, pairs of setae. VENTER with lanceolate body setae moderate in length (longest seta on abdom- inal segment VIII from 39-50 (45) u long, on segment III from 47-62 (55) u), medial setae stout, capitate. Enlarged setae nor- mally of small size, straight; present near body margin from abdominal segment VIII through head. Macrotubular ducts of 2 kinds: larger size present on lateral and sub- lateral areas; smaller size normally only slightly smaller than large size, present in medial areas from segment VII or VI through head. Microtubular ducts present over en- tire surface, most abundant near lateral margins. Multilocular sessile pores of 3 kinds: septeloculars least common, if pres- ent, usually on abdominal segment IX or VII; quinqueloculars most abundant, pres- ent over entire surface; triloculars present on anterior abdominal segments, thorax, and head. Cruciform pores present along lateral margin of anterior abdominal segments, on VOLUME 93, NUMBER 2 341 Fig. 3. Acanthococcus eriogoni (Ehrhorn). Flagstaff, Arizona, date and host unknown. 342 lateral areas of thorax and head, and ante- rior of each leg. Legs: hind coxae dorsally with 17-45 (27) pores, ventral surface with 4—20 (12); hind femora dorsally with 3-14 (8) pores, ventral surface with 0-5 (2); tibiae with 5 setae; inner, apical, tibial setae robust; tarsi longer than tibiae (hind tibia/tarsus ratio 0.78-0.93 (0.88)); claws with denticle near tip. Anten- nae 6-segmented, rarely with third segment divided forming seventh segment, third seg- ment longest. Apical segment with 2 or 3 sensory setae; second segment from apex with sensory setae absent; third segment from apex with only | which is short and robust. Notes: This is an extremely variable spe- cies. I believed for some time that it was a synonym of A. dubius, but after careful con- sideration, I have concluded that they are distinct. Because of the extreme variability within these species, it is not possible to use one character to separate them, instead, it is necessary to use a combination of char- acters. Acanthococcus eriogoni possesses: enlarged setae with rounded apices; strongly curved medial and sublateral setae; medial and sublateral setae all of approximately the same size, particularly on abdominal seg- ment V forward through head; front tibiae with 5 setae; and microtubular ducts with long area of sclerotization; A. dubius, on the other hand, possesses: enlarged setae with acute apices; straight or slightly curved me- dial and sublateral setae; medial and sub- lateral setae of 2 sizes, large size forming 3 pairs of longitudinal lines from abdominal segment VIII through posterior thorax; front tibiae normally with 6 setae; and microtu- bular ducts normally with short area of scle- rotization. See “notes” under EF. arenosus for an ad- ditional comparison. Specimens examined: ARIZONA, CO- CHISE Co.: 21 mi. N. Bisbee, VIII-2-66, on (?), D. R. Miller (1 ad. female) UCD. COCONINO Co.: Flagstaff, date (?), on (?), PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON O. E. Bremner (4 ad. female of 4 sl.) CDA, UCD; on Eriogonum sp. (Polygonaceae), collector (?) (1 lectotype ad. female and 7 paralectotypes on | sl.) USNM; IX-(?)-1900, on Eriogonum sp., O. E. Bremner (5 ad. female on 4 sl.) UCD; Yuba City, on Hap- lopappus acradenius (?) (Compositae), H. L. McKenzie (4 ad. female on 3 sl.) UCD. MARICOPA Co.: quarantined at Phoenix from Alamo, Texas, I-21-69, on Echinopsis sp. (Cactaceae), P. F. Min and Hancock (2 ad. female on 2 sl.) UCD, CDA; 5 mi. E. Phoenix, [X-3-68, on (?) (Chenopodiaceae), D. R. Miller and J. E. Lauck (2 ad. female) UCD. PIMA Co.: 35 mi. S. Tucson, Santa Rita Range, on Eriogonum wrightii, H. S. Haskell (17 ad. female, 7 second instar fe- male, 65 first instars, 3 second instar male on 6 sl.) UCD. CALIFORNIA, IMPERIAL Co.: 1 mi. W. Glamis, I-28-65, on Eriogonum sp., D. R. Miller (2 ad. female on 2 sl.) UCD; 12 mi. E. Holtville, XII-1 1-58, on Ephedra cal- ifornica (Ephedraceae), G. L. Osborn (12 ad. female, 5 second instar female, 3 first instar, 1 ad. male, 1 fourth instar male, 3 second instar male on 7 sl.) CDA, UCD; Imperial, XII-28-63, on “cactus,” L. Phipps and G. Skaggs (3 ad. female on 3 sl.) CDA. LOS ANGELES Co.: Lancaster, VI-12-18, on Haplopappus (= Isocoma) venetus, G. F. Ferris’ (6 ad. female on: 2°sl)) UCD: RIV- ERSIDE Co.: 18 mi. W. Blythe, I-29-65, on Palafoxia linearis (Compositae) and Atri- plex sp. (Chenopodiceae), D. R. Miller (4 ad. female on 4 sl.) UCD; 4 mi. W. Desert Center, II-20-58, “‘in soil,”’ E. I. Schlinger (3 ad. female) UCD; Desert Center, IV-18- 66, on Euphorbia sp. (Euphorbiaceae), R. J. Gill and L. L. Johnson (1 ad. female) CDA; X-5-66, on Eriogonum deflexum, D. Fis- kaali (1 ad. female, 3 second instar female, 3 second instar male on 6 sl.) CDA. SAN BERNARDINO Co.: 5 mi. S. Kramer Junc- tion, XII-28-64, I-24-65, on Ceratoides lan- ata (Chenopodiaceae), D. R. and J. F. Miller (3 ad. female on 2 sl.) UCD. SAN DIEGO VOLUME 93, NUMBER 2 Co.: Borrego Springs, I-27-65, on Eriogo- num inflatum, D. R. Miller (2 ad. female on 2 sl.) UCD. SISKIOU Co.: 7 mi. S. Yreka, VII-20-66, on Gutierrezia sp. (Compositae), D. R. Miller (1 ad. female) UCD. TULARE Co.: above Mineral King, date (?), on Er- iogonum sp., G. F. Ferris (2 ad. female) WED: FLORIDA, COUNTY (?): Fruit Cove, VII-27-66, on (?) (Cactaceae), A. E. Graham (2 ad. female on 2 sl.) CDA. NEVADA, LYON Co.: 5 mi. S. Wabus- ka, VII-5-68, Ceratoides lanata, D. R. Mil- ler and R. F. Denno (4 ad. female on 2 sl.) UCD. WASHOE Co.: Reno, I-2-59, on “cactus,” R. F. Rebuffo (6 ad. female on 3 sl.) CDA. TEXAS, BAILEY Co.: Muleshoe, (7?)-(?)- 21, on Meriolix serrulata (Onagraceae), G. Eo erernas (5 vad. female on 3) sl.) UCD: BREWSTER Co.: Chisos Mountains, (?)- (?)-21, on Paronychia jamesii (Caryophyl- laceae), G. F. Ferris (4 ad. female on 3 sl.) UCD. EL PASO Co.: near El Paso, date (?), on Sida hederae (Malvaceae), collector (?) (12 ad. female on 7 sl.) UCD. PECOS Co.: near Shefhield, Pecos River, (?)-(?)-21, on Croton sp. (Euphorbiaceae), G. F. Ferris (3 ad. female on 3 sl.) UCD. Host and distribution: Found on many plant genera: Atriplex, Croton, Echinopsis, Ephedra, Eriogonum, Euphorbia, Eurotia, Gutierrezia, Haplopappus, Meriolix, Pala- foxia, Paronychia, and Sida. There seems to be no particular host pattern. Distributed in warm areas of southwest- ern United States. Acanthococcus froebeae Miller, NEw SPECIES Froebe eriococcin Fig. 4 Type material: Adult female holotype (1 specimen on slide), left label ‘““Eriococcus froebeae Miller TYPE”; right label “‘5 mi. n. Baker, San Bernardino Co. Calif., on Franseria sp. IV-13-63, D. R. Miller’ 343 (UCD). In addition there are 2 paratypes (WED): Field features: This species occurs on the foliage of its host. Recognition characters: Adult female ho- lotype, mounted 2.2 mm long, 1.4 mm wide (paratypes 2.1—2.7 (2.4) mm long, 1.5-1.8 (1.7) mm wide). Anal lobes slightly pro- truding, acute, slightly sclerotized; each lobe dorsally with 4 enlarged setae (anterolateral seta smallest, remaining 3 setae approxi- mately equal), with 4 or 5 microtubular ducts; each lobe ventrally with 3 body setae and | sessile pore. DORSUM with setae of | size; largest seta 31 w long (paratypes 34-35 (35) uw), smallest seta 16 w long (paratypes 18-19 (18) uw); longest seta 2.0 times longer than smallest seta (paratypes | .8—1.9 (1.9) times). All setae slightly curved, slender, with rounded apices; setal rings thin. Enlarged setae abundant—e.g. abdominal segment V with 60 (paratypes 52-78 (62))—with no longitudinal pattern. Macrotubular ducts scattered over surface. Microtubular ducts moderate in length (6 u long) (paratypes 5— 6 (6) uw), with area farthest from dermal or- ifice sclerotized and divided into 2 parts, apical portion small, approximately 0.2 times length of remaining sclerotized por- tion; total sclerotized area shorter than un- sclerotized area; dermal orifice only weakly sclerotized. Microtubular ducts numerous over surface. Anal ring bent around abdomen apex, with 4 pairs of setae. VENTER with lanceolate body setae moderate in length (longest seta on abdom- inal segment VIII 37 uw long (paratypes 41- 44 (43)), on segment III 44 u (paratypes 56- 62 (58) u), medial setae with apices acute. Enlarged setae present along lateral margin from abdominal segment VIII through head. Macrotubular ducts of 2 kinds: larger size same as on dorsum, present along lateral margins and on medial and sublateral areas of thorax and head; smaller size present on 344 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Acanthococcus froebeae Miller. 5 miles N. Baker, California, April 13, 1963, on Franseria sp. VOLUME 93, NUMBER 2 medial and sublateral areas of abdomen. Microtubular ducts most abundant along lateral margins, also present on medial areas of anterior abdominal segments, thorax, and head. Multilocular pores of 2 kinds: quin- queloculars present over entire surface ex- cept thorax; triloculars rare. Cruciform pores absent. Legs: hind coxae dorsally with 20 and 25 pores (paratypes with 6-17 (12)), absent on ventral surface; hind femora dorsally with 4 and 6 pores (paratypes with 2-5 (4)), ab- sent on ventral surface; tibiae with 4 setae; inner, apical, tibial setae robust, tarsi slight- ly longer than tibiae (hind tibia/tarsus ratio 0.90) (paratypes 0.85—1.00 (0.9)); claws with small denticle near tip. Antennae 7-seg- mented, third segment longest. Segment 7 with 3 slender sensory setae; segment 6 with 1 slightly longer and more slender than sin- gle sensory seta on segment 5. Variation: The paratypes agree well with holotype. Notes: I take great pleasure in naming this species A. froebeae in honor of Judith Froebe Miller, my wife, who has helped me collect scale insects for more than 30 years, in- cluding this species. She has been especially tolerant and supportive of my obsession for Coccoidea systematics and for that I am eternally grateful. This species is distinct in possessing the following combination of characters: 4 setae on each tibia; enlarged seta of essentially | size scattered over entire dorsum; 4 en- larged setae on each anal lobe. Specimens examined: CALIFORNIA, RIVERSIDE Co.: 7 mi. N. Indio, III-26-64, on Atriplex sp. (Chenopodiaceae), D. R. Miller and J. A. Froebe (2 ad. female para- type) USNM; 11 mi. N. Indio, VIII-24-68, on Atriplex sp., D. R. Miller (1 ad. female paratype) UCD. SAN BERNARDINO Co.: 5 mi. N. Baker, IV-13-63, on Franseria sp. (Compositae), D. R. Miller (1 ad. female holotype) UCD. Host and distribution: Known on Atriplex and Franseria. 345 Probably occurs throughout warm area of southwestern United States. Acanthococcus salarius (Ferris), New ComBINATION Salt eriococcin Fig. 5 Eriococcus salarius Ferris, 1955; Hoy, 1963. Type material: | have examined the ho- lotype which is labeled “Eriococcus salarius n. sp. On Atriplex TYPE Salt Dale, near Mojave, Calif. Apr. 26, 1936 G. F. F. Stan- ford University Natural History Museum” (UCD). In addition I have seen 8 paratypes (UCD, USNM). Field features: Adult females rotund, pur- ple. No ovisac has been observed. This species is found on the roots of its host. Recognition characters: Adult females, mounted, 2.1—-3.1 (2.6) mm long, 1.5—2.6 (2.0) mm wide. Anal lobes rounded, not protruding, lightly sclerotized; each lobe dorsally with 3 enlarged setae (relative sizes variable), with 2 or 3 microtubular ducts; each lobe ventrally with 2 or 3 body setae and from 3-9 (6) sessile pores. DORSUM with enlarged setae of 2 pr- mary sizes: | larger seta on margin of each abdominal segment, also present along mar- gin of thorax and head; remaining setae small. Largest large seta 34—42 (37) u long, largest small seta 10-30 (24) uw; on abdom- inal segments VIII through III largest large seta 1.3—1.7 (1.5) times longer than longest small seta. All enlarged setae straight, fu- siform, apices rounded to blunt; setal ring broad. Enlarged setae abundant—e.g. ab- dominal segment V with 39-54 (47)—large setae showing no longitudinal pattern. Mac- rotubular ducts moderate in size, scattered over surface. Microtubular ducts moderate in length 6-7 (7) uw long, with area farthest from dermal orifice sclerotized and divided into 2 parts, apical portion rounded, from 0.5-0.8 (0.6) times length of remaining 346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Acanthococcus salarius (Ferris). 15 miles N. Kramer Junction, California, December 27, 1964, on Atriplex sp. VOLUME 93, NUMBER 2 sclerotized portion, approximately 4 times length of unsclerotized portion; dermal or- ifice sclerotized. Microtubular ducts nu- merous over surface. Anal ring normally ventral, with 4 or 5 pairs of setae. VENTER with lanceolate body setae elongate (longest seta on abdominal seg- ment VIII from 50-59 (57) u long, on seg- ment III from 53-69 (63) uw), medial setae apically acute. Enlarged setae of small type only, present along margin of abdominal segment VIII through head. Macrotubular ducts of 2 kinds: larger size present on lat- eral areas of abdomen and scattered over thorax and head; smaller size present on medial and sublateral areas of abdomen near transverse row of body setae. Microtubular ducts on lateral areas of abdomen and scat- tered over thorax and head. Multilocular sessile pores of 3 kinds: noveloculars and septeloculars scattered in small numbers over entire surface; quinqueloculars abun- dant. Cruciform pores present only in large clusters below each anterior spiracle. Legs: hind coxae dorsally with 30-55 (42) pores, ventral surface with 22-47 (38); these pores absent on femora; tibiae with 5 setae; inner, apical tibial setae unenlarged; hind tarsi longer than tibiae (hind tibia/tarsus ra- tio 0.89-0.97 (0.92); claws with small den- ticle near tip. Antennae 6- or 7-segmented, third segment longest. Apical segment with 2 or 3 sensory setae; second segment from apex with 1 slightly longer and more slender than single sensory seta on third segment from apex. Notes: This species is distinct from all other species in the United States in pos- sessing: fusiform enlarged setae and char- acteristic pattern of cruciform pores. Specimens examined: CALIFORNIA, LOS ANGELES Co.: near Mojave, Salt Dale, IV-26-36, on Atriplex sp. (Chenopo- diaceae), G. F. Ferris and P. C. Ting (1 ad. female holotype on | sl., 8 ad. female para- types, | first instar nymph on 4 sl.) UCD, USNM. SAN BERNARDINO Co.: 15 mi. 347 N. Kramer Junction, XII-28-64, on Atriplex sp., D. R. Miller (3 ad. female on 3 sl.) UCD. Host and distribution: Known only from Atriplex on the high deserts of southern Cal- ifornia. Acanthococcus tinsleyi (Cockerell), NEw COMBINATION Tinsley eriococcin Fig. 6 Eriococcus tinsleyi Cockerell, 1898, 1898a; Tinsley, 1898, Cockerell, 1899, 1900; Fernald, 1903; Cockerell, 1906; Cockerell and Robinson, 1914; Ferris, 1919, 1921; Lobdell, 1929, Ferris, 1955; Hoy, 1963; McDaniel, 1964. Nidularia tinsleyi, Lindinger, 1933. From the published records of this species the following specimens have been exam- ined and are considered misidentifications: NEW MEXICO, DONA ANA Co.: Mesilla Park, VIII-(?)-1898, on Atriplex canescens (Chenopodiaceae), T. D. A. Cockerell (see E. arenosus). TEXAS, BAILEY Co.: Mule- shoe, (?)-(?)-21, on Meriolix serrulata (On- agraceae), G. F. Ferris (see FE. eriogoni). BREWSTER Co.: Chisos Mountains, (7?)- (?)-21, on Paronychia jamesii (Caryophyl- laceae), G. F. Ferris (see E. eriogoni). PE- COS Co.: near Sheffield, Pecos River, (?)- (?)-21, on Croton sp. (Euphorbiaceae), G. F. Ferris (see E. eriogoni). Type material: | have chosen from the syntypes and marked as lectotype 1 of 2 adult females mounted on a slide labeled **Eriococcus tinsleyi Ckll., On roots of Atri- plex canescens Mesilla Park N. M. April 30 1898 (J. D. Tinsley)’ (USNM). The speci- men nearest the old label on the right side of the slide is the lectotype. In addition there are 3 lectoparatypes on | slide (USNM). Field features: Female broadly oval. Body pale brown to light purple with trace of two longitudinal purple lines dorsally; legs light brown. Covered with many crystalline rods. Ovisac yellowish-white, enclosing adult fe- male and many yellow eggs. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Acanthococcus tinsleyi (C ockerell). New Mexico, quarantined at Blythe, California, October 25, 1960, on Atriplex sp. VOLUME 93, NUMBER 2 Found on crown and roots of host. Recognition characters: Adult female, mounted, 1.3-1.4 (1.3) mm long, 0.7-0.8 (0.7) mm wide. Anal lobes apically acute, protruding, unsclerotized; each lobe dor- sally with 3 enlarged setae (lateral and pos- teromedial setae equal, anteromedial seta shortest), with 2 or 3 microtubular ducts; each lobe ventrally with 3 or 4 body setae and from | to 3 sessile pores. DORSUM with enlarged setae of | pri- mary size with 1 seta on margin of each abdominal segment slightly longer than re- maining setae. Largest seta varying from 44— 47 (45) w long, smallest seta varying from 25-28 (27) uw; on abdominal segments VIII through III largest seta from 1.6-1.7 (1.6) times longer than smallest seta. All enlarged setae slightly curved, apices rounded; setal rings thin. Enlarged setae abundant—e.g. abdominal segment V with 44-56 (50)— large type setae showing no longitudinal pattern; enlarged setae distributed in char- acteristic pattern. Macrotubular ducts scat- tered over dorsum. Microtubular ducts long (8-10 (9) w long), with area farthest from dermal orifice sclerotized and divided into 2 parts, apical portion rounded, from 0.3- 0.5 (0.5) times length of remaining sclero- tized portion; total sclerotized portion slightly longer than length of unsclerotized portion; dermal orifice unsclerotized. Mi- crotubular ducts scattered over surface. Anal ring apical or ventral, with 3 or 4 pairs of setae. VENTER with lanceolate body setae moderate in length (longest seta on abdom- inal segment VIII from 22-36 (29) uw long, on segment III from 28-41 (37) uw), medial setae apically capitate. Enlarged setae pres- ent along margin from abdominal segment VIII or VII through head. Macrotubular ducts of 2 kinds: larger size present on lat- eral areas; smaller size present on medial and sublateral areas. Microtubular ducts on lateral areas of abdomen, scattered over thorax and head. Multilocular pores of 3 kinds: septeloculars rare or absent; quin- 349 queloculars scattered over entire surface; triloculars most numerous on thorax. Cru- ciform pores present on lateral areas from anterior abdominal segments forward to head, a few such pores anterior of meso- thoracic pair of legs. Legs: hind coxae dorsally with 9-25 (12) pores, ventral surface with 5-10 (7); hind femora dorsally with 4-6 (5) pores, absent ventrally; tibiae with 4 setae; inner, apical, tibial setae slightly larger than remaining leg setae; hind tarsi longer than tibiae (hind tib- ia/tarsus ratio 0.78—0.83 (0.80)); claws with conspicuous denticle near tip. Antennae 6- or 7-segmented, when 7-segmented, seg- ment 4 longest. Apical segment with 3 sen- sory setae; second segment from apex with 1 longer and more slender than single sen- sory seta on third segment from apex. Notes: This species is quite similar to A. eriogoni (Ehrhorn), but differs in possessing: characteristic dorsal setal pattern, different- ly shaped microtubular ducts and 4 setae on each tibia; A. eriogoni, on the other hand, possesses: 5 setae on each tibia. Specimens examined: ARIZONA, APACHE Co: Springerville, VII-4-18, on Atriplex canescens (Chenopodiaceae), G. F. Ferris (2 ad. female) UCD. CALIFORNIA, RIVERSIDE Co: 11 mi. N. Indio, VIII-24-68, on Atriplex sp., D. R. Miller (3 ad. female) UCD. IDAHO, POWER Co.: 5 mi. S. American Falls, VIII-19-64, on Chrysothamnus sp. (Compositae), D. R. and J. F. Miller (4 ad. female on 2 sl.) UCD. NEW MEXICO, quarantined at Blythe, California, X-25-60, on Atriplex sp., D. R. Dilley (3 ad. female on 2 sl.) CDA. DONA ANA Co.: Mesilla Park, X-9-1896, on Atri- plex sp., Townsend (3 ad. female) USNM; IV-30-1898, on A. canescens, J. D. Tinsley (lectotype, 3 paralectotypes on 2 sl.) USNM; X-13-1898, on A. canescens, J. D. Tinsley (20 spm. on 2 sl.) USNM. TEXAS, EL PASO Co.: near Fabens, VIII- (?)-21, on Atriplex sp., G. F. Ferris (1 ad. female) UCD. 350 Host and distribution: Known on Atriplex and Chrysothamnus. Probably occurring throughout the warm areas of the western United States. Acanthococcus whiteheadi Miller, NEw SPECIES Whitehead eriococcin Figs 7 Type material: Adult female holotype (1 specimen on slide) with left label ‘“‘Eriococ- cus whiteheadi Miller TYPE”; right label “8 mi. E. Hawthorne, Mineral Co., NE- VADA 6-VII-1968 ex Atriplex sp. D. R. Miller & R. F. Denno 1212” (UCD). In ad- dition there is a single paratype. Field features: This is a very unusual erio- coccin. In appearance it resembles a fuzzy seed. The body is dark green and 1s covered ventrally by a light white secretion. Many slender, waxy, crystalline rods are produced from the dorsum giving the body a seed- like appearance. This species is found on the main roots of its host. Recognition characters: Adult female ho- lotype, mounted, 2.0 mm long, 1.2 mm wide (paratype 2.0 mm long, 1.1 mm wide). Anal lobes strongly protruding, acute, slightly sclerotized; each lobe dorsally with 3 en- larged setae (lateral and posteromedial setae approximately equal, anteromedial seta shortest and most slender), with 3 micro- tubular ducts; each lobe ventrally with 4 body setae and 4 or 5 sessile pores. DORSUM with enlarged setae of 2 sizes: larger setae present medially, sublaterally, and laterally, from abdominal segment VIII through thorax, with 2—4 (3) such setae in medial cluster in each abdominal segment, 1 or 2 on sublateral area, and 2 or 3 on each margin; remaining setae of small size. Larg- est large seta 50 u long (paratype 58 wu), larg- est small seta 37 uw long (paratype 41 yu); longest large seta 1.3 times longer than lon- gest small seta (paratype 1.4 times). All se- tae straight, conspicuously broad basally, with acute apices; setal rings thin, often ap- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pearing fused to remainder of seta. Enlarged setae abundant—e.g. abdominal segment V with 59 (paratype with 69)—large setae forming 5 longitudinal lines (1 medially, 2 sublaterally, and 2 laterally). Macrotubular ducts in small numbers over surface. Mi- crotubular ducts elongate (12 u long) (para- type 14 uw), with area farthest from dermal orifice sclerotized and weakly divided into 2 parts, apical portion rounded apically, ap- proximately 0.3 times length of remaining sclerotized portion; total sclerotized area approximately 0.3 times length of unscler- otized portion; dermal ring weakly sclero- tized, thin, although similar to bifurcate or- ifice of other species in the genus such as A. azaleae, apparently with single opening only. Microtubular ducts in small numbers over surface. Anal ring bent around abdominal apex, with 4 setae. VENTER with lanceolate body setae moderate to elongate (longest seta on ab- dominal segment VIII 50 yu long (paratype 62 «), on segment III 51 u (paratype 63 y)), medial setae with capitate apices. Enlarged setae present along lateral margin from ab- dominal segment VIII through head. Mac- rotubular ducts of 2 kinds: larger size re- stricted to lateral areas; smaller size present on medial and sublateral areas near body setae. Microtubular ducts most abundant along lateral margins, also present in small numbers on medial and sublateral areas of thorax and head. Multilocular sessile pores in unusually small numbers on thorax; ses- sile pores of 2 kinds: quinqueloculars pres- ent over entire surface, most abundant on posterior abdominal segments; triloculars present near spiracles and on anterior ab- dominal segments. Cruciform pores present on sublateral areas of anterior abdominal segments, and on thorax, and head. Legs: hind coxae dorsally with 22 and 26 pores (paratype with 21 and 26), ventrally with 18 and 23 (paratype with 31 and 23); hind femora dorsally with 2 and | pores (paratype with 3 and 5), ventrally with 3 VOLUME 93, NUMBER 2 351 , ed — a a-4 ar \ AVY 7 LN | fo) / Ke Ne ‘ i / / > SN Md wrens bade Nee " 2] “XS vo yibes = \z \ wow ie : aye Wee ida i a2 vss Rif Ie LE ATTY _, \ eae wet eg oat — or a “p Vir “=. ppp Pill : Wis 49 Zh: ‘ eS ee = o—®) ao o ~) 2 2 44 sg igs ° < a a 45 al A plein 4 el! pe -. ‘22> “ay | a) —2_ 2 Fy LAC Al sp Aa og PPA | = Bia Py on) a Pe , 2 Aaa ne ° os 4 A 4 pp * D° ww E A Whe pA fl | ral le OF maa oe I (ane ee git ti 44 ZPOV EG |e) | ‘ BD ‘ae agp yt i . yy mG? : \ a3 ae I as ’ vey. (paw * y ea sags : eee ; ov Se ee — i \ a y \ar See Fig. 7. Acanthococcus whiteheadi Miller, n. sp. 8 miles E. Hawthorne, Nevada, July 6, 1968, on Atriplex sp. 352 and 2 (paratype with 2 and 3); tibiae with 5 setae; inner, apical, tibial setae robust; tar- si slightly longer than tibiae (hind tibia/tar- sus ratio 0.87) (paratype 0.90); claws with small denticle near tip. Antennae 7-seg- mented, third segment longest. Segment 7 with 3 sensory setae; segment 6 with | lon- ger and more slender than single sensory seta on segment 5. Variation: The paratype agrees with the holotype in nearly all respects except it pos- sesses one 6-segmented antenna. Notes: I take great pleasure in naming this species A. whiteheadi in honor of the late Donald R. Whitehead, Systematic Ento- mology Laboratory, ARS, USDA. He was a good friend who had significant impact on my career. See appendix I for a eulogy that I presented during the memorial service in his honor. The unusual elongate microtubular ducts are more similar to the microtubulars on species from New Zealand or Australia than to the microtubulars cf North American species. Specimens examined: NEVADA, MIN- ERAL Co.: 8 mi. E. Hawthorne, VII-6-68, on Atriplex sp. (Chenopodiacae), D. R. Mil- ler and R. F. Denno (1 ad. female holotype, 1 ad. female paratype on 2 sl.) UCD. Host and distribution: Probably occurring throughout warm areas of the southwestern United States on Atriplex. CONCLUSIONS There are seven species of Acanthococcus that feed on Atriplex in western North America. This host has more species of Acanthococcus than any other plant genus in the region. The genus Eriogonum has five species and Artemisia, Gutierrezia, and Eu- phorbia each have four (Miller 1969). It is interesting to note that of the seven Acan- thococcus species that occur on Atriplex, five are restricted to this host or are found on one other, probably incidental, genus and only two occur on four or more host genera. Of the species that occur on Eriogonum, only one Acanthococcus species is restricted PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to Eriogonum and four occur on four or more host genera. For Artemisia there are two species restricted to this host and two that are polyphagous. For Gutierrezia and Euphorbia all four Acanthococcus species are polyphagous. It appears that some interesting evolu- tionary trends have occurred in the Acan- thococcus species that inhabit Atriplex and perhaps Artemisia. The relatively host-spe- cific species that occur on these hosts are quite distinctive and are readily recognized even without a key. The significance of this phenomenon may become apparent when phylogenetic relationships among Acantho- coccus species are analyzed. LITERATURE CITED Borchsenius, N.S. 1948. On the revision of the genus Eriococcus Sig. Akademia Naukowe Doklady 60: 501-503. (In Russian.) Cockerell, T. D. A. 1897. New insects from Embudo, New Mexico. Annals of the Magazine of Natural History 20: 510-515. 1898. Supplementary note, and notice of a new Eriococcus. Canadian Entomologist 30: 246— 247. 1898a. A bright new parasite of Coccidae. Canadian Entomologist 30: 276. 1899. Article VII.—First supplement to the check-list of the Coccidae. Bulletin of the Illinois State Laboratory of Natural History 5: 389-398. 1900. A new Eriococcus, with remarks on other species. Entomological News 11: 594-596. 1906. The South American Coccidae of the genus Eriococcus. Proceedings of the Entomolog- ical Society of Washington 8: 32. Cockerell, T. D. A. and E. Robinson. 1914. Descrip- tions and records of Coccidae. American Museum Natural History 33: 327-335. Ehrhorn, E. M. 1911. New Coccidae, with notes on other species. Canadian Entomologist. 43: 275- 280. Fernald, M. E. 1903. A catalog of the Coccidae of the world. Massachusetts Agricultural Experiment Station Bulletin 88, 360 pp. Ferris, G. F. 1919. A contribution to the knowledge of the Coccidae of southwestern United States. Stanford University Publications, University Se- ries, Biological Sciences, 68 pp. . 1920. Scale insects of Santa Cruz— Peninsula. Stanford University Publications, University Se- ries, Biological Sciences, 60 pp. 1921. Report upon a collection of Coccidae from Lower California. Stanford University Pub- VOLUME 93, NUMBER 2 lications, University Series, Biological Sciences |: 61-132. 1955. Atlas of the scale insects of North America. Volume VII. The families Aclerdidae, Asterolecaniidae, Conchaspididae, Dactylopiidae, and Lacciferidae. Stanford University Press, Stan- ford, California, 233 pp. Hoy, J. M. 1963. A catalogue of the Eriococcidae of the world. New Zealand Department of Science and Industrial Research Bulletin 50, 260 pp. Lindinger, L. 1933. Beitraége zur Kenntnis dir Schild- lause. Entomologische Anzeiger 13: 77-78, 107- 108, 116-117, 143, 159-160, 165-166. Lobdell, G. H. 1929. Two new species of Eriococcus from Mississippi. Annals of the Entomological So- ciety of America 22: 762-767. McDaniel, B. 1959. An undescribed Eriococcus from Mexico. Proceedings of the Entomological Society of Washington 61: 137-138. . 1964. Key to Texas species of Eriococcus and a description of a new species. Texas Journal of Science 15: 111-114. Miller, D. R. 1969. A systematic study of Eriococcus Targioni-Tozzetti with a discussion of the zoo- geography of the Eriococcidae. Ph.D. Dissertation submitted to the Department of Entomology, Uni- versity of California, Davis, 253 pp. 1984. Terminology. Scale 10: 47-49. Miller, D. R. and Gonzales, R. H. 1975. A taxonomic analysis of the Eriococcidae of Chile. Revista Chi- leana Entomologia 9: 131-163. Miller, D. R. and McKenzie, H. L. 1967. A system- atic study of Ovaticoccus Kloet and its relatives, with a key to North American genera of Eriococ- cidae. Hilgardia 38: 471-537. Nelson, C. R., Haws, B. A., and Nelson, D. L. (in press). Incidence and distribution of mealybugs and related Homoptera of shadscale: Possible agents in the dieoff problems in the intermountain west. Sasscer, E. R. 1912. Catalogue of recently described Coccidae—IV. United States Department of Ag- riculture Bureau of Entomology, Technical Series 16: 83-97. Tinsley, J. D. 1898. Notes on Coccidae, with de- scriptions of new species. Canadian Entomologist 30: 317-320. APPENDIX I A Eulogy presented May 7, 1990 at the Memorial Service held at the Grace Presbyterian Church in Lanham, Maryland Donald R. Whitehead was a unique char- acter—I mean that as a double positive. He was a character among characters, for I am 353 told that systematic entomologists have a reputation for being somewhat unusual hu- man beings. In the next several minutes | would like to describe why I characterize Donald as special and why he had an im- portant influence on my life and career even during the last days of his life. Please forgive me if I take too long in this task, but this will be the last time that we will take the opportunity to think and talk about him for any extensive length of time. I think he de- serves our thoughts. Donald had a diverse combination of in- terests and personality traits and I certainly was not privy to them all. The words that describe him by my perception are: intel- lectual, unswerving, willpower, dedicated, researcher, insect identifier, curator, mil- lipeds, stamp collector, computer, self dep- recating, tropics, bowler, ground beetles, idea person, zoogeographer, seed weevils, wee- vils, West Virginia, Costa Rica, Mexico, quiet, bald, immigrant, colleague, friend. Donald was first and foremost an intel- lectual, and, by my perception, he was quite bright. His more than 60 scientific papers reflect his probing interest in finding a pat- tern or explanation for generally accepted phenomena. For example, he was keenly interested in pests and later became es- pecially curious about immigrant pests. He searched for characteristics that were shared by pest species of weevils so that he could predict which species were most likely to become pests in the future. He also inves- tigated how to determine if a pest was an immigrant or naturally occurred in the U.S. Until Donald started asking these ques- tions, I, at least, would never have consid- ered questioning such basic terms as pest and immigrant. Donald could make almost anything an intellectual endeavor. At one point, he was asked to write a position paper on the mission of the Systematic Entomol- ogy Laboratory for presentation at a retreat. If I had been given this assignment, I would probably have copied an earlier mission statement, modified it to a limited extent, and been done with it. But not Donald! He 354 started completely fresh, talked with others in the Laboratory and produced so much interest that the mission generated more discussion than any other subject at the re- treat. The point is that Don, by his ques- tioning intellect, stimulated thought about a subject that was basic to the entire un- derpinnings of the Laboratory but never was given serious thought or consideration. He was an idea person. His contributions were many and diverse and he had an unswerving dedication, even self-depricating devotion, to the areas of his vocation. In research, he frequently became frustrated with his inability to find the per- fect answer to a problem. He had very high expectations of himself, and, when he didn’t achieve the unachievable, he would wait and gather more data rather than publish his already very significant contributions. He expected to develop a new theory of rela- tivity, and, when he did not, he treated his findings as insignificant and unimportant. His colleagues tried to make him under- stand the importance of his discoveries, but because he expected more, he usually didn’t accept our opinions. The research that he did publish is quite important, but I hate to think of the many discoveries that must be rediscovered in the future. His research contributions and ideas were far from his only contributions at work. In 1985 he was given an award for outstanding achievement in providing insect identifi- cation services and information. If a U.S. port of entry, a homeowner, or a researcher required information and the name of a beetle of concern, Donald would spend hours or even days trying to find the answer. The information that he provided was always extensive and well researched and was far beyond what was expected. His curatorial contributions to the Smithsonian’s Nation- al Insect Collection have been unheralded to now, but are deserving of special recog- nition. As Donald’s supervisor, I have al- ways been impressed during his yearly eval- uation with the massive portions of the weevil collection that he had reorganized or PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON curated. Most scientists in the Laboratory have expressed their curatorial accomplish- ments in terms of drawers. while Donald only thought in terms of cabinets. With no assistance other than his own hands, he made large portions of the weevil collection available and usable for future research and scientific discovery. For me, Donald had an important influ- ence early in our relationship as a knowl- edgeable colleague, even teacher. I was a staunch believer in the old approach to sys- tematics and was taught that cladistic ap- proaches were useless. Donald, Chris Thompson, and I had lengthy, vigorous dis- cussions about the subject. I can still re- member Donald’s animated gesticulations in his pursuit of making a point. What fun we had in learning from one another, even though to our wives it seemed that we were going to end up hating or even punching each other. It was during these sessions that Donald was given the nickname Baldhead. Several months after putting this label on him, I had second thoughts, since some are self conscious about such attributes. I should have known better, since Donald seemed to pride himself in his polished dome and he often referred to himself as Baldy when he sent memos and mail messages. This man had a willpower much stronger than most. For many years, Donald enjoyed the vices of smoking and drinking, but as time passed they became more and more of a problem. So as was typical of him, he simply decided to quit. Not only did he de- cide to quit both at the same time, against all advice, but he also decided to quit when it is most dificult—at Christmas and New Years—and of course, to tempt himself even more, by bringing a bottle of scotch whiskey as his contribution to the Christmas party downtown. As far as I know, Donald never wavered from his decision to quit. He never smoked or drank again. Donald also was intense. When he decid- ed to do something, he did it all the way even if the activity was recreational. At some point, he joined us in bowling duck pins one VOLUME 93, NUMBER 2 evening a week. I don’t think he even really wanted to bowl, but once he got started, he went at it with a vengeance. For most of us, bowling provided a change of pace one night each week. If we did well, that was fine, but if we didn’t, it wasn’t a concern. As often was the case, Donald wasn’t satisfied with doing well. Each time he bowled he ex- pected to do excellently; if he didn’t he was upset with himself. In an attempt to over- come this problem, he decided to practice and I think he may have practiced as often as twice a week. In the scientific arena, he had a broad array of interests from patterns of distri- bution as they translate to species recogni- tion to revisionary studies of acorn weevils. He studied ground beetles, seed weevils, weevils, millipeds, parasitic wasps, true bugs and was very much enamored by the trop- ics, especially in Mexico and Costa Rica. He enjoyed doing field work and derived much pleasure from his work with George Ball, Dan Janzen, and John Kingsolver. The sounds, smells, and feel of the tropics had a special importance to Donald. But another area closer to home also was a favorite, namely West Virginia. The milliped fauna of West Virginia is especially diverse, and in the past several years he formulated a hypothesis of milliped mimicry in West Virginia. I thought it especially fitting that the presentation that I am giving here was written in his beloved West Virginia. 355 Even to his avocation, Donald brought an unusual amount of vigor and intensity. He was an expert on the post marks of Mex- ico during the period from 1870 to 1915 and made major contributions to a book on the subject. He also had special collections of post marks from Tasmania and recently started a West Virginia collection. When I asked him about his collection two weeks ago, he launched into a discussion that was well beyond his energy level. If he had been able, his arms would have been gesticulating as only Donald could make them, in his excitement over a subject that was dear to him. Up to the very end, he had Jo bidding on collections in mail auctions. During the early morning hours when Donald couldn’t sleep because of chemotherapy treatment, he was working away on a database on Mex- ican post marks. He told me that it was of no value and not worth looking at, but I suspect otherwise. In the end, Donald would not give up. He beat the odds for the last year of his life, a year that seemed especially important to him. In that year, he allowed himself some of the pleasures that his dedication would not allow previously. I am told that he pur- chased especially extravagant gifts for those he loved at Christmas. He allowed himself the time to visit with colleagues and he spent a lot of time in West Virginia studying mil- lipeds. I personally received a lot from old Baldy, and I will miss him greatly. PROC. ENTOMOL. SOC. WASH. 93(2), 1991, pp. 356-389 THE TRIBE CICINDINI BANNINGER (COLEOPTERA: CARABIDAE): COMPARATIVE MORPHOLOGY, CLASSIFICATION, NATURAL HISTORY, AND EVOLUTION DAvip H. KAVANAUGH AND TERRY L. ERWIN (DHK) California Academy of Sciences, San Francisco, California 94118; (TLE) Na- tional Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Abstract. —The two species of the Gondwanian carabid beetle tribe, Cicindini, are re- described in detail and their relationships within the Carabidae and among themselves are reassessed. On the basis of several autapotypic features, a new genus, Archaeocindis, is erected for the Persian Gulf species, Cicindis johnbeckeri Banninger. The type locality for Cicindis horni Bruch is restricted to Guanaco Muerto, Cordoba Province, Argentina. For each taxon, a synthesis of available literature on both taxonomy and natural history is presented. The name of the tribe is determined to be Cicindini, not Cicindisini, according to the rules of zoological nomenclature. Based on a detailed analysis of characters of external structure and genitalia and consideration of past and present classifications of carabidae, cicindines are placed in a separate supertribe, the Cicinditae, near the Nebriitae and Elaphritae (sensu Kryzhanovskiy 1976). Present geographical distributions of the tribe and genera suggest that cicindines are a western Gondwanian lineage, the distribution of which was divided by development of the South Atlantic Basin in the Late Mesozoic. Resulting South American and African isolates gave rise to Cicindis and Archaeocindis, respectively. Members of the former taxon occupy interior saline lake shore habitats, those of the latter occur in tidal flats of bays in the Persian Gulf. Based on interpretation of a suite of unusual structural features, cicindines probably behave like diurnal tiger beetles that also can survive submersion in and swim on the surface of salt water. Key Words: Cicindini, Cicindis, Archaeocindis, Gondwanian lineages, carabid classifi- cation In 1979, one of us (TLE) wrote that spe- cies of the tribe Cicindini “‘surely represent the most obscure group of carabid beetles remaining today. Their bizarre features in combination with lack of available material make them nearly impossible to deal with effectively.’’ Eleven years later, we reject the latter, rather defeatist notion, and take up the challenge to better understand this ob- scure group, even though only one more specimen 1s now available to aid in the study. Luckily, this specimen came with important new biological information that should en- able us to gather additional specimens and data in the future. It is with this prospect in mind that we provide a synthesis of cur- rent knowledge and add new information on the structure, distribution, and phylo- genetic relationships of these beetles, based on our study of available specimens. Bruch (1908) recognized his new species as something peculiar and placed it near the tiger beetles. He did so without formally classifying them, probably because he was not a carabid specialist. Banninger (1925, 1927a, b) studied the Argentine specimen VOLUME 93, NUMBER 2 described by Bruch and another specimen collected at Baishehr, Iran, which he de- scribed as new, concluding that the genus was related to the basal lineages of carabids, as understood in the early part of this cen- tury. He classified the lineage near the Ozae- nini, with which he was very familiar (Ban- ninger 1927b), as an independent group within the Isochaeta. He mentioned in his 1925 paper that the lineage should be given tribal status, but did not provide a name; subsequently, he did so in his 1927a paper. For several decades, Banninger’s classifi- cation was either accepted or not dealt with (for example, see Csiki 1927, Crowson 1955, Blackwelder 1944, Kryzhanovskiy 1976). Reichardt (1977) elevated the group’s status to the rank of subfamily, but left it in a position near the Paussinae (which contains the Ozaenini). Ball (1979) and Erwin (1979), without recourse to the specimens, also tac- itly accepted this classification, but later Ball and McCleve (1990) removed the lineage from the paussines assemblage, leaving its status, in their opinion, as incertae sedis. However, after studying one of the speci- mens, Erwin (1985: 467) concluded that the group, at the rank of tribe, belonged to the Nebriitae lineage of the Carabinae, near the tribe Notiophilini. Reichardt (1977) reported that specimens of Cicindis horni Bruch had been collected by A. Martinez near salt lakes in desertic areas. We were informed that the specimen collected in 1979 by R. A. Ronderos (pers. comm.) listed below was found at a light, at night, near the enormous salt lake bed of Salinas Grandes in the northern part of Cor- doba Province, Argentina. Stork (1982) re- ported on a specimen of Cicindis johnbeck- eri Banninger received for identification. It had been collected as part of a study of ““mud skippers”’ in the Persian Gulf, taken from a tidal flat, probably esturine, in a bay some 20 km north of Kuwait City. The type spec- imen of this species, collected almost di- rectly across the Persian Gulf from Kuwait, at Bashehr, Iran, also may have been col- 357 lected from a similar tidal flat, perhaps in the Bay of Soltani (Khowr-e Soltani). Limits of the present study were deter- mined by the paucity and condition of available specimens. One species is known only from females and both species are known to us from only two specimens each (additional specimens of C. horni are re- portedly in the UNLP or MACN but have not been made available to us). One of the specimens studied is in poor condition with most appendages missing, a second speci- men has been damaged by previous dissec- tions, and a third is teneral. Thus, complete disarticulation of one or more specimens, to properly study all structural features, was necessarily ruled out. Nonetheless, we feel that our review of the form and structure of cicindines, through conservative dissec- tion, the results of which are presented here, provides new information that better rep- resents structural diversity within the group and relationships of its members to other lineages of Carabidae. MATERIALS AND METHODS General procedural methods are those which we have used before (Erwin and Ka- vanaugh 1981). Measures for body length, and pronotal and elytral dimensions are coded as follows and are presented in the species descriptions as ranges based on the smaller and larger of specimens studied. All specimens were measured with an ocular micrometer in a Wild microscope and mea- sures are presented in millimeters. SBL, standardized body length = the sum of head length (midline distance from apical margin of clypeus to a point opposite posterior mar- gin of eyes) + pronotal length (midline dis- tance from anterior [apical] to posterior [basal] margin) + elytral length (midline distance from apex of scutellum to a point opposite apex of longer elytron) (see Ka- vanaugh 1979, Erwin and Kavanaugh, 1981). TW, total width across the widest portion of the elytra = width of left elytron, measured at widest point, and doubled to 358 obtain value. We use the term forebody to indicate the head and pronotum together. Accept where noted, scale lines in all figures are equal to 1.0 mm length. For conciseness, we have grouped the characters and their states in three catego- ries, depending on their perceived utility in outgroup comparisons with equivalent rank taxa. For example, if the state of some char- acter is of general importance at the tribal level we place it in the tribal diagnosis, whereas minor characteristics such as color are placed within the species descriptions, because these have value only among spe- cies level groups. We do not repeat descrip- tors at successive higher or lower taxon lev- els. Specimens were borrowed from the fol- lowing institutions for this study: BMNH Department of Entomology, Brit- ish Museum (Natural History), Cromwell Road, London, En- gland Institut fiir Pflanzenschutzfor- schung (BZA) der Akademie der Landwirtschaftswissenschaften der Deutschen Demokratischen Republik, DDR 13, Eberswalde - Finow | Museo Argentina de Ciencias Na- turales, Av. Angel Gallerdo 470, Buenos Aires, Argentina Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Paseo del Bosque 1900, La Plata, Argentina DEIE MACN UNLP Tribe Cicindini Banninger Cicindisini Banninger 1927a: 119. Cicindisini Banninger 1927b: 177. Cicindini Csiki 1927: 425. Cicindini Blackwelder 1944: 22. Cicindini Crowson 1955: 6. Cicindisini Kryzhanovskiy 1976: 56. Cicindisinae Reichardt 1977: 375. Cicindisini Reichardt 1977: 357, 1979: 319, 32): PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cicindisini Ball 1979: 91, 95, 100. Cicindisini Erwin 1979: 589, 1985:467. Tribal nomenclature. — Banninger (1925) first proposed that the genus Cicindis be classified in its own tribe, but did not men- tion a name for such until later, where he used Cicindisini (Banninger 1927a). In the same year, Csiki (1927), citing Banninger’s 1925 paper, listed Cicindis under the tribal name Cicindini. Both spellings have been used since, and repeatedly, in the literature. Because Banninger’s paper was published on November 10 and Csiki’s on December 22, 1927, Banninger’s use of a tribal name, Cicindisini, has priority. However, Bannin- ger applied the tribal ending, -ini, to the complete generic name rather than its stem, whereas Csiki’s name, Cicindini, is formed correctly. Based on Article 11f (ii) of the International Code of Zoological Nomen- clature, the valid tribal name is Cicindini, and Banninger is its author. Tribal diagnosis.—Adults: Slightly hy- pognathous. Ventral surface of body (except for gular region of head) and coxae, tro- chanters, and femora covered with fine pu- bescence of slightly to moderately decum- bent and curved setae. Head without or with one pair of supraorbital setiferous punc- tures; frons without longitudinal ridges; ver- tex slightly concave; gena with a sharp-edged longitudinal flange ventral to eye, postgenal groove present, distinctly biarcuate; com- pound eye very large, longitudinal diameter equal to or more than 1.5 times width of labrum, dorsomedial margin markedly con- cave; clypeus narrower than distance be- tween antennal sockets; mandible (Figs. 6, 7) with terebral blade long and markedly down-curved, scrobe asetose and delimited dorsomedially by a prominent elevated ridge extended apically onto blade, terebral tooth triangular, retinaculum with both anterior and posterior retinacular teeth (the former larger and more acuminate on right man- dible than left), small accessory tooth (ho- mology unknown) on each mandible be- VOLUME 93, NUMBER 2 tween terebral and posterior retinacular teeth, molar region undeveloped, setiferous ventral groove present, longer on left than on right mandible; maxilla, mentum, and submentum without setiferous spiniform processes or ridges; lacinia of maxilla (Fig. 8) with apical tooth articulated; ligula of labium (Fig. 10) with paraglossae short but distinct. Prothorax with front coxal cavities bridged (biperforate) (Fig. 14A) and sepa- rate internally (Fig. 14B), closed posteriorly by narrow medial extension of propleuron with prosternal projection overlapped pos- teriorly; front tibia (Fig. 19) with apex only slightly oblique (posterior angle slightly dis- placed proximally) and both spurs apical, anteroventral margin with row of short fine densely arranged setae along apical two- thirds, setae progressively longer apically, antenna cleaner simple, sulcate, with two long, curled setae proximal to posterior api- cal spur. Pterothorax with elytron (Fig. 15) with nine complete, shallowly impressed in- terneurs, scutellar interneur short, present only on basal one-seventh; interval 2 much narrower than intervals 3 and 4 together; discal setae absent or present only on in- terval 3, umbilicate series present on inter- val 9, with 12 to 14 setae positioned as in Fig. 15); internal plica simple, keel-like, faintly defined (especially near apex), not extended to epipleuron apically; metatho- racic wing (Fig. 16) with Oblongum Cell narrowed posteriorly, stalked, M4 vein po- sitioned slightly anterior to middle of Ob- longum Cell, Third Radial Cell larger than Anterior Sector Cell; metepimeron (Fig. 17) present, triangular; middle coxal cavities disjunct, confluent (through very small hole); hind coxal cavities conjunct, confluent, de- limited laterally by metepimera and second (first visible) abdominal sternum; hind tar- someres (Fig. 25) 1 to 4 with ventroapical margin lobate anteriorly, lobes and associ- ated setae longer on successive tarsomeres; tarsal claws (Fig. 23) asymmetric in length, the anterior claw longer. Female genitalia with eighth sternum (Fig. 28) and ninth/ 359 tenth tergum (Fig. 29) undivided; gonan- gulum (Fig. 30) with scattered setae dor- somedially; coxostylus (Fig. 31) short, stout, with three or four spines lateroventrally and two or three spines dorsomedially. Larvae: Unknown. Tribal distribution. — The widely disjunct distribution (Fig. 33) includes two rather circumscribed areas along the bays of the northern Persian Gulf and among the lakes of the great salt flats of northwestern Ar- gentina. KEY TO SPECIES 1. Anterior pair of supraorbital setigerous punc- tures present; elytral margin serrate, more so apically .. Archaeocindis johnbeckeri (Banninger) 1’. Supraorbital setigerous punctures absent; ely- tral margin smooth ....... Cicindis horni Bruch Archaeocindis Kavanaugh and Erwin, NEw GENUS Type species. — Cicindis johnbeckeri Ban- ninger 1927a: 119. Derivation of genus name.—From the Greek, archaeon, meaning ancient, and a part of the genus name, Cicindis, referring to the occurrence of this genus in the Pa- laearctic Region, particularly near the mouth of the historically important Tigris and Eu- phrates Rivers. Diagnosis.—Body deep, subcylindrical. Head (Fig. 3A) with one pair of supraorbital setiferous punctures near medial margin of eye (anterior one-sixth); area between pos- terior margin of eye and postgenal groove slightly depressed; antennomeres | to 3 and basal four-fifths of 4 glabrous (except for apical whorl of fixed setae), apical one-fifth of antennomere 4 with a few, small setae, antennomeres 5 to 11 densely and evenly pubescent (Fig. 4A); dorsal surface of man- dible smooth, without macrosculpture; last (fourth) maxillary palpomere (Fig. 8) with a sensory pit ventromedially in apical one- half; ligula (Fig. 10A) deeply emarginate apically, paraglossae, short, much shorter than fused glossae; mentum (Fig. 11A) with anteromedial emargination deep, dentate, 360 tooth broadly entire and with one pair of paramedial setiferous punctures at base, epilobes narrowly rounded, broadly toothed anteromedially and without setae; sub- mentum broad anteriorly, only slightly nar- rower than mentum, with one pair of lateral setiferous punctures, gular portion with sides subparallel. Pterothoracic elytron-locking mechanism (Fig. 18A) with deeply emar- ginate internal and short external meta- pleural flanges, external flange with poste- rior bulge inserted in elytral epipleural cavity (when elytra are closed), elytral epipleuron markedly notched. Femora longitudinally concave, front femoral and hind femoral concavities shallow anteriorly and deep posteriorly, middle femoral concavities deep both anteriorly and posteriorly. Middle and hind tibiae (Figs. 20A, 21A) without acces- sory setae; hind tibia straight; tibial spurs of anterior tibia longer than tarsomere 1, posterior spur of middle tibia and anterior spur of hind tibia subequal to length of re- spective first tarsomeres. Tarsi (Figs. 22A, 24A, 25A) without fringes of accessory se- tae; posterior claw (Fig. 23A) of all tarsi at least two-thirds the length of anterior claw. First visible (second) abdominal sternum without lateral concavity. Geographic distribution.—Known only from both sides of the north end of the Per- sian Gulf. Archaeocindis johnbeckeri (Banninger), New ComMBINATION Cicindis johnbeckeri Banninger 1927: 119. Holotype female, in DEIE, labelled: ‘““Holotypus” [red label]/ ““Bushere Beck- er 22.1V.27”/ “Type [written vertical] Ci- cindis John-Beckeri m.” [‘‘8.1927” on underside]. Type locality: IRAN, Bish- ehr [28°59’N 50°50’E]. Diagnosis.—Clypeus narrower than dis- tance between antennal sockets, anterior su- praorbital setiferous punctures present, and pronotal and elytral margins finely serrate. Description of form and structure of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON adults. —Size medium, standard body length of females 9.6 and 10.2 mm. Form (Fig. 1) with elytra proportionately short in relation to forebody (ratio: elytral length/forebody length = 1.63 [mean]). Color: Head, body, and appendages pale yellowish tan, except eyes piceous and api- ces and medial margins of mandibles in- fuscated; elytra with very pale marginal pat- tern (Fig. 15A). Luster: Entire body surface moderately shiny, except elytra slightly duller. Microsculpture: Entire body with isodia- metric meshes, well impressed on labrum, legs, and elytra, more faintly impressed on forebody and venter. Head: Smooth dorsally; frons slightly convex, frontal furrows absent; antenna (Fig. 4A) with scape long, length about equal to width of labrum, markedly narrowed ba- sally, and form slightly sinuate; clypeus (Fig. 5A) with anterior margin straight; penulti- mate labial palpomere (Fig. 10A) with three setae anteriorly. Prothorax: Pronotum (Fig. 12A) broad, greatest width wider than width of head across eyes, slightly cordate, slightly nar- rowed basally, smooth medially, without obliquely transverse and shallow rugulae laterally; disc markedly convex; apical mar- gin markedly bisinuate; lateral margin cren- ulate or faintly serrate and arcuate, with bas- al sinuation shallow and long; lateral explanation narrowed medially, broadened apically and basally; basal margin markedly bisinuate; apical angles long, narrow, mark- edly projected anteriorly; basal angles ob- tuse and sinuate, slightly projected poste- riorly; apical margination deeply impressed laterally, absent from middle one-sixth; lat- eral bead narrow, poorly defined anteriorly, effaced near apical angle, elevated and well defined at basolateral setiferous puncture, absent from sinuation of basal angle; basal margination effaced medially and laterally, faintly defined paralaterally; anterior trans- verse impression broadly and shallowly de- fined; median longitudinal impression nar- VOLUME 93, NUMBER 2 LY as 361 Seas a= \ (7) » 2 y Z Fig. 1. Habitus, dorsal aspect, Archaeocindis johnbeckeri (Banninger). rowly and finely defined; posterior transverse impression smooth to slightly rugulose, nar- rowly and deeply defined, extended laterally to basal foveae, distinctly arcuate; basal fo- veae deep anteriorly, posteriorly moderate- ly deep, slightly divergent, and confluent with lateral explanation; mid-lateral setif- erous puncture absent, basolateral setifer- 362 ous puncture inserted on posterior end of elevated lateral margination distinctly an- terior to basal angle. Prosternal intercoxal process (Fig. 13) broadly spatulate in ven- tral aspect. Pterothorax: Elytra (Fig. 15A) with sil- houette subfusiform, widest anterior to middle, lateral margins faintly and apical margins distinctly serrate; elytron convex, with basal margination complete, deeply impressed and markedly sinuate, lateral margination complete and very narrow, slightly broadened near middle, effaced from apex, humerus rounded, lateral and basal margination joined at markedly obtuse an- gle, with humeral angle produced anteriorly, interneurs | to 7 finely striate, 8 and 9 very faintly punctulate, intervals smooth, im- punctate, slightly convex or flat; metatho- racic wing (Fig. 16A) full-sized, with stalk of Oblongum Cell short. Legs: All legs long and slender; front tar- someres | to 4 slender and without pads of adhesive setae ventrally in female (male un- known). Abdomen: Last visible (seventh) sternum moderately emarginate apicomedially and with two pairs of long paramedial marginal setae in female (male unknown). Male genitalia: Male unknown. Female genitalia: Coxostylus (Fig. 31A) slightly rounded apically, with four ventro- lateral and two dorsomedial spines. Form and structure of bursa copulatrix and sper- mathecal apparatus unknown (one speci- men damaged, the other too teneral for dis- section). Dispersal potential.— The wings are fully developed, and as with the following spe- cies, we believe that adults have strong pow- ers of flight. Natural history.—Stork (1982) presented all the available information on the habitat distribution of this species, and we use these data in the discussion below. A specimen collected in April, 1982, was teneral, which suggests that adult emergence occurs at that time. Nothing else is known about the life PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON history or behavior of members of this spe- cies. Locality records. —(Fig. 34). Known only from the type locality (in IRAN) and KU- WAIT, Al Jahrah (29°20’E 47°40'N) (BMNH; | female). Cicindis Bruch Cicindis Bruch 1908: 497. Type species. — Cicindis horni Bruch 1908: 499. Diagnosis. — Body moderately depressed. Head (Fig. 3B) without supraorbital setifer- ous punctures; area between posterior mar- gin of eye and postgenal groove markedly depressed; antennomeres | to 4 glabrous (except for apical whorl of fixed setae), 5 to 11 densely and evenly pubescent (Fig. 4B); dorsal surface of mandible (Figs. 6, 7) with obliquely transverse grooves and ridges; last (fourth) maxillary palpomere (Fig. 9) with- out sensory pit ventromedially in apical one- half; ligula of labium (Fig. 10B) with apical margin deeply emarginate, paraglossae short but equal in length to fused glossae; mentum (Fig. 11B) with anteromedial emargination shallow, dentate, tooth bifid, with two pairs of paramedial setiferous punctures at base, epilobes broadly rounded, narrowly toothed anterormedially and with six or seven pairs of marginal and several pairs of basal setif- erous punctures; submentum narrow ante- riorly, much narrower than mentum, with three or four pairs of anterolateral setiferous punctures, gular portion with sides diver- gent posteriorly. Pterothoracic elytron- locking mechanism (Fig. 18B) with long in- ternal and external metapleural flanges, external flange without posterior bulge (ely- tron with poorly defined receptive cavity), epipleuron entire. Femora with both ante- rior and posterior surfaces convex. Middle tibia (Fig. 20B) with posterodorsal fringe of long, fine accessory setae; hind tibia (Fig. 21B) markedly arcuate; tibial spurs of an- terior tibia shorter than tarsomere 1, pos- terior spur of middle tibia and anterior spur VOLUME 93, NUMBER 2 of hind tibia less than or equal to one-half length of respective first tarsomeres. Front tarsomere 1 (Fig. 22B) with posterodorsal fringe of accessory setae, middle tarsomeres 1 to 5 (Fig. 24B) with anteroventral and posterodorsal fringes of accessory setae, hind tarsomere | (Fig. 25B) with anteroventral and posterodorsal fringes of accessory setae, tarsomeres 2 to 5 with anteroventral fringe only; posterior claw (Fig. 23B) of all tarsi equal to or less than one-half the length of anterior claw. First visible (second) abdom- inal sternum (Fig. 17) with deep lateral con- cavity. Geographic distribution.—Known only from the great salt lake region of north- western Cordoba Province, Argentina. Cicindis horni Bruch Cicindis horni Bruch 1908: 499. Holotype female, in MACN, labelled: ““Rep Argen- tina Prov. Cordoba [illegible number] C. Bruch’’/ “Col. C. BRUCH” [white label with black border]/ ““Cicindis Horni Bruch C. BRUCH DETERM.” [white label with black border] ‘““Typus” [light green label]/ ‘“HOLOTYPUS” [red label with black border]. Type locality. ARGENTINA, Cordoba Province, Guanaco Muerto [30°29’S 65°03'W], herewith restricted. Diagnosis.—Clypeus narrower than dis- tance between antennal sockets, supraor- bital setiferous punctures absent, and pro- notal and elytral margins smooth. Description of form and structure of adults. —Size medium, standard body length of male 10.2 mm, female 10.5 mm. Form (Fig. 2) with elytra proportionately long in relation to forebody (ratio: standardized elytral length/forebody length = 1.95 [mean]). Color: Head, body, and appendages pale yellowish tan, except eyes piceous and api- ces and medial margins of mandibles in- fuscated; elytra with very pale marginal pat- tern (Fig. 15B). Luster: Entire body surface moderately shiny, slightly duller on elytra of female. 363 Microsculpture: Entire body with small, isodiametric meshes, well impressed on la- brum, legs, and venter of both sexes and elytra of female, effaced or nearly so from forebody of both sexes, less impressed on elytra of male especially on crowns of in- tervals. Head: Smooth dorsally; frons slightly convex, frontal furrows broadly impressed, parallel, and limited to area medial to an- terior half of eye; antenna (Fig. 4B) with scape short, length about 0.7 times width of labrum, cylindrical, and form straight; clyp- eus (Fig. 5B) with anterior margin slightly emarginate; penultimate labial palpomere (Fig. 10B) with four setae anteriorly. Prothorax: Pronotum (Fig. 12B) narrow, greatest width less than or equal to width of head across eyes, markedly cordate, nar- rowed basally, smooth medially, with obliquely transverse and shallow rugulae laterally; disc markedly convex; apical mar- gin markedly bisinuate; lateral margin smoothly arcuate, with basal sinuation moderately deep and abrupt; lateral expla- nation narrow, slightly broadened apically and basally; basal margin slightly bisinuate; apical angles long, narrow, markedly pro- jected anteriorly; basal angles subrectan- gular, not at all projected posteriorly; apical margination deeply impressed laterally, ab- sent from middle one-third; lateral bead narrow, poorly defined, effaced near apical and basal angles; basal margination absent; anterior transverse impression broadly and deeply defined; median longitudinal im- pression narrowly and finely defined; pos- terior transverse impression smooth to slightly rugulose, narrowly and deeply de- fined, extended laterally to basal foveae, straight; basal foveae deep anteriorly, pos- teriorly very shallow, slightly divergent, and confluent with lateral explanation; midlat- eral setiferous puncture absent, basolateral setiferous puncture inserted slightly ante- rior to basal angle. Prosternal intercoxal process (Fig. 14A) narrowly sublanceolate in ventral aspect. 364 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Habitus, dorsal aspect, Cicindis horni Bruch. VOLUME 93, NUMBER 2 Pterothorax: Elytra (Fig. 15B) with sil- houette subrectangulate, widest posterior to middle, lateral and apical margins smooth; elytron slightly depressed, with basal mar- gination complete, deeply impressed and slightly sinuate, lateral margination com- plete and very narrow throughout, humerus squared, lateral and basal margination joined at slightly obtuse angle, interneurs | to 7 finely striatiopunctulate, less distinctly so apically, 8 and 9 finely punctulate, in- tervals smooth, impunctate, | and 3 to 8 moderately convex to apex, 2 and 9 slightly convex or flat; metathoracic wing (Fig. 16B) full-sized and functional, with stalk of Ob- longum Cell long. Legs: All legs long and slender; front tar- someres | to 4 slightly expanded laterally and with pads of adhesive setae ventrally in male (front tarsi of female specimen miss- ing). Abdomen: Last visible (seventh) sternum slightly emarginate apicomedially in female but entire in male, with two pairs of long paramedial marginal setae in female, and apparently one pair in male (only male available for study has one such seta on one side). Male genitalia: Ring sclerite (Fig. 26) ovoid, slightly asymmetrical, slightly nar- rowed anteroventrally; median lobe (Fig. 27) short, with broad apical orifice. Parameres slightly asymmetrical, with left paramere slightly longer and narrower than the right, both bisetose apically. Female genitalia: Coxostylus (Fig. 31B) pointed apically, slightly serrate apicolater- ally, with three ventrolateral and three dor- somedial spines. Bursa copulatrix (Fig. 32) with anterodorsal lobe broad, apex deflect- ed right of midline anteriorly, short and wide posteroventral lobe present; spermatheca subequal in diameter to spermathecal duct, slightly twisted; spermathecal duct slender, inserted anteriorly on anteroventral lobe of bursa copulatrix. Sexual dimorphism: In addition to leg and 365 abdominal characters described above, fe- male slightly larger overall than male. Dispersal potential.—The wings are fully developed and one specimen was recorded as having flown into a light (R. A. Ronderos, pers. comm.). Although fringe setae on the legs suggest that these beetles are swimmers, we believe they have strong powers of flight as well. Natural history.—These beetles are re- corded only from salt lakes and we suspect that they are confined to such areas. Their swimming hairs, elevated eyes, and mouth- parts lead us to suggest that they are su- perspecialist, amphibious predators at the edges of shallow alkaline lakes, the carabid equivalent of a crocodile. The adult speci- men, taken at light in December, 1979, is unusually soft for a mature adult carabid, which suggests that it may have been slight- ly teneral when collected. With the possible exception of this indication of adult emer- gence period, nothing is known about the life history or behavior of members of this species. Locality record.—(Fig. 35). ARGENTI- NA, Cordoba Province (MACN; 1 female) Guanaco Muerto (UNLP; | male). DISCUSSION A review of the descriptions provided above for the tribe, genera, and species of cicindines reveals an array of features of form and structure that, even for the trained specialist on carabid beetles, represent a unique and surprising combination of traits. What justification is there for assigning the two known species to distinct genera and these genera to a distinct tribe? What evi- dence is available in support of the mono- phyly of the tribe Cicindini as here defined and how is this taxon related to other tribes and more inclusive groups of carabids? What, if anything, can we infer about the biogeographic history of the group from present knowledge of the geographical and habitat distributions of extant cicindine taxa? To what extent can we relate apparent 366 specializations in form and structure to hab- itat or behavior? What future research ef- forts are required to generate the kinds of additional data needed to better answer all of these and other questions? In the follow- ing discussion, we address each of these top- ics. PHYLOGENETIC RELATIONSHIPS AND CLASSIFICATION In his review of the present system of classification of carabid beetles and its his- torical development, Ball (1979) suggested that “higher-ranking taxa are in part clade- based, and in part grade-based.” He iden- tified five evolutionary trends in carabid evolution, with reference to which the basic framework of that classification has been constructed. Trends included were (1) de- velopment of a more complex antenna- cleaning organ on the front tibia; (2) mod- ifications of the pterothorax; (3) loss of a setiferous puncture from the scrobe of the mandible; (4) reduction of the apex and oth- er modifications to the parameres in males; and (5) reduction of elytral apices and en- hancement of the ability to take flight. To- gether, the presumed starting points (i.e. plesiotypic states) of each of these trends (characters or suits of characters) define the basic ground plan of the hypothetical an- cestral carabid. Attempts to establish phy- logenetic relationships among suprageneric carabid taxa using the characters involved in these trends (Sloane 1923, Jeannel 1941, Crowson 1955, Bell 1965, 1967, Kryzhan- ovskiy 1976) have all failed, at least par- tially, due to homoplasy (convergence) in these traits, as shown by the discordant dis- tributions of character states among the taxa considered. As demonstrated below, clado- grams generated with reference to different characters suggest very different patterns of relationships. The present classification is therefore a compromise arrangement, the analog of a consensus tree, the different nodes of which are a mixture of grades and clades, just as Ball (1979) suggested. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Kavanaugh and Négre (1982) tried to de- termine phylogenetic relationships of the tribe Notiokasiini with other member tribes of supertribe Nebriitae but noted that the “main difficulty with [phylogenetic] studies of nebriite genera is in establishing a suit- able outgroup for character analysis,” against which character transformations can be po- larized. Failure to agree on a suitable out- group for the Carabidae results in the same problem at a higher level. For example, Ka- vanaugh (1986) followed Ponomarenko (1977) in suggesting that trachypachines (as presumed living representatives of the Ju- rassic-Cretaceous eodromine radiation) are the sister group of the remaining Carabidae and, with them, form a monophyletic group. Acceptance of this view results in a clado- gram for the Carabidae that approximates, in broad outline but not in detail, a tree reflecting the present classification (Kava- naugh, study in progress). However, Bell (1966, 1982), Hammond (1979), and Roughley (1981) interpreted available data as supporting a sister group relationship be- tween trachypachines and at least some Hy- dradephaga (the dytiscoid families in par- ticular). Further, Bell (1966) suggested a sister group relationship between trachy- pachines + some hydradephagans and Me- triini + Ozaenini + Paussini. If this inter- pretation is correct, then Carabidae, including metriines, ozaenines, and paus- sines, but excluding either trachypachines + some hydradephagans or these hydra- dephagans alone, is a paraphyletic group. A suitable outgroup for this entire assemblage must be sought at or outside the basal ra- diation of suborder Adephaga. Acceptance of this view requires a radical reclassifica- tion of Adephaga in general and of Carab- idae in particular. Preliminary cladistic analyses with char- acters and/or character systems tradition- ally used in carabid classification above the genus level have failed to provide an un- ambiguous resolution of phylogenetic rela- tionships among even the better known VOLUME 93, NUMBER 2 higher taxa of carabids (Ball 1979, Kava- naugh 1986 and study in progress). Such a basic framework of caraboid relationships, which is prerequisite to understanding the relationships of cicindines and their proper place in a classification, must await more uniform, detailed, and broad-based study of form and structure across the Adephaga, particularly with internal organ systems of adults and internal and external structures of immatures. Molecular studies may also aid in this effort. For the present, we limit ourselves to placement of cicindines within the present hybrid cladistic/gradistic carabid classifi- cation, based on our study of their form and structure and our reinterpretation of tradi- tionally used characters and their transfor- mations. To do this, we examine the char- acters individually and then place the tribe, on the basis of synapotypic features shared with other tribes, in the existing classifica- tion. For these purposes, we accept trachy- pachines as the sister group of all other cara- bids (Kavanaugh 1978, 1986) and character transformations are polarized accordingly. Coxal cavities and thoracic structure. — Bell (1965, 1967) provided the most thor- ough analysis of the arrangement of thoracic sclerites that form the cavities surrounding the three pairs of coxae in Adephaga, al- though several other workers (Sloane 1923, Jeannel 1941, Ball 1960, 1979, Lindroth 1969, Hlavac 1972, 1975, Ponomarenko 1977) have also discussed these features and their use in classification. In our discussions here, Bell’s (1967) analysis and nomencla- ture are used, except as noted, and each tho- racic segment is considered separately. Prothorax: In cicindines, the front coxal cavities are enclosed behind by a medial extension of the propleuron (not proepi- meron; see Hlavac 1975) that makes con- tact, with a slight posterior overlap, with the prosternal intercoxal process (Fig. 14); right and left coxal cavities are completely separated medially by a sclerotized internal septum (Fig. 14B); and the dorsum of each 367 cavity is spanned by a narrow sclerotized bridge that divides the dorsal unsclerotized opening into two parts (Fig. 14A), the so- called biperforate (Sloane 1923) or bridged (Bell 1967) condition. The states of each of these three characters found in cicindines, namely closed, separate, and biperforate coxal cavities, traditionally have been con- sidered the apotypic state (Sloane 1923, Bell 1967) in relation to their respective alter- nate state (i.e. open, confluent, and un- bridged). However, Hlavac (1975) reported the bridged condition in several groups (e.g. in Leistus, Carabus, Hiletus and cicinde- lines) whose members were thought to have uniperforate coxae only (Bell 1967, Erwin and Stork 1985). The distribution of bridged front coxal cavities as reported by Hlavac (1975) suggests that this condition repre- sents the plesiotypic state of this character among Adephaga. Also, detailed examina- tion of the area of contact between pro- pleuron and prosternum that forms the pos- terior closure of the cavity suggests that the method of closure differs in different groups (Bell 1967, Hlavac 1975) and hence, that closure may have evolved several times in- dependently. Cicindines share the apotypic pair of fea- tures, closed and separate coxal cavities, with a majority of the tribes of Carabidae, but not with hydradephagans, trachypachines, or a few other carabid-tribes— namely those thought to represent the basal grade of cara- bid evolution. Among the groups he ex- amined, Bell (1967) found five of the eight possible combinations of states for the three two-state characters, and Hlavac (1975) re- ported on a sixth combination (e.g. open, separate, bridged) in Hiletus. Clearly, ho- moplasy is involved in the present distri- bution of these character states. Mesothorax: The lateral wall of the mid- dle coxal cavities in cicindines includes the medial margin of the mesepimeron (Fig. 17), the so-called disjunct condition (Sloane 1923, Jeannel 1941). Medially, mesosternal and metasternal processes meet but do not 368 completely separate the right and left coxal cavities, which therefore remain at least partly confluent internally (Fig. 17). Sloane (1923) grouped those tribes whose members had disjunct middle coxal cavities under the name Carabidae Disjunctae and called the remaining carabids, in which the mesepimeron is separated from the coxal cavity by a posterolateral extension of the mesosternum, the Carabidae Conjunctae. However, the disjunct and confluent middle coxae probably represent the plesiotypic states of each of these characters (Bell 1967, Kavanaugh 1986), so the monophyly of the Carabidae Disjunctae is unsupported. Bell (1967) suggested that conjunct middle coxal cavities evolved independently in Notio- philini and that Gehringiini may have evolved the disjunct condition secondarily. Aside from those possible exceptions, the distribution of states of this character among carabid tribes suggests that little or no ho- moplasy is involved, and the Carabidae Conjunctae may well be a monophyletic group. If so, then most of the tribes with which the Cicindini share closed and sep- arate front coxal cavities form a monophy- letic group that does not include them, and independent closure of front coxal cavities in the cicindines is again suggested. Metathorax: In cicindines, a distinct met- epimeron forms the lateral wall of the hind coxal cavities (Fig. 17). Bell (1967) called this condition disjunct, and recognized three other states of this character: conjunct, in which the metepimeron is apparently ab- sent; /obate, in which the posterior edge of the metepimeron is free and partially over- laps the first visible sternum; and incom- plete, in which the coxa extends to the mar- gin of the body and the metepimeron is apparently absent. The hind coxal cavities are confluent medially in cicindines, as in all other Adephaga except gehringiines and rhysodines. Bell (1967) suggested that the disjunct condition is the ancestral (plesiotypic) state and that the other states are apotypic, not PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON necessarily in any graded sequence. He pro- posed that the incomplete condition, found only in trachypachines, gehringiines, rhy- sodines, and hydradephagans among extant Adephaga, is a synapomorphy for the trachypachines and at least some hydra- dephagans (Gehringiini and Rhysodini were excluded on other grounds; Bell 1964, 1967). However, this arrangement of hind coxae is found also in fossil protocoleopterans, fossil and extant archostematans, and fossil eo- dromines, as well as all extant hydradepha- gans and trachypachines (Ponomarenko 1977). Kavanaugh (1986) suggested that this trait was plesiotypic for Adephaga, with other states evolved from it. Bell’s hypoth- esis for the transformation of this character rests on the assumption that the ancestral metathoracic arrangement of pleural scler- ites was similar to, if not serially homolo- gous with, the mesothoracic arrangement found in extant forms with disjunct middle coxal cavities (see Bell 1967: Fig. 1). If, how- ever, the incomplete condition is plesiotyp- ic, then the ancestral adephagan may have lacked a distinct metepimeron, and the sclerite found in extant cicindines and many other carabids (and called the metepimeron) may not be serially homologous with the mesepimeron. Metepimera of the type that define the disjunct condition of hind coxal cavities are found in varied form among cicindines, cic- indelines, elaphrines, loricerines, and scar- itines, but they are best developed in the first and last of these groups. If this character state 1s apotypic, as we suggest, then it may be either synapotypic for these tribes or in- dependently evolved in some or all of them. Because few other synapotypic features ap- pear to support close relationships between cicindines and any of these particular tribes, we suspect that homoplasy is involved in the character state distribution of this char- acter. Front tibial spurs and antenna cleaner. — Jeannel (1941) divided Carabidae into two groups, based on the location of the front VOLUME 93, NUMBER 2 tibial spurs, form of the tibial apex, and form and location of the antenna cleaner. Those beetles with both spurs positioned apically and the tibial apex truncate or near- ly so, but with the antenna cleaner (setal band; Hlavac 1971) extended and devel- oped proximal to both spurs were included in his Isochaeta. Among these were trachy- pachines, gehringiines, metriines, ozae- nines, and paussines. Ball (1979) added no- totylines and cicindines (but see below) to this group. The remaining carabids form a second group, which Jeannel did not name, but which has come to be called the Ani- sochaeta (see Ball 1979). As Hlavac (1971) noted, Anisochaeta includes a few groups like the cicindelines and opisthiines in which both tibial spurs are apical, the tibial apex is only slightly oblique, and the antenna cleaner is of the sulcate type (simply a trans- verse setal band across the concave tibial apex between the spurs). However, it also includes groups in which the posterior tibial spur is displaced proximally with the pos- terior end of the setal band to form an an- tenna cleaner more or less removed from the tibial apex. Jeannel (1941) was the first to recognize that, among carabids, a proximal shift of the antenna cleaner occurred both with and without proximal displacement of the pos- terior tibial spur. However, it was Hlavac (1971) who suggested that the arrangement of tibial spurs, tibial apex, and setal band found in cicindelines, opisthiines (i.e. the sulcate type of antenna cleaner) could serve as a Starting point from which development of a proximally displaced antennal cleaner could evolve along two different lines: one involving proximal displacement of the posterior tibial spur (the anisochaetous mode) and one independent of spur dis- placement (the isochaetous mode). The an- isochaetous antenna cleaner has probably evolved several times independently among Carabidae, perhaps the most striking single example of which is its occurrence in Pam- borini, a group that is certainly closely re- 369 lated to cychrines and carabines (Moore 1966), all of which have sulcate antenna cleaners. Cicindines have the antenna clean- er sulcate (Fig. 19B), the plesiotypic state for this character. Mandibular setae. —Cicindines lack a se- tiferous puncture in the scrobe of the man- dible. Presence of this seta is considered ple- siotypic, its loss apotypic. However, loss of the seta has probably evolved in several lin- eages independently. Carabid with special- ized mandibles (Loricera, hiletines, and Promecognathus, for example) lack a scro- bal seta. Cicindine mandibles have a well- developed scrobe but modified terebral blade, which may account for absence of the seta. Although Jeannel (1941) and others have relied heavily on this character in their classifications (Ball 1979), it contributes lit- tle to an understanding of cicindine rela- tionships. Labral setae.—Bell (1964) identified the number of setiferous punctures on the an- terior margin of the labrum dorsally as an important character for corroborating Jean- nel’s Isochaeta. He suggested that six was the plesiotypic number of setae in carabids and that a higher number characterized the Isochaeta. Trachypachines, metriines, paussines, and many ozaenines have ap- proximately 12 such setae. Mainly on this basis, Bell was able to exclude gehringiines, which have only six labral setae, from the Isochaeta (with which they share incom- plete hind coxal cavities). Labrum with six setae, the number found in cicindines, is certainly the most wide- spread condition among carabids, and it also may be the plesiotypic state for this char- acter at the family level, as Bell suggested. However, some cicindelines and carabines also have a higher number of labral setae, up to 10 or 12. Phylogenetic relationships among genera and species groups within these tribes are still unclear, so the plesio- typic number of labral setae for each of them cannot be determined at present. Among ozaenines, Ball and McCleve (1990) found 370 a range of from 0 to 17 setae and suggested that the highest numbers were plesiotypic, the lower numbers apotypic. These data and the fact that trachypachines, our choice as the sister group for all other carabids, have a high number of seta, suggest that this may be the plesiotypic state for Carabidae. Six or fewer (as in loricerines, for example) la- bral setae may be apotypic within carabids, but certainly not synapotypic for all taxa with this trait (e.g. the independent reduc- tion in number of setae within ozaenines already cited). Other assorted setae.—Presence or ab- sence of several other fixed setiferous punc- tures has been used widely in carabid clas- sification (see Ball 1979 for summary). The supraorbital setae, pronotal (midlateral and basolateral) setae, and the discal and um- bilicate series of elytral setae have been most often considered. In cicindines, the poste- rior supraorbital and midlateral pronotal setae are absent. The anterior supraorbital setae are also absent from Cicindis horni specimens examined. A few very short and fine discal setae occur on elytral interval 3 in C. horni, but the elytral disc is asetose in Archaeocindis johnbeckeri. Both species have a well-developed umbilicate series of 12 to 14 very short, fine setae on interval 9. With each of the setal characters just mentioned, absence (or a reduction in num- ber of setae in a series) is probably apotypic in relation to the number and arrangement in the hypothetical ancestral carabid. The usefulness of these characters as indicators of phylogenetic relationship, however, is varied and probably less important than their use as aides for identification, at least for the present. The pattern of presence and absence of these setae in cicindines suggests no particular phylogenetic affinity with one or more other tribes of carabids. Metathoracic wing.—Ward (1979) ex- amined metathoracic wing venation pat- terns in Adephaga and found several char- acters that he suggested are useful indicators of phylogenetic relationship. Among these PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON are (1) shape of the Oblongum Cell (OC) and relationship of 4m-cu and 5m-cu cross- veins to the M4 and Cubitus veins; (2) point of insertion of M4 on the distal side of the OC; (3) relative sizes of the Third Radial (3RC) and Anterior Sector (SAC) cells. He proposed that the plesiotypic states of these characters are as follows: (1) OC transverse- ly rectangular, with 4m-cu and 5m-cu cross- veins clearly separated posteriorly; (2) M4 inserted in the anterior one-third of the dis- tal wall of the OC; and (3) 3RC subequal in size to SAC. In cicindines (Fig. 16), the OC is nar- rowed posteriorly, with the 4m-cu and 5m- cu crossveins fused anterior to the Cubitus to form a stalk for the OC; M4 1s inserted at or slightly anterior to the middle of the distal wall of the OC; and 3RC appears to be larger than SAC. This combination of traits is not represented among examples that Ward provided; and whether or not it is shared with any of the many other carabid groups that he did not discussed we cannot judge. The degree of usefulness of venational characters in phylogenetic reconstruction that Ward suggested remains untested in our view. For example, among the nebriines alone, shape of the OC varies from nearly rectangular to distinctly triangular, with the 4m-cu and 5m-cu crossveins separate at the Cubitus in some species, or fused for greater or lesser distances anterior to the Cubitus, resulting in shorter or longer stalks for the OC in other species (Kavanaugh 1978). In most, but not all, nebriines, M4 is inserted on the distal wall of the OC distinctly pos- terior to its midpoint (Kavanaugh 1979, Figs. 33-35), whereas insertion closer to the midpoint is at least suggested by Ward’s cladogram (Ward 1979, Fig. 22). Clearly, additional detailed and comparative study of venation patterns is needed before the value of these characters can be determined. Parameres of male genitalia.—Jeannel (1941) made a detailed study of the para- meres of male carabids and relied heavily VOLUME 93, NUMBER 2 on differences found among them in con- structing his classification. Important fea- tures included the degree of symmetry of right and left parameres in shape and size and the presence and distribution of setae on them. Symmetrical and setose parameres have been considered plesiotypic by most workers following Jeannel. In the Cicindis horni male examined, the parameres are moderately long and slender, only slightly asymmetrical (Fig. 27), with the left slightly longer and narrower than the right, and both apparently with two se- tae apically. This arrangement is very sim- ilar to that seen among bembidiines, tre- chines, pogonines, and patrobines among Jeannel’s (1941) Stylifera, a group with which cicindines share few other apotypic features. Coxostyli of female genitalia. — Bell (1982) and others have suggested that the apparent absence of a gonostylus (or stylomere two) from the ovipositors of female hydradeph- agans, trachypachines, metriines, ozae- nines, and paussines (i.e. the Isochaeta of Bell 1967) may be a synapotypic feature for this group. However, Kavanaugh (1986) noted that opisthiines, nebriines, notioka- slines, and, in fact, many basal-grade cara- bid groups also have females with a gono- stylus either absent or fused with the gonocoxite (or stylomere one) to form an unjointed ovipositor. He suggested that this condition was apotypic for the suborder Adephaga, not just for the Isochaeta (sensu Bell), and that structures called gonostyli (second stylomeres) in female cicindelines, carabines, cychrines, and most intermedi- ate- and advanced-grade carabids may not, in fact, be homologous with the gonostyli of female Archostemata and Polyphaga. In cicindines, the ovipositor blades, which we refer to as the coxostyli, are unjointed, the condition that we view as plesiotypic within Carabidae. Placement of Cicindini in relation to past and present classifications. — We provided a brief history of the placement of Cicindini 371 in carabid classification in our introduction. But where would cicindines have been placed in some of the more important clas- sification schemes in which they were not considered, and where should they be placed now? In the classification proposed by Sloane (1923), Cicindini would be grouped with the Carabidae Disjunctae-Clausae, but they cannot be identified using Sloane’s key. The only tribes listed whose members have dis- junct middle coxal cavities, closed front coxal cavities, and terminal front tibial spurs are the ozaenines and metriines; but ozae- nines have the lateral elytral margin “‘with a process [= the flange of Coanda; Ball and McCleve 1990] on each side,” and me- triines have a mandibular scrobal seta. In addition, the antennal cleaner is of the 1so- chaetous type in members of both of these groups. Cicindines cannot be placed within any of the supratribal groups proposed by Jean- nel (1941). The sulcate antenna cleaner ex- cludes them from Isochaeta, the large, dis- tinct metepimeron from the Simplicia, shape and vestiture of the parameres of males from the Scrobifera, and the disjunct middle cox- al cavities from the Stylifera, Conchifera, and Balteifera. They would have to be placed in a separate group within the Limbata, near the Scrobifera. In his informal classification scheme, Bell (1967) recognized a group that he called An- isochaeta-Isopleuri, members of which have front coxal cavities closed, separate, and un- bridged, middle coxal cavities disjunct and confluent, metacoxal cavities disjunct and confluent, and antenna cleaner sulcate or proximally displaced in the anisochaetous mode (i.e. posterior tibial spur also dis- placed proximally). Among these traits, the only one not found in cicindines is un- bridged front coxal cavities. As noted above, distribution of the bridged condition, found in cicindines, is still too poorly known among adephagans to justify exclusion or inclusion of cicindines on this basis alone. 372 Cicindelines, loricerines, elaphrines, and scaritines were included in the Isopleuri as defined by Bell. Ball (1979) suggested that Bell’s groupings of tribes may represent grades attained independently by different clades, and we suspect that, in this view, he is at least partially correct. Our reinterpre- tation of some of the character polarities reinforces this opinion. The classification of carabids that most closely reflects both our present understand- ing and our ignorance of phylogenetic re- lationships (Ball 1979) among extant supra- generic carabid taxa is that proposed by Kryzhanovskiy (1976). In that classifica- tion, the four tribes in Bell’s Isopleuri are distributed among two subfamilies (Cicin- delinae [= Cicindelini sensu Bell, and as we have used the name in this report] and Ca- rabinae) and three supertribes in the latter subfamily (i.e. the Elaphritae, Loriceritae, and Scarititae). We suggest that the best placement of Cicindini in Kryzhanovskty’s scheme is in a supertribe of its own, the Cicinditae, between the Nebriitae and Ela- phritae. Monophyly of cicindine taxa and justifi- cation for their ranking. — Our placement of the Cicindini in the present carabid classi- fication is both tentative and unsubstan- tiated by an unambiguous set of nested syn- apomorphies with other suprageneric taxa. Nonetheless, the evidence for the mono- phyly of the group and justification for its recognition as a distinct tribe are provided by a suite of autapotypic features that in- cludes the following: (1) orientation of mouthparts slightly hypognathous; (2) ex- cept for gular region of head, entire ventral surface of body and all surfaces of coxae, trochanters, and femora covered with fine pubescence; (3) gena with a sharp flange ventral to eye; (4) compound eye very large; (5) mandible with terebral blade long and markedly down-curved; (6) mandibular scrobe without a setiferous puncture; (7) front coxal cavities closed and separate; (8) tarsal claws asymmetric in length, with an- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON terior claw longer than posterior; and (9) eighth sternum and ninth/tenth tergum un- divided. A tenth feature, hind coxal cavities disjunct (metepimeron present and large), may also be synapotypic for the group. Two of these characters, (6) and (7), have been used repeatedly as distinguishing features at the tribal level. The remaining seven or eight characters represent significant evolution- ary novelties among carabid beetles that, in concert, suggest a group with a long and unique evolutionary history. Justification for recognition of a distinct genus for each of the two cicindine species is based on autapotypic traits for each of these taxa that suggest a degree of differ- entiation between them at least as great as between most pairs of closely related cara- bid genera in other tribes. The following autapotypic features are recognized for Ar- chaeocindis: (1) body deep, subcylindrical; (2) apical maxillary palpomere with large sensory pit ventromedially in apical one- half; (3) pterothoracic elytron-locking mechanism and elytral epipleuron as in Fig. 18A; and (4) femora longitudinally concave. Autapotypic features for Cicindis include (1) head without supraorbital setiferous punc- tures; (2) dorsal surface of mandible with obliquely transverse grooves and ridges; (3) epilobes of mentum broadly rounded, nar- rowly toothed anteromedially, and with six or seven pairs of marginal and several pairs of basal setiferous punctures; (4) submen- tum narrow anteriorly, much narrower then mentum, with three or four pairs of antero- lateral setiferous punctures; (5) middle tibia with a posterodorsal fringe of long, fine ac- cessory setae; (6) hind tibia markedly ar- cuate; (7) one or more tarsomeres on each leg with fringes of accessory setae (see Figs. 22B-25B for distribution of fringes); and (8) first visible (second) abdominal sternum with deep lateral concavity. HISTORICAL BIOGEOGRAPHY Jeannel (1942) recognized several carabid groups whose present distributions include VOLUME 93, NUMBER 2 at least parts of the Guyana and Brazilian Shields of South America, tropical and southern Africa, Madagascar, and India. He noted that, during the Mesozoic, these areas were part of the western portion of the su- percontinent, Gondwanaland, prior to the development of the South Atlantic oceanic basin by the end of that era. He called groups with such extant distributions /ignées ina- brésiennes (Jeannel 1941, 1942) and sug- gested that these disjunctions reflect the fragmentation of what were more wide- spread ranges in western Gondwanaland through development of the southern At- lantic Basin [vicariant event] in late Me- sozoic time. Reichardt (1977, 1979) and Noonan (1985) discussed the distribution of cicindines and listed several other cara- bid tribes with South American/African disjunctions (e.g. hiletines, siagonines, and apotomines). They agreed with Jeannel in dating the initiation of these disjunctions [vicariance] to the opening of the South At- lantic, at least 65 million years ago. If this timing of vicariance is correct, as we also suggest, then Archaeocindis and Cicindis have had at least that long to evolve inde- pendently. Development of hypotheses concerning the origins of the Cicindini and their pre- or early-Gondwanian history must await discovery (or at least recognition) of their sister group. Available data suggest that the two known cicindine species differ in their habitat dis- tributions: A. johnbeckeri adults apparently occur on saline, intertidal mudflats of bays in the Persian Gulf, and C. horniadults have been found only in the vicinity (microhab- itat still unknown) of salt lakes in the in- terior of Argentina. Were ancestral cicin- dines coastal, sea beach inhabitants that subsequently invaded interior saline lake beds, or were they originally interior forms that radiated into coastal areas? Analogous to this apparent difference be- tween cicindines are the different habitat distributions of three pogonine species in North America. Diplochaetus lecontei Horn 373 ranges from the Gulf Coast of Texas east to Florida and north to New Jersey. Its mem- bers are confined to sandy sea beach areas, where they occur at the margins of standing freshwater pools in dune areas (TLE, per- sonal observations). Diplochaetus desertus Van Dyke is known only from the shores of the Salton Sea in southern California. This saline water body was connected with the Gulf of California until the late Tertiary (early Pliocene) (Hunt 1974). Throughout Quaternary time, its water levels have fluc- tuated greatly, due to subsidence of its ba- sin, sedimentation, continued development of the Colorado River delta across its south- ern end, and changes in the channel of that river itself (Shelton 1966). Pogonistes pla- natus Horn is found throughout the interior of the western United States, from Okla- homa and Kansas northwest through Utah to eastern Oregon, where it occurs at the margins of saline lakes. These three habitat types form a series, from coastal sea beach to interior saline lake shore, with shores of the relatively recently isolated Salton Sea perhaps representing an intermediate stage between these extremes. Has D. desertus evolved from an ancestral, coastal form that became isolated from the sea coast with the separation of the Salton Sea from the Gulf of California? Could similar (yet unknown) events account for the evolution of other forms, living at the margins of saline lake beds, that have relatives in coastal areas? Although the three pogonine species are probably closely related (Van Dyke 1953), phylogenetic relationships among them have not been analyzed, nor has a suitable sister group for them been proposed. Without ref- erence to the habitat distributions of re- spective sister groups, the polarity of ap- parent transformations in habitat cannot be determined for either the pogonines or the cicindines considered. It is also possible that present habitat distributions of one or both groups are relictual, and that extinct or yet unknown sister groups occupied (or pres- ently occupy) completely different (e.g. non- 374 saline) habitats. The recognition of the sister group of cicindines 1s the necessary next step to understanding this and other aspects of the evolutionary history of this group. STRUCTURAL EVOLUTION, HABITAT, AND BEHAVIOR Several features of adult form or structure in one or both cicindine species are con- spicuous or otherwise of special interest and warrant at least brief comment here con- cerning their possible function or signifi- cance. With a cicindine specimen in hand, one is at first struck by its great similarity in overall form to that of some tiger beetles (Cicindelini). With many species in several genera of cicindelines, it shares a similar general body form, large compound eyes, long legs, and an elytral pattern of marginal pale markings. Additional similarities are apparent under microscopic examination. Cicindines and cicindelines both have closed front coxal cavities and at least most species of the latter group also have an apparent metepimeron. Many tiger beetles (e.g. Platychila pallida Fabricius and Megaceph- ala limata Perty) have genal flanges, serrate elytral margins (especially apically) and long tibial spurs, just as in cicindines. In all tiger beetles we examined, we found a metapleu- ral elytron-locking mechanism very similar to that occurring in cicindines. Based on many other important features in which cic- indines and cicindelines differ (see previous discussion), it is clear that most, if not all, of these similarities represent independent (convergent) developments in the two groups, probably in response to similar be- havioral and/or habitat distribution pat- terns. In fact most of these features are shared with other carabid taxa as well. Enlarged eyes, similar in form to those in cicindines, are found also in Notiophilus, Elaphrus, and Graphipterus adults and in at least some genera of many other tribes (e.g. Bembidiini and Lebiini) in addition to cic- indelines. Members of most, if not all, of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON these other groups are mainly day-active, visually-oriented predators, although indi- viduals are sometimes attracted to lights at night. Genal flanges seen in at least some cic- indelines (e.g. Megacephala limata) appear to be posterior extensions of the carinate rim of the maxillary fossa across the gena. Those in cicindines (Fig. 3) are separate from and dorsal to a line drawn posterior to the rim of the fossa. The cicindine flange appears to serve as a forward extension of the thin, fin- like plane established by the anteriorly ex- tended apical angle of the pronotum under the eye (see additional comments below). Further expansion of this flange on the gena, beyond the condition seen in extant cicin- dines, might result in a functional arrange- ment similar to that seen in most gyrinids, where a broad, sharp flange on the sclerite that divides each compound eye into dorsal and ventral fields continues the trimline of the pronotum anteriorly on the head. This is no doubt a streamlining (perhaps also hy- drofoil) adaptation in gyrinids for swimming rapidly at the surface. Its function in cicin- dines, where it is only partially developed, may be similar if less efficient. Closure of the front coxal cavities provides better protection ventrally for the membra- nous intersegmental connection between pro- and mesothoraces and permits increased ventral motion of the prothorax (Hlavac 1975). Because tiger beetles often assume a stance in which the venter is well above the substrate (see below) and exposed, coxal clo- sure may be an important protective adap- tation in this group, and perhaps in cicin- dines as well. The functional significance of the metepimeron is unknown. Very little comparative study has been made of elytron-locking mechanisms in ca- rabid beetles to date. The metapleural mechanisms in cicindines and cicindelines are at least similar in general form. Tiger beetles depend on their ability to take flight instantly, mainly for predator avoidance in exposed areas. Their elytron-locking mech- VOLUME 93, NUMBER 2 anism must facilitate rapid release; and the arrangement of metapleural and elytral epi- pleural parts that form this observed mech- anism would seem to provide such an op- portunity. It appears that release would involve simply lifting the elytra slightly, or depressing the abdomen and posterior por- tion of the metathorax slightly, or both. The presence of such a mechanism in cicindines suggests that they too can take flight quickly. Serrate elytral margins occur among many species and genera of cicindelines and also in several bembidiine taxa (e.g. especially among subtribes Anillina and Tachyina), some carabines (e.g. a few Calosoma spp.), and in single species in several other tribes. Carabids with serrate elytra occupy a wide range of habitats, from sandy sea beaches and the open shores of lakes and streams to the margins of forested swamps in tropical regions. No pattern of co-occurrence with a particular habitat or behavior pattern is ap- parent to us at present, but this feature must have some important function to have de- veloped independently in several different lineages. Long, slender legs occur in many carabid groups, but particularly in cicindelines, ne- briines, cychrines, and platynines. Members of each of these groups are fast runners, but among these, cicindelines are probably the fastest. Tiger beetles, the only one of these groups with diurnally active members, also have legs most similar in form to those in cicindines. Long legs not only facilitate lon- ger strides, hence speed, but also may permit the beetle to lift itself (behavior known as stilting) well above the substrate, an impor- tant advantage for life in open, exposed areas where daytime temperatures at the substrate surface may be extremely high (Pearson 1988). Exceptionally long tibial spurs (especially on the hind leg) are found in several different carabid groups, including many cicindelines, all masoreines, and in the genus Nemotarsus among lebiines. These cicindelines and some masoreines (e.g. Tetragonoderus spp.) run on 375 open, sandy substrates, whereas Nemotarsus adults and other masoreines (e.g. Sarothro- crepis spp.) are foliage or tree-trunk runners. The habitat distributions of cicindines spe- cies suggest that, like certain tiger beetles (e.g. Megacephala and Platychila spp.), they run on loose, particulate substrate. Long tibial spurs may assist in gripping such substrates and thereby facilitate running. The only feature shared uniquely by cic- indines and cicindelines (especially Cicin- dela and Megacephala spp.) is the elytral pattern of marginal pale spots. In fact, the patterns are different in detail in the two groups; but, overall, they are more similar to each other than either is to any other carabid with which we are familiar. The pat- tern is probably cryptic, an example of dis- ruptive coloration, in the respective habi- tats of these beetles. Cicindines share other features with a di- verse array of other adephagans, again, clearly through independent development of these traits. A pale body and appendages is Shared with many carabids, especially with species that live in sea beach, desert playa, or other exposed habitats. Such species in- clude Nebria diversa LeConte and Euryne- bria complanata (Linnaeus) (Nebriini), Platychila pallida (Cicindelini), Pogonistes planatus (Pogonini), and Tetragonoderus pallidus Horn (Masoreini). Among cara- bids, only cicindines are known to have asymmetrical tarsal claws; however, several group of dytiscids (e.g. Colymbetini, Hy- driphini, and a few genera in other tribes) have adults with asymmetry in claw length. Setal fringes on tibiae and tarsi, which aid in swimming, are widespread among hy- dradephagans, but among carabids, they are known to us only in C. horni adults. An- terior projection of the apical pronotal an- gles, lateral or ventral to the compound eyes, as thin, fin-like planes that are closely fit against the head, occurs in cicindines, some cicindelines (e.g. Platychila pallida), omo- phronines, amphizoids, many dytiscoids, and gyrinids. 376 One feature of adults of both C. horni and A. johnbeckeri that occurs elsewhere among adephaga chiefly in a few genera or species of the higher-grade carabid tribes (e.g. har- palines or chlaeniines), is the presence of dense setae on the venter. These setae may be hydrofuge in function and serve to trap air in a functional gill (Hinton 1976) for respiration during submersion underwater. Alternately, they may contribute to ther- moregulation by providing insulation from heat rising from the substrate. In fact, they may serve both functions under appropriate conditions. Although almost nothing has been ob- served directly about the microhabitat re- quirements or behavior of cicindines, con- sideration of their unusual suite of structural features suggests to us several hypotheses about their biology that can be tested by subsequent observations of these beetles in the the field. We suggest that they behave, in many ways, like many diurnal tiger bee- tles, as fast-running, visually oriented pred- ators that inhabit hot, open, saline habitats and take to flight rapidly, at least when dis- turbed. Both species have features that sug- gest an ability to withstand submersion (hy- drofuge hairs) and to swim (tibial and tarsal setal fringes in C. horni, concave femora in A. johnbeckeri, asymmetrical tarsal claws in both), perhaps chiefly at the water surface (genal flanges and anteriorly projected api- cal pronotal angles below the compound eyes). In general, structural features inter- preted as adaptations for swimming (i.e. se- tal fringes, asymmetry of tarsal claws) are more highly developed (relatively apotypic) in C. horni adults, those for running on hot, loose substrates (longer legs, exceptionally long tibial spurs) are better developed in A. Johnbeckeri adults. PROSPECTUS FOR FUTURE STUDY Further advance in our understanding of cicindine carabids must await additional specimens, both dead and alive, and field observations of behavior and habitat dis- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tributions. Additional museum specimens will permit greater freedom for dissection, thereby allowing a review of structures not yet properly examined, especially the male and female genitalia of both species. Live adults are needed for rearing cicindine lar- vae, which remain unknown. Larval fea- tures may provide additional clues to rela- tionships of this tribe with others. Field observations of habits and habitats are needed to confirm our suggestion, for ex- ample, that C. horni adults are excellent sur- face swimmers, using their specialized legs for this purpose. Comprehensive morpho- logical studies of both adults and larvae are urgently needed throughout the Adephaga to broaden and refine the base of compar- ative data available about carabid form and structure. Characters used traditionally in carabid systematics have helped to establish a Classification that functions moderately well but has failed to provide us with a clear understanding of relationships. New char- acters must be identified and surveyed and new techniques employed to resolve present conflicting observations. ACKNOWLEDGMENTS We thank curators L. Dieckmann (DEIE), A. O. Bachmann (MACN), R. A. Ronderos (UNLP), and N. E. Stork (BMNH) for the loan of specimens in their charge and their patience during this protracted study. We also thank George L. Venable for the com- puter-drawn habitus illustrations and anon- ymous reviewers and Michael G. Pogue for their helpful suggestions for the improve- ments for early drafts of the manuscript. George E. Ball added significantly to this study by generously sharing with us his in- sights and experience concerning carabid classification and diversity and by provid- ing unending encouragement to us over the decades of our respective professional as- sociations with him. Finally, to the memory of our departed friend and colleague, Donald R. Whitehead, we dedicate this paper. VOLUME 93, NUMBER 2 377 Figs. 3,4. Fig. 3. Head, left lateral aspect; A, Archaeocindis johnbeckeri (Banninger); B, Cicindis horni Bruch; aa = antennal articulation; ce = compound eye; cl = clypeus; gf = genal flange; lb = labrum; md = mandible; mxf = maxillary fossa; pgg = postgenal groove; pn = pronotum; pp = propleuron; ps = prosternum; ss = supraorbital seta. Fig. 4. Right antenna, dorsal aspect; A, Archaeocindis johnbeckeri (Banninger); B, Cicindis horni Bruch. 378 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON B tb C Figs. 5-7. Fig. 5. Labrum and clypeus, dorsal aspect; A, Archaeocindis Johnbeckeri (Banninger); B, Cicindis horni Bruch; cl = clypeus; lb = labrum. Figs. 6, 7. Mandibles, Cicindis horni Bruch: ; A, dorsal aspect; B, ventral: C, lateral aspects. Fig. 6. Left mandible. Fig. 7. Right mandible. At = accessory tooth (homology unknown); art = anterior retinacular tooth; prt = posterior retinacular tooth; sc = scrobe: Svg = setiferous ventral groove; tb = terebral blade; tm = terebral margin; tt = terebral tooth. 5 VOLUME 93, NUMBER 2 379 Figs. 8-11. Fig. 8. Left maxilla, ventral aspect, Archaeocindis johnbeckeri (Banninger); sp = sensory pit. Fig. 9. Left maxillary palpus, ventral aspect, Cicindis horni Bruch. Fig. 10. Labium, ventral aspect; A, Archaeocindis Johnbeckeri (Banninger); B, Cicindis horni Bruch. Fig. 11. Mentum and submentum, ventral aspect; A, Archaeo- cindis johnbeckeri (Banninger); B, Cicindis horni Bruch; mnt = mentum; sbm = submentum. 380 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 12 13 BS B g Figs. 12-14. Prothorax. Fig. 12. Pronotum, dorsal aspect; A. Archaeocindis johnbeckeri (Banninger); B, Cicindis horni Bruch. Fig. 13. Archaeocindis johnbeckeri (Banninger), ventral aspect. Fig. 14. Cicindis horni Bruch; A, ventral aspect; B, left ventrolateral oblique aspect. Cx = front coxa; fe = front femur; pcb = dorsal bridge of front coxal cavity; pec = front coxal cavity; pip = prosternal intercoxal process; pn = pronotum; pp = propleuron; ps = prosternum; tr = front trochanter; stippled areas = non-sclerotized areas. VOLUME 93, NUMBER 2 382 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SAC 3R Figs. 16-18. Fig. 16. Left metathoracic wing, dorsal aspect; A, Archaeocindis johnbeckeri (Banninger), B, Cicindis horni Bruch; Cu = Cubitus; M4 = fourth branch of the Media; O = Oblongum Cell; SAC = Anterior Sector Cell: 3RC = Third Radial Cell; 4m-cu = fourth medio-cubital crossvein; Sm-cu = fifth medio-cubital crossvein. Fig. 17. Pterothorax, left ventrolateral oblique aspect, Cicindis horni Bruch; eep = elytral epipleuron; lc = lateral concavity of first visible sternum; mscc = middle coxal cavity, msem = mesepimeron; mses = mesepisternum; mss = mesosternum; mtc = metacoxa; mtem = metepimeron; mtes = metepisternum; mts = metasternum: $1 = first visible sternum; stippled areas = nonsclerotized. Fig. 18. Metathoracic elytron-locking mechanism, left ventrolateral oblique aspect; elytron lifted slightly dorsad of resting position; A, Archaeocindis johnbeckeri (Banninger); B, Cicindis horni Bruch; eep = elytral epipleuron; msem = mesepimeron; mtem = metepimeron; mtes = metepisternum; S1 = first visible sternum. VOLUME 93, NUMBER 2 383 Figs. 19-21. Left tibiae. Fig. 19. Front tibia, Cicindis horni Bruch; A, anterodorsal oblique aspect; B, ventral aspect. Fig. 20. Middle tibia, dorsal aspect; A, Archaeocindis johnbeckeri (Banninger); B, C: icindis horni Bruch. Fig. 21. Hind tibia, dorsal aspect; A, Archaeocindis johnbeckeri (Banninger);, B, Cicindis horni Bruch. 384 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON A B —_—$<_—— Se Figs. 22-25. Left tarsi; A, Archaeocindis johnbeckeri (Banninger), female; B, Cicindis horni Bruch, male. Fig. 22. Front tarsus, dorsal aspect. Fig. 23. Front tarsal claws, apical aspect. Fig. 24. Middle tarsus, dorsal aspect. Fig. 25. Hind tarsus, dorsal aspect. VOLUME 93, NUMBER 2 385 6 27 | Figs. 26, 27. Male genitalia, Cicindis horni Bruch. Fig. 26. Ring sclerite, dorsal aspect. Fig. 27. Median lobe and parameres; A, ventral aspect; B, dorsal aspect; C, left lateral aspect. ST) 386 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 28-32. Female genitalia; stippled areas = unsclerotized areas. Fig. 28. Eighth sternum, ventral aspect; A, Archaeocindis johnbeckeri (Banninger); B, Cicindis horni Bruch. Fig. 29. Ninth/tenth tergum, dorsal aspect, Cicindis horni Bruch. Fig. 30. Left gonangulum, ventrolateral aspect, C: icindis horni Bruch; scale line = 0.1 mm. Fig. 31. Left coxostylus, ventral aspect; A, Archaeocindis johnbeckeri (Banninger); B, Cicindis horni Bruch; scale line = 0.1 mm. Fig. 32. Bursa copulatrix and spermathecal apparatus, C icindis horni Bruch; A, dorsal aspect; B, left lateral aspect; C, schematic, left lateral aspect; be = bursa copulatrix; cs = coxostylus; gp = gonopore; sd = spermathecal duct; sp = spermatheca; scale line = 0.1 mm. VOLUME 93, NUMBER 2 387 33 vi ae toa f Y 3 f) 18: a es \ \ bor 5 CASPIAN =~} \ { a \ x L Eaeeh dD P van ; Pas S